SCANDINAVIAN FISHES

SECOND EDITION

JV» 0036

Carded

Division of Fishes,
if. S. National Museum

SMITT. SCANDINAVIAN FISHES

A HISTORY OF

BY

B. FRIES, C, U. EKSTROM, AND C. SUNDEYALL

WITH COLOURED PLATES

BY

W. YON WEIGHT

AND TEXT ILLUSTRATIONS

SECOND EDITION

REVISED AND COMPLETED BY

Professor F. A. SMITT

A * I

MEMBER OF THE ROYAL SWEDISH ACADEMY OF SCIENCE

T E X T

PART II

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CONTENTS.

Ordo: PISCES TELEOSTEI (continued).
Subordo: Teleostei Physoclysti (continued).
Phalanx: Physoclysti Eleutherognathi (continued).
Cohors: Eleutherognathi Malaeopterygii (continued).

Series: Onomorphi (continued).

Familia: Ammodytidce pag. 5(17.

» Macruridce » 580.

» Ophidiidce » 594.

» Lycodidce - » 602.

Phalanx: Physoclysti Plectognathi » 618.

Series: Gyrnnodontes . » 622.

Familia: Orthagoriscidce ... » 622.

Series: Sclerodermi .. » 631.

Familia: Balistidce » 631.

Phalanx: Physoclysti Hemibranchii. » 635.

Familia: G aster ostei doe » 637.

Phalanx: Physoclysti Lophobranchii.. _ ... » 661.

Familia: Syngnathidce » 663.

Subordo: Teleostei Physostomi. » 689.

Series: Glanomorphi . — » 690.

Familia: Siluridce . . .. - » 692.

Series: Cyprinomorphi. — » 702.

Familia: Vobitidce » 703.

» Cyprinidce » 714.

Series: Thrissomorphi » 826.

Familia: Salmonidce — » 827.

» Scopelidce - » 920.

» Chipeidce ... » 946.

Series: Esociformes 1. Lycomorphi pag. 91)7’.

Familia: Esocidce . » 997.

Series: Enchelyrnorphi .. » 1011.

Familia: Anguillidoe » 1022.

Ordo: PISCES CHONDROSTEI » 1043.

Familia: AcipenseiHdce . » 1044.

Ordo: PISCES ELASMOBRANCHII .... » 1063.

Subordo: Elasmobranchii Holocephali . » 1078.

Familia: Chimceridce » 1078.

Subordo: Elasmobranchii Plagiostomi- » 1085.

Phalanx: Plagiostomi Batoidei » 1086.

Familia: Myliobatidce » 1093.

» Trygonidce » 1096.

» Rajidce ... » 1100.

Phalanx: Plagiostomi Selachoidei.. » 1127.

Series: Asterospondyli » 1128.

Familia: Garchariidce... » 1128.

» Lamnidce » 1135.

» Scylliidce » 1147.

Series: Cyclospondyli » 1156.

Familia: Spinacidce » 1157.

Ordo: PISCES CYCLOSTOMi » 1172.

Familia: Petromyzonidce » 1172.

» Alyxinidce » 1195.

Ordo: PISCES LEPTOCARDIP. » 1210.

Familia: Amphioecidce » 1211.

SAND-EELS.

567

I am. AMMODYTIDfE.

Body elongated , fusiform , terete or compressed, covered with thin cycloid scales or partly naked. Caudal fin
separated from the other vertical fins. Jaws without teeth*. G'dl-openings large: branchiostegal membranes more
or less completely free from each other and from the isthmus. Pseudohranchice well- dev eloped and distinct.

Air-bladder wanting. Pyloric appendages rudimentary .

The place of this family in the system has long
been a debated question. The original opinion of mo-
dern systematists, that of Aetedi, was that the Sand-
Eels — with their long dorsal tin occupying the greater
part of the back — should be ranged beside the genus
Coryphcena. This opinion, borne out by the Mackerel-
like coloration of the Sand-Eels, still survived in 1839
in Swainson b, who pointed out the external resemblances
between these fishes and Lepidopus. Linnjeus had ima-
gined that he had made an improvement by uniting
all fishes without ventral fins into an order ( Apodes ),
and thus in 1817 — 1829 the Sand-Eels assumed in
Cuvier’s works" the rank of a genus within the family
of the Eels, and in 1832 — 1841, in Bonaparte**, that
of a subfamily ( Ammodytini ) of the Ophidiidce among
Malacopterygii apodes. When MulleiC formed the order
Anacanthini , he did not hesitate to include in this or-
der the family Ophidiidce , but he declared himself
unable to give a decided opinion as to the place of
Ammodytes, though he positively denied the relationship
between this genus and the Eels. In his later work7,
however, Bonaparte ranged the Sand-Eels, as a distinct
family (Ammo dy tides), among the Gadi, and Gunther0,
did not hesitate to include these fishes among the
Anacanthini as a subfamily (Ammodytina) of the Ophi-
diidce.

This diversity of opinion has been caused by the
absence in the Sand-Eels both of the ventral fins and the

air-bladder. The reduction and eventual disappearance
of the ventral fins is a characteristic which, as we have
seen above, may occur within several piscine orders.
The absence of the air-bladder may be explained in
the same way. We must, therefore, look to other
characters in order to discover the nearest relations of
the family, and as often happens in such cases, we may
have recourse to characters apparently of minor im-
portance. The Sand-Eels are approximated by their
form and coloration not only to the Mackerels but,
still more closely, to the Garpikes. To the latter fishes,
which are Pharyngognate Anacanthini, it is impossible
to unite the Sand-Eels, in which the lower pharyngeals
are free from each other. Still, where the lateral line
lies in the Garpikes and Elying-fishes, along the ventral
margin, at the boundary between the sides and the
belly, and also along the base of the anal fin, here we
find in the Sand-Eels a dermal ridge, somewhat raised
in the same way. The physiological signification of
this ridge is indeed unknown, but in a morphological
respect it shows at least a trace of resemblance to the
Garpikes and Flying-fishes, such a resemblance as we
have seen above in other Anacanthini, in the Couchia-
stage of Onos. The Garpike-like coloration of these
last fishes, in its sharp contrast to that of adult Rock-
lings, may also be a trace of their original relationship
to the Sand-Eels. Among the osteological peculiarities
which we have above remarked in the Gadoid family,

a Day ( Fishes of India, p. 420) observed, however, “a few, fine teeth opposite the symphysis in either jaw” in Ammodytes ( Blee -
keria ) kallolepis.

b Nat. Hist. Fish., Amplu, Rept., vol. II, p. 254.
c R'egne Animal , tome II, ed. 1, p. 240; ed. 2, p. 360.
d Iconogr. Fn. Ital., tom. Ill, Pesci , Introd., p. 15.
e Abh. Akad. Wiss. Berk 1844, p. 177.

I Uatalogo Metodico dei Pesci Europei , Napoli 1846, pp. 6 and 40.
g Cat. Brit. Mus., Fisli, vol. IV, p. 384.

72

Scandinavian Fishes.

568

SCANDINAVIAN FISHES.

there recurs in the Sand-Eels the characteristic, lobate
process on the intermaxillary bones, though it is re-
moved farther forward, to about the end of the first
third ( Ammodytes tobianus ) or fourth (Am. lanceolatus )
of the length of the bones. This resemblance may be
not without importance in a morphological respect. With
this exception, however, the intermaxillary bones of the
Sand-Eels are very unlike those of the Codfishes, not
only in their narrow and terete, almost needle-like
shape, but also in the more or less complete freedom
of the nasal processes, which vary considerably in length
and mobility, and are united to the anterior end of
the bones only by cartilage and ligaments. The ske-
leton of the Sand-Eels is also distinguished from that
of the Codfishes in two other essential respects. Ribs
are attached to the abdominal vertebral from the very
first of these bones; and in the caudal fin, which is far
more differentiated than in the Codfishes, and the base
of which is composed exclusively of the last two ver-
tebras and the urostyle, the development of the hypural
bones is quite as typical as in the rest of the Teleosts.

Still, though we may find in the above-mentioned
points of resemblance to the Codfishes and the Gar-
pikes fully valid reasons of morphological significance
for the opinion advanced by Gunther and other mo-
dern writers, that the Sand-Eels are Anacanthine fishes,
we are not destitute of grounds for a close comparison
of these fishes with the Eels, though the latter are
assigned to a far distant place in the system by the
arrangement of the jaws and shoulder-girdle, as well
as by their character of Physostomous fishes. This
comparison is suggested by the scales. Their structure
most strongly reminds us partly of the Eels and partly
of Enchelyopus. In the Scandinavian Sand-Eels, as in
most fishes, and in a manner that especially calls to
mind the simple scales of our common Flatfishes, the
anterior part of the scale is quite different from the
posterior; but here the difference is so marked that the
former resembles in structure an entire scale of Enche-
lyopus, with dense, concentric stria?, interrupted by

grooves radiating from the nucleus, while the latter
resembles the scales of the Eels, with continuous con-
centric striae, but with the grooves broken up into
more or less irregular, round or oblong, small patches.
On the anterior part of the scale too, the concentric
striae are about twice as dense (numerous) as on the
posterior part. The scales of the Sand-Eels also vary
considerably in form and development, not only in
different species but even in the same fish. All of
them are comparatively small and thin. The most
developed are set on the back of the fish, the largest
of them, as usual, on the hind part of the body. These
scales are imbricated, with the posterior part free; and
they vary in shape from rounded to oblong or lingui-
form. On the sides of the body the scales lie in der-
mal folds that run in an oblique transverse direction
downward and backward from the lateral line proper,
which is situated high on the back, to the raised dermal
ridge that coasts each side of the belly, forming a
boundary between the latter and the sides of the body.
On the belly itself, between these dermal ridges, the
folds are less sharply marked, but the scattered (not
imbricated) scales lie hidden in the skin, in rows that
run from each dermal ridge obliquely forward and in-
ward, towards a similar but lower dermal ridge at the
middle of the belly. The scales which lie in the skin
are of a broad oval (linguiform) shape, with the hind
extremity pointed".

In their manner of life the Sand-Eels remind us
both of the Garpikes and of the Eels: in the open sea
they are active and eager in their pursuit of small
fishes and fry, but now and then they hide themselves
in the sand to escape their numerous enemies, just as
the Eels burrow in the sand and mud or creep into
crevices between the stones.

The family contains remarkably few forms: among
the 4 or 5 known species'' only two genera can with
reason be distinguished. Most of the species belong to
the Northern Hemisphere, both to the Atlantic and the
Pacific; but one species occurs in the Indian Ocean.

a In the Indian species, which Gunther has adopted as the type of a distinct genus ( Bleekeria ), the scales are said to be larger (of
moderate size), and the said dermal ridges are wanting.

6 Five distinct species have been described from America. But the Greenland species is undoubtedly identical with our Sand-Launce,
and Jordan and Gilbert (Bull. U. S. Nat. Mus., No. 16, pp. 414 and 909), who have rejected one species as based merely on a description
of a damaged specimen, advance the opinion that all the others are hardly more than varieties of the Sand-Launce. Brown-Goode, however
{Fisher., Fisher. Industr. U. S ., sect. I, p. 244), insists upon maintaining a rigid distinction between the European and American Sand-Eels.

SAND-EELS.

569

Genus AMMODYTES.

Body more or less terete ( only slightly compressed).

Branchiostegal

The extremely similar species that form this germs,
have long been known. At the middle of the fifteenth
century Salvianus described and figured the ‘ sandilz
(Sand-Eel) of English waters, and gave an account of
its habits and the manner in which it was taken. The
figure and description recur in Gesner", who gave the
genus its Greek name* 6 * * 9. Ray6 and JagcG mention two
species; but although we can see, from the statement of
the latter as to the length of the fish described and figured
by him (1572 Gi.), that he referred to a species regarded
as .distinct in modern times {Ammodytes lanceolatus), still
the supposed specific distinction was based merely on
defects in the figure of Salvianus. Artedi described6
the smaller of our forms both fully and accurately,
but did not distinguish between it and the larger one.

In his Sy sterna Natures f Linnaeus adopted only one spe-
cies {Am. Tobianus), the name of the species being
clearly derived from Artedi’s quotation of Sciione-
velde?; but in his Fauna Suecica h he draws attention
to the fact that in his travels through Oland' he had
found more fin-rays in the Sand-Eel than Artedi, to
which fact he probably refers when he remarks, in the

A longitudinal dermal ridge on each side of the belly,
or 8.

twelfth edition of his Systema (1766): “In Sweden there
seem to be two distinct species, as Ray once conjec-
tured was the case in England.” There is no doubt,
however, that in 1744-' Klein had already distinguished
between Artedi’s Tobianus and Jago’s “ Ammodytes
Anglorum verus, The. Bounce, sive True Sand-Eel’
though without giving them binomial names. After-
wards, in 1810*, when Rafinesque described his Ammo-
dytes cicerelus, the three European species now accepted
became known; but the largest of them was without
any binomial specific name, which it first received in
1824 of Lesauvage'. The natural relations between
these three species, as they are distinguished at present,
may be expressed as follows:

A: Pectoral fins shorter than the lower jaw:
a: The whole body behind the head
scaly, with the scales on the sides
of the body hidden in transverse

dermal folds even on the forepart Ammodytes lanceolatus.
b: Forepart of the body naked, with-
out transverse dermal folds Ammodytes cicerelus.

B: Pectoral fins longer than the lower

jaw Ammodytes tobianus.

rays 7

a Paralipomena , p. 3.

6 From ay yog, sand and dvzyg, diver.
c Syn. Meth. Pise., p. 38.

d Ibid., p. 165, fig. 12.
e Spec. Pise., p. 55.

f Ed. X, tom. I, p. 247 ; ed. XII, tom. I, p. 430.

9 To which form Schoneveede himself referred, is a question difficult to decide. Ichth. Slesv. Hols., p. 76.
h Ed. I, p. 114; ed. II, p. 109.
i It. CEl., p. 87.

j Hist. Pise. Nat. Prom. Miss. IV, pp. 55 and 56, tab. XII, figs. 8 — 10.
k Caratteri d. ale. n. gen., p. 21, Tab. IX, fig. 4.

^ Bull. Sc., Soc. Philom. 1824, p. 140.

570

SCANDINAVIAN FISHES.

THE SAND-EEL (sw. hvittobisen or tobiskungen).

AMMODYTES LANCEOLATUS.

Plate XXIII, fig. 4 and Fig. 135.

Upper jaw not at all (or only slightly ) protrusile: nasal processes of the intermaxillary hones short — measuring
about 11 — 13 % of the length of the intermaxillary hones — and more or less firmly united hy ligaments to the
ethmoid hone. Pectoral fins short, their tip extending scarcely to the perpendicular from the beginning of the dorsal
fin, and their length being less than that of the lower jaw, less than 46 % of that of the head, and at most 33 %
of the distance between the dorsal fin and the tip of the snout. Head of the vomer f urnished with two pointed,
crooked teeth, diverging at the tip. Dorsal and anal fins with straight margin. A black spot generally visible

on the middle of the sides of the snout.

Fig. 135. Ammodytes lanceolatus, with the body in transverse section, a scale, and a head with open mouth. After Bhnecke.

R. hr. 7; D. 53— 59«; A. 28 6 — 33; P. 12—14; V. 0;
C. *+l + 13 + l+jc; Lin. lat. 172 — 1 8 3 c ; Vert. 66 — 67.

Syn. Enchelyopus No. 7; Klein, 1. c., p. 56, tab. XII, fig. 10;

Ammodytes tobianus, p. p., Swartz, Svensk Zoologi , H. 11,
No. 64.

Ammodytes lanceolatus, Lesatjv. 1. c.; Sundev., SIcand. Fisk.,
ed. 1, p. 209, tab. 54; Gthr, Cat. Brit. Mus., Fish., vol.
IV, p. 384; Malmgr., Finl. Fisk. (disp. Plels.) p. 32; Coll.,
Forb. Vid. Selsk. Christ. 1874, Tillsegsh., p. 126; ibid. 1879,
No. 1, p. 69 (corr. N. Mag. Naturv. Christ., Bd. 29 (1884),
p. 94); Malm, Gbgs, Boli. Fn., p. 500; Winth., Naturh.
Tidskr. Kbhvn, ser. 3, vol. XII, p. 35; Bncke, Fisch.,
'Fischer., Fiscliz. 0., W. Preuss., p. 99; Mor., Hist. Nat.
Poiss. Fr., tom. Ill, p. 217; Day, Fish. Gt. Brit., Irel.f
vol. I, p. 329, tab. XCII, fig. 1; Mob., Hoke, Fisch. Osts.,
p. 85; Lillj., Sv., Norg. Fislc., vol. II, p. 211.

Ammodytes tobianus, Cuv., Regne Anim., ed. 2, tom. II, p.
360; Nilss., Prodr. Iclxthyol. Scand., p. 63; Ekstr., Vet.-
Akad. Handl. 1834, p. 67; Kr., Damn. Fislce, vol. Ill, p.
575; Nilss., & ’hand. Fn., Fisk., p. 653.

The Sand-Eel is the largest Scandinavian species
of this genus, and attains a comparatively considerable

size: Sundevall met with a specimen 32 cm. long;
Day gives 33 cm., and Jago (in Ray) 394 mm., as
the maximum length of the species. The ordinary length
on our coasts, however, is only 12 — 20 cm.'

The body is elongated, the greatest depth, across
the belly, measuring in ordinary cases only about 6 or
7 % of the length of the body to the end of the outer-
most caudal rays, though in gravid females the belly
is of course deeper. The body is also of nearly uni-
form depth, with gradually tapering tail and conical,
pointed head, and almost terete, the greatest breadth
being usually at least more than 3/i (81 — 84 %) of the
greatest depth. The compression of the tail is confined
to the base of the caudal fin.

The conical head is also only slightly compressed,
but above and on the sides somewhat flattened and
smooth, without scales. The depth across the occiput
is someAvhat less than the greatest depth of the body.

° Sometimes 61, according to Gunther and Moreau.

,, 26, ,, ,, Lilljeborg.

e ,, 169, ,, ,, Malm.

,, 203, „ „ Day.

SAND-EEL.

571

The length from the tip of the upper jaw to the occi-
put is twice the depth at the occiput, and in a small
specimen (about 15 cm. long) nearly Vs °f the total
length (including the whole of the caudal fin). The
length of the head — according to the method of mea-
suring it usually employed in this work, from the tip
of the snout to the hindmost point of the opercular
(here the subopercular) flap — measures in specimens
between 11 and 14 cm. long about 21 % (20'8 — 20‘7
%, according to our measurements) of the length of the
body (from the tip of the snout to the end of the
middle caudal rays), and in specimens between 26 and
28 cm. long about 19 % (18*5 — 19*6 %) of the same
length. The eye is fairly round, but closely surrounded
by a sharp, soft orbital margin, the opening of which
is usually transversely oblong (the height greater than
the length). Both in large and small specimens the
hind margin of the eye lies half-way between the tip
of the lower jaw and that of the gill-cover; but in
large specimens the size of the eye is less in proportion
to that of the body than in smaller ones, the diameter
of the eye in a specimen 30 cm. long being only twice
as great as in a specimen 7 V2 cm. in length. During
the growth of the body from 11 to 28 cm. the longi-
tudinal diameter of the eye decreases from about 3 1/2
to l3/4 % of the length of the body or from 16 to 9 %
of the length of the head from the tip of the snout.
The length of the snout measures about 60 % (61 V2 —
5 9 1/ 2 %) of the postorbital length of the head, or about
V3 (324/2 — 35 %) of the total length of the head from
the tip of the snout, in specimens between 11 and 28
cm. long. The breadth of the interorbital space, on
the other hand, shows even relative increase during
growth, in the said specimens from about 2 V2 * to
about 33/4 % of the length of the body, or from 1 2 1 / 2
to I8Y3 % of the length of the head.

The nostrils are small, oblong, not raised, and only
slightly more than twice as large as the small muci-
ferous pores which retain their normal position in the
supraorbital, suborbital, and preoperculo-mandibular
branches of the cephalic system of the lateral line. The
anterior nostrils are set on each side somewhat behind
the middle point between the eye and the tip of the
snout, the posterior somewhat higher but much nearer
to the anterior nostrils than to the eye. The tip of the
snout, seen from above, is parabolic, flattened, and flex-
ible; it projects considerably, almost concealing the upper
jaAV (the intermaxillaries) when the mouth is closed.

The mouth is rather large, but capable of hardly
any protrusion. When the jaws are opened, the upper
jaAv thus assumes a vertical position and bends the tip
of the snout upwards. In young specimens, however,
the mouth is someAvhat protrusile. The intermaxillary
bone is rather narroAv, and extends someAvhat behind
the nostrils Avhen the mouth is closed. The maxillary
bone is much more robust; it articulates beloAV the sides
of the tip of the snout and, Avhen the mouth is closed,
lies hidden under a dermal fold of the cheek (the lower
preorbital margin). The total length of the upper jaw,
from the tip of the snout to the hind extremity of the
maxillary bones, increases Avith age, in the specimens men-
tioned above — between 11 and 28 cm. long — from 6 to
Q2/3 % of the length of the body or from 29 to 34 % of that
of the head. The loAver jaAV is fairly strong and much
longer than the upper. In the specimens just mentioned
its length measures about 7m (10’2- — 9‘1 %) of that of
the body or about 7a (46*8 — 50 %) of that of the head,
and is greater than the length of the pectoral fins
(113 — 155 % thereof). Its articulation lies vertically
beloAV the eye, and its tomial margin is deeply sinuated.
The conical tip of the loAver jaAV, Avhich projects beyond
the upper jaw, and thus forms the tip of the head,
measures as much as 7 5 — Vi °f the snout proper, or
even a little more. The lips are Avell-developed. Trans-
verse palatal folds are present both in the upper jaAV
and in the lower. Teeth are entirely wanting in the
jaAvs, on the palatine bones, and on the tongue. O11
the head of the vomer alone Ave find two teeth, Avhich
are strong, set close together, and curved. The pha-
ryngeals, Avhich are small (3 above and 1, elongated in
form, beloAV on each side), are furnished Avith close-set,
fine, cardiform teeth. The gill-rakers are setiform,
numbering about 25 on the front surface of the first
branchial arch. The tongue is of a shape not so com-
mon among fishes: it is free, broad but thin, and con-
cave at the top, Avith depressed, rounded tip.

The gill-openings are large, nearly the Avhole of
the gill-cover being free at the top, Avhile beloAV the
opening is continued doAvn to the articulation of the
loAver jaAV. The branchiostegal membrane is thus com-
pletely divided, without being united to the narroAv
front part of the belly (the isthmus) between the gills.
It is furnished with 7 rays, which are, hoAvever, diffi-
cult to count, as the outermost ray is rather short and
fine, and is hidden by the extraordinarily strong muscles.
The rays are covered by the margin of the fold and

572

SCANDINAVIAN FISHES.

extend nearly to its extremity. The opercular appa-
ratus as a whole is fairly large, but the operculum
proper is the smallest part (about 1/3) thereof. The
operculum is smooth, convex, triangular, and almost
equilateral. The greatest part of the opercular apparatus
consists of the suboperculum, which forms the whole
of the inferior and posterior parts, and with its broadly
rounded point projects somewhat behind and above the
base of the pectoral tin. It is united to the operculum
by an oblique, straight line; the lower margin forms
a large, rounded sinus, to receive the base of the pec-
toral tin; and the surface is smooth, with 9 distinct,
radiating lines on the lower part. The four branchial
arches are complete, and a -well-developed pseudobranchia,
composed of from 12 to 14 threads, occupies the inner
surface of each gill-cover.

On the body itself Ave observe several longitudinal
lines, first the lateral line proper, Avhich lies high on
the back, the distance between it and the dorsal fin
being about 1/2 of that between it and the median line.
The lateral line is parallel to the dorsal tin, straight,
and depressed in the rather large scales, each of which
contains a branch of the muciferous canal Avith its
opening. The line begins above the gill-cover, Avithout
its usual external continuation on the head, and ends
at the termination of the dorsal tin. — Along the middle
of the sides there runs a someAvhat depressed median
line along the middle angles formed by the aponeurotic
septa of the large lateral muscles. This line is quite
straight and consists of rather small dots, pricked, as j
it Avere, Avitli a needle, and invisible Avithout the help
of an ordinary magnifying-glass. They form an irre-
gular double (or multiplex) toav, equal in Avidth to half
a scale. The line resembles a lateral line, but is less
marked and destitute of muciferous ducts. — On the
sides of the belly Ave find a still more remarkable carina
or dermal fold, which begins beloAv the base of the
pectoral fin, folloAvs a perfectly straight course, and gra-
dually disappears half-Avay between the vent and the
caudal fin. Its height is about equal to the greatest
breadth of a scale; it is thin and soft, and does not
seem to contain any muciferous canal; but, in speci-
mens preserved in spirits at least, it lies with the free
margin bent outwards, covered by the mucus of the
skin, and thus appears itself to surround a canal. — On
the belly, between the tAvo lateral carina?, three fine
longitudinal lines run from beloAv the pectoral fin to
the vent, the tAvo lateral ones slightly depressed, but

the middle one very fine, somewhat raised, of clear
colour, and scaleless. In middle-sized specimens the
anal aperture lies at the end of the third fifth of the
length of the body, in the largest specimens a little
farther back, its position thus groAving more posterior
Avith age. It is only slightly, if at all, prominent,
hardly more so than the smaller opening for the genital
organs that lies behind it.

The dorsal fin is of almost uniform height, its
height in large specimens being scarcely half the depth
of the body. It begins at the end of the first fourth
of the body, or usually a little farther back, about as
far (in young specimens someAvhat less, in old someAvhat
more) behind the occiput as the latter from the tip of
the snout. It ends some Avay from the caudal fin, the
distance betAveen them being about equal to the least
depth of the tail, or a little greater. The rays are
about 56 in number, the variations being apparently
confined to 2 or 3 more or less; all the rays are fine
and articulated, but perfectly simple and pointed though
not pungent. The membrane is thin and fragile.

The anal fin begins just behind the genital opening
and ends opposite to the termination of the dorsal fin.
It is equal in height to the latter, and contains about
30 rays of the same structure as those of the dorsal fin.

The caudal fin is deeply forked; its greatest length
is about equal to the depth of the body. It contains
15 true (articulated) rays. Thirteen of these are bran-
ched, though not very deeply, the outermost ray above
and beloAv being simple and only slightly shorter than the
corners of the fin. Outside these rays there lie several
small, unarticulated or supporting rays: in a specimen
3 dm. long Sundevall counted 1 1 above and 9 beloAv.

In middle-sized specimens the pectoral fins do not
extend further back than to the perpendicular from the
first ray of the dorsal fin. In the very largest speci-
mens they are relatively much shorter. In the speci-
mens mentioned above, between 11 and 28 cm. long,
the length of the pectoral fins varies Avith increasing
age between 9 and 6 1 / 3 % of that of the body, from
33 to 22 % of the distance betAveen the dorsal fin and
the tip of the snout, or betAveen 81 and 56 % of the
postorbital length of the head. They are also invariably
shorter than the loAver jaw, their length varying with
increasing age in the said specimens betAveen 88 and
64 % of that of the latter. They contain 14 rays, all
articulated; the first is simple and measures only slightly
more than half the length of the fin, the second is in-

SMOOTH SAND-LAUNCE.

573

distinctly branched at the extreme tip, the others are
doubly branched, and the fifth (4 — 6) is the longest.
— The ventral fins are wanting.

The small scales cover the entire body, but not the
head. On the back and belly they are set more closely
and irregularly, on the sides of the body they lie ar-
ranged in the regular, oblique, simple rows described
above. These rows run from the lateral line on the
back, obliquely backwards, down to the lateral ventral
carinas. Their number varies considerably: in a large
male Sundevall found only 172; in another large spe-
cimen we found 176, in a middle-sized one 183; Kroyer
has found as many as 195. The usual number is pro-
bably about 180. The first ones, about 5, above the
gill-cover, and the last 6 or 7, at the caudal fin, are
short and incomplete. Furthermore, on the front and
middle parts of the body these rows are a little way
apart from each other, while on the tail they are set
close together. In each row the scales are set so den-
sely that they overlap each other to a small extent of
their surface. At the middle of the body each row
contains about 46 scales, 18 or 19 of which lie between
the lateral line and the median line, the other 27 or
28 between the median line and the lateral carina on
the belly. In the largest specimens (about 3 dm. long)
7 scales in one of these rows occupy a space 5 mm.
long.

The coloration is greenish on the back and white
on the rest of the body, shifting into all the colours of
the rainbow; but in the living fish the whole body is
somewhat transparent. All the fins are pale. A large
blackish spot occurs on the sides of the snout, half-way
between the tip of the snout and the eye.

No distinct external difference between the sexes
has been discovered.

With regard to the internal organs we merely re-
mark that the peritoneum is white with a silvery lustre,
the stomach rather small, firm, and wide, with a long,
narrow continuation in a backward direction, which
when empty resembles a narrow, cylindrical appendage,
but when full of food becomes rather large and extends
back to the anal region. The liver is small and white, and
lies in front of the stomach, round the oesophagus. The
intestine is narrow, with a sharp bend. Only one short,
conical appendage occurs at the pylorus. The ovaries
are separate in front, but towards the vent completely
united into one mass. The eggs are flame-yellow, fairly
numerous, and fine: their diameter is only slightly more
than l/s urn i . , 7 ripe eggs occupying a space 4 mm. long.

As all the Sand-Eels, so far as is known, live in
the same manner, and as the two species that occur in
any abundance in Sweden, are found in company with
each other, we shall shortly return to the habits and
development of the Sand-Eel.

THE SMOOTH SAND-LAUNCE (sw. slattobisen a).
AMMODYTES CICERELUS.

Fig. 136.

Upper jaiv protrusile. Pectoral fins short , as in the preceding species. Head of the vomer toothless. Dorsal and
anal fins with undulating margin. No regular transverse folds across the sides of the body.

Fig. 136. Ammodytes cicerelus, natural size. Taken at a depth of 30 fathoms off Grip (Norway), by Lilljeborg.
Specimen in the possession of the Zoological Museum of Upsala University.

Ammodytes cicerelus , Rafin., Caratt , d. ale. n. gen ., p. 21,
tab. IX, lig. 4; Mor., Hist. Nat. Poiss. Fr., tom. Ill, p.
219; Day, Fish. Gt. Brit., Irel., vol. I, p. 333, tab. XCII,
fig. 3; Lill.j., Sv., Norg. Fisk., vol. 2, p. 228.

a Lilljeborg, 1. c.

6 According to Day, Moreau, and Lilljeborg.

15 according to Costa.

16 ,, ,, Swainsok.

c According to Costa.

R. br. 7; D. 53—59; A. 28—30; P. 13— 14* 6; V. 0;
C. x + 1 + 1 3 + 1 + x ; Vert. 6 8 c.

Syn. Gicirellus Messanensis, Boccone, Recli., Observ. Nat., Amster-
dam 1074, p. 294, fig. in tab. ad p. 287.

574

SCANDINAVIAN FISHES.

Ammodytes tobianus, Risso, Ichth. Nice , p. 95; Costa, Fn.

Nap., Pesci, Gen. Ammod., pp. 1 — 10, tab. LI.

Ammodytes Siculus, Swainson, Zool. Illustr., vol. I, tab. 63;

Bonap., Cat. Met. Pesci Fait., p. 40; Gtiir, Cat. Brit. Mus. ,

Fish., vol. IV, p. 386; Ann. Mag. Nat. Hist., ser. Ill, vol.

XX, p. 290; Steind., Stzber. Akad. Wiss. Wien. Math. Naturw.

Cl., vol. LV1I, i (1868) p. 712, tab. II, fig. 3; Gigl., Espos.

Intern. Pesca, Berl. 1880, Catal. Sez. Ital., p. 97.

The chief characteristic of this species is the weak
development of the scales. Distinct scales occur only
on the hind part of the tail, and the sides of the body
are without the oblique, transverse, dermal folds in
which the rest of the scales are hidden in the other two
species. To this we may add the apparently charac-
teristic shape of the dorsal and anal tins". The former
is furnished with two sinuses, one in the front part of
the upper margin and one in the hind part thereof;
while the anal tin contains one sinus, corresponding to
the posterior one in the dorsal fin. In other respects
the Smooth Sand-Launce is an intermediate form be-

tween our other two species: the pectoral tins are short
as in the Sand-Eel, but the structure of the mouth and
of the head of the vomer is the same as in the Sand-
Launce. The back is greenish (olive) or bluish. A broad
band with a strong, silvery lustre extends along the
whole length of the sides. A blue spot often occurs,
according to Moreau, on the top of the head. The snout
is flesh-coloured, according to Costa. The iris is sil-
very white.

According to Gunther, Moreau, and Giglioli the
Smooth Sand-Launce is the only Mediterranean species
of the genus. Gunther states that in 1867 Gwyn-
Jeffreys took a specimen 15 cm. long in a dredge, off
the Shetland Islands, in 80 or 90 fathoms of water. In
1858 and with the same tackle Lilljeborg took a spe-
cimen 12V2 cm. long, on a bottom of shell sand, at
a depth of about 30 fathoms, off Christiansund in Nor-
way. The former specimen had attained the maximum
size assigned to the species by Moreau.

THE SAND-LAUNCE (sw. blatobisen).

AMMODYTES TOBIANUS.

Fig. 137.

Upper jaw protrusile: nasal processes of the intermaxillary hones long — measuring about 2/3 (64 — 67 %) of the
length of the intermaxillary hones — and mobile , with a gliding motion, in the groove in the upper surface of the
ethmoid hone. Pectoral fins comparatively long, their tip extending distinctly behind the perpendicular from the
beginning of the dorsal fin, and their length, being greater than that of the lower jaw, more than 46 % (47 — 60
%) of that of the head , and at least 37 % (37 — 43 %) of the distance between the dorsal fin and the tip of the
snout. Head of the vomer toothless. Dorsal and anal fins of uniform height.

Fig. 137. Ammodytes tobianus,

with the body in transverse section, a scale, and a head with opened mouth. After Benecke.

R. hr. 7; I). (51) 55 — 62 (64); A. (26) 29—33; P. 12—15;
V. 0; C. x + 1 + 13 + 1 + x ; L. lat. 120 — 145; Vert. 62 — 63.

Syn. Enchelyopus No. 6, Klein, 1. c., p. 55, tab. XII, fig. 8 et 9 ;

Ammodytes, Art., 1. c.; The Launce , Penn., Brit. Zool.
(London, Warrington, 1776) vol. Ill, p. 137.

Ammodytes tobianus, Lesauv., 1. c.; Gthr, 1. c., p. 385;
Steind., 1. c.; Coll., Forh. Vid. Selsk. Christ. 1874,
Tilleegsh., p. 126; Malm, Gbys, Boh. Fn., p. 500; Win-
ther, 1. c.; Bncke, 1. c., p. 100; Mor., 1. o., p. 218;
Day, 1. c., p. 331, tab. XCII, fig. 2; Mob., Hcke, 1. c.,

p. 86.

This character is not noticed by Costa, either in his description or figure.

SAND-LAUNCE.

575

Ainmodytes lancea , Cuv., Regne Anim., ed. 2, tom. II, p. 300;
Nilss., Proclr. Ichthyal. Scand ., p. 03; Kb., 1. c., p. 593;
Nilss., Skaiul. Fn., Fisk., p. 650; Malmgr., 1. c.; Lindstr..
Goil. L. Hush. Sallsk. Arsber. 1860, p. 24 (sep.); Mela,
Vert. Fenn., p. 296, tab. IX; Lillj., I. c., p. 221.

The Sand-Launce never attains so great a size as the
maximum size of the Sand-Eel. K royer’s largest spe-
cimen was 187 mm. long; a specimen from Greenland,
brought home by O. Torell, measures 204 mm. from
the tip of the lower jaw to the end of the longest rays
of the caudal tin, or 197 mm. from the tip of the snout
to the end of the middle rays of the same fin.

The body is generally somewhat deeper and more
compressed than that of the Sand-Eel, though no constant
character can be drawn from this relation. The greatest
depth of the body, across the belly, varies in ordinary
cases between about 9 and 10 V8 % of the length from
the tip of the lower jaw to the end of the outer rays
of the caudal tin, and the greatest breadth between
about 66 and 75 % of the greatest depth. We have
found the depth at the beginning of the anal fin to vary
between 6 1/5 and 9 vi %a of the length of the body
from the tip of the snout to the end of the middle
caudal rays; and, in the same specimens, the greatest
breadth varied between 4 and 773 %h of the same length.

In specimens of equal size the head is of essen-
tially the same shape as in the Sand-Eel; but as a rule
it is shorter. In specimens between 95 and 197 mm.
long we have found its length to vary between 19 and
173/4 % of that of the body. The longitudinal diameter
of the eye varied in these specimens between 20 and
13 % of the length of the head. The length of the
snout varied between 64 and 56 % of the postorbital
length of the head or between 33 and 30 % of the total
length thereof. The least breadth of the interorbital
space varied between 2V3 and 3 2/5 % of the length of
the body or between 121/ 2 and 1972 % of the length
of the head.

In the nostrils we find scarcely any difference from
those of the Sand-Eel; but the structure of the mouth,
as we have remarked above, affords one of the most
important characters for the Sand-Launce and the forms
akin to it in this respect. The mobility of the inter-
maxillary bones is produced in the same manner as in
the case of most other fishes, and in striking contrast
to the Sand-Eel. The difference lies partly in the long

a In the Sand-Eel between 62/3 and 8 %.

b al, o/

1111 ii ii ° 13 ii ° 13 /”•

c ,, ,, ,, from 16’3 % to 15'3 %.

nasal processes, which glide forward and backward in a
groove in the upper surface of the ethmoid bone, partly
in the greater length of the muscular band that extends
under each of these processes, starting from the inner
(posterior) surface of the front of the intermaxillary
bone and from its articular process (cf. the explanation
of fig. 118, p. 463, above), and attached to the inward
vomeral process of the maxillary bone, which process is
closely united by ligaments to the anterior end of the
vomer. These two muscular bands serve to draw back
the intermaxillary bones. The protrusion of the inter-
maxillary bones is effected by their union to the lower
jaw by skin, muscles, and ligaments, and they thus
follow the lower jaw when it is depressed. The anterior
end of the maxillary bones is rendered independent of
this downward motion by the union of the vomeral
processes to the vomer; and on casual examination these
processes, with their sharp, transverse, osseous points,
present a confusing resemblance to the teeth on the
head of the vomer in the Sand-Eel — they have several
times been confounded with those teeth. The length of
the upper jaw shows even relative increase with age
from the earliest stages until the fish is of middle size,
but it subsequently seems to be retrogressive, unless
our observations are based on individual variations. In
a specimen 30 mm. long we find this length to be 2 2 2/3
% of that of the head, and in larger specimens, up to
a length of about 13 cm., this percentage increases to
about 33; but in our largest specimen the proportion
is no more than 28 V2 %. The lower jaw is usually
shorter than in the Sand-Eel, its length being about
8 % (between 8'6 — exceptionally 9'3 — and 7‘6 %) of
that of the body, or about 45 % (between 47'8 — in our
smallest specimen 49 — and 42*8 %) of the length of
the head, and never so much as 15 % ( 14*2C — 12*2 %)
of the length of the base of the dorsal fin. The conical
tip in which the lower jaw projects beyond the tip of
the snout, is also generally less than in the Sand-Eel,
and seems at most not to exceed V4 of the length of the
snout proper. The gill-rakers are setiform in this species
too, and number 22 or 23 on the first branchial arch.
The lips, tongue, palatal folds, pharyngeal bones, and
gill-covers resemble those of the Sand-Eel, but the oper-
culum is more scalene, with the lower side (along the
suboperculum) perceptibly greater than the upper. The

Scandinavian Fishes.

73

576

SCANDINAVIAN FISHES.

lateral line contains about 125 scales; and the oblique
dermal folds on the sides of the body number about
130, being thus fewer than in the Sand-Eel. It is also
easy to see, on comparing specimens of equal size, that
the scales of the Sand-Launce are larger. The anal
aperture lies nearer the middle of the body, the distance
between it and the tip of the lower jaw measuring,
according to Kroyer, about 53 — 56 % of the length of
the body to the end of the outermost caudal rays. We
have found the distance between it and the tip of the
snout, in specimens from 97 to 130 mm. long, to rise
from 5 6 3/4 to 5872 % of the length of the body to
the end of the middle caudal rays'*.

The dorsal fin generally begins somewhat further
forward than in the Sand-Eel, usually in front of the
end of the first fourth of the length of the body, but
sometimes behind it. On the other hand, its extent
is apparently always somewhat greater, sometimes as
much as 2/3 of the length of the body. Consequently,
the distance between it and the tip of the snout, accord-
ing to our measurements, never exceeds 40 % of the
length of its base6. As a rule too, its length increases
even relatively with age, the length of the head in
specimens about 9 cm. long measuring 28' 7 % of the
length of the base of the dorsal fin, in specimens be-
tween 13 and 20 cm. long only 26*7 % thereof0. The
anal fin generally begins somewhat further forward and
in most cases is of somewhat greater extent than in
the Sand-Eel; but in both species the length of its
base varies between 48 and 43 % (in the Sand-Launce
exceptionally 41 '3 %). In the caudal fin we find no
characteristic peculiarity worthy of mention, the least
depth of the tail in this species, too, being generally
somewhat more'' than half the length of the middle
rays of the caudal fin.

In the Sand-Launce the pectoral fins always extend
distinctly (for V4 — 1/8 of their otvn length) behind the

perpendicular from the beginning of the dorsal fin.
Their length is at least somewhat more than 9 % (ac-
cording to our measurements 9'1 — 10'3 %) of that of
the body, than 37 % (43 — 37'2 %) of the distance be-
tween the dorsal fin and the tip of the snout, or than
90 % (1 13*6 — 90' 1 %) of the postorbital length of the
head. They are also always longer than the lower jaw,
the length of which we have found to vary between
80 and 95 % of that of these fins.

The coloration is the same as that of the Sand-
Eel, but is often more bluish, olive-green with steel-
blue lustre, on the back. The young specimens in par-
ticular are finely punctated with round, dark brown
spots, one row above the lateral line, one row below
it, and a third row along the base of the dorsal fin
composed of somewhat larger pigmental spots. This
marking is, however, by no means peculiar to the Sand-
Launce. The upper part of the silvery white iris is in
most cases more shaded with black than is usually the
case in the Sand-Eel, where the whole iris is generally
silvery. In tins species too, the lower part of the sides
has a silvery lustre, but the belly is of a duller white.
The youngest specimens have a row of dark brown,
pigmental spots along each side of the base of the anal
fin. The snout is without the black spot on the sides,
or possesses merely a trace of this spot in the form of
scattered pigmental spots, larger and denser than on
the rest of the snout. The tip of the lower jaw is also
destitute of the black colour which we find there in
most Sand-Eels, at least in old specimens. The caudal
fin is darkened at the base above and below by brown
pigment between the rays. The peritoneum is silvery
white as in the Sand-Eel, but more densely punctated
with small, round dots of black pigment, sometimes so
dense that the preponderating colour of the membrane
becomes coal-black.

In order to ascertain the natural relations between
these three species of European Sand-Eels, we shall now
notice some points in the manner in which the most

a In specimens of the Sand-Eel between 135 and 289 mm.
to advance with fair regularity.

b From 40 to 36 % in the Sand-Launce;
c In Sand-Eels between 11 and 29 cm.
the percentage was
d Sometimes

significant characteristics appear in these species. In
general Ammodytes lanceolatus has a longer head, measur-
ing more than Vs °f the length of the body; but

long, this proportion rises from 59'2 to 60'2 %, the increase thus seeming
33 to 30’6 /. In a Sand-Launce 56 mm. long, however,

from 45 to 44 % in the Sand-Eel.
long this proportion sinks from

32-1.

perceptibly less, however, according to Kroyer.

AMM0DYTID7E.

577

this relative length decreases with increasing age, and
in this respect Am. lanceolatus thus retains a juvenile
character. Nor can the character be employed as a
specific distinction, for in a young specimen of the Sand-
Launce 56 mm. long, taken by Fries in Bohusktn, the
length of the head is nearly 21 % of that of the body.
In the same specimen the length of the lower jaw is
9‘3 % of that of the body, while in other cases the
boundary between the species in this respect lies at 9 %.
The length of the pectoral fins, where the line of di-
stinction is the same, but the direction of distinction
reversed — the percentage less in Am. Icmceolatus than
in Am. tobianus — in a specimen of Am. tobianus 20
cm. long is only 9'1 % of the length of the body, and
in a specimen of Am. lanceolatus 11 cm. long 9 % of
the same length. In this respect, too, the percentages
decrease with increasing age, and in this respect Am.
tobianus thus retains a juvenile character. In his de-
scription of Ammodytes americanus “ Storer says the
dorsal fin begins just at the end of the pectoral fin,
and that the length of the latter fin is only 1/3 of that
of the head. It was b}7 this statement that I was in-
duced to refer the Sand-Eels brought home by the Vega
Expedition from Pitlekaj, north-west of Behring Strait,
to Am. lancea, var. americancd, for in a specimen 77
mm. long the beginning of the dorsal fin lay at a
distance from the tip of the snout measuring 29‘3 % of
the length of the body, and extremely little in front
of the tip of the pectoral fin, though this fin measured
1 1 % of the length of the body. Later investigations,
however, especially those of Jordan and Gilbert, show
that Storer’s description and figure must have been
based on some exceptional specimen, for, unless this
were so, such a character could not have escaped ob-

servation. Still, this shows that even the characters
most important in a systematic respect are subject to
variation. According to Storer Am. americanus may
also attain a size of 3 dm., though from a specimen
which the Royal Museum has received through the
Smithsonian Institution from Woods Hole (Mass.), it
appears to be quite identical with Am. tobianus. The
other two American species that have been ranged be-
side Dekay’s Ammodytes americanus c, but which are
regarded by Jordan and Gilbert merely as varieties
of this species, show among their characters a variation
of the transverse dermal folds on the sides of the body
between 130 and 182, thus filling the gap between the
normal numbers in Am. tobianus and Am. lanceolatus ,
and showing how the form-differentiation may bring
about a resemblance to Am. lanceolatus, without altera-
tion in the structure of the mouth and without the
development of teeth on the head of the vomer. The
development of the said dermal folds, the absence of
which is one of the most important characters of Am.
cicerelus, in Am. tobianus is a character of growth ex-
tremely irregular in its appearance. Even in specimens
7 cm. long it is sometimes impossible to discover them.
Of the other character which should serve to distinguish
Am. cicerelus, the undulating margins of the dorsal and
anal fins, I have found at least a trace in a slight con-
cavity of the margin of the anal fin in a young Sand-
Launce. All this goes to show, not only that all the
species are extremely closely related — so closely that
we may well be tempted to regard them merely as
varieties of the same species, or as species at the be-
ginning of their differentiation from each other — but also
that this relationship has its origin in a form essentially
resembling Am. tobianus , or perhaps in this very species.

At least one of the species described above, the
Sand-Launce, must be regarded as circumpolar. The
Royal Museum possesses examples of this species from
Norwegian Firnnark, the Murman Coast, the White Sea,
North-East Siberia, Greenland, and Iceland. In Spits-
bergen, however, it is unknown. According to Machado
(quoted by Steindaciiner) the range of this species
extends southward to Cadiz. Day assumes that it may

occasionally wander into the Mediterranean; but he gives
no observation of its occurrence there. In the Baltic
the Sand-Launce penetrates at least to the island-belt
of Stockholm and, according to Mela, to the islands

o o

round Aland and Abo and into the Gulf of Finland
up to the island of Hogland. In the first locality Sun-
devall could not find it; but during the investigations
which I caused to be made this summer (August, 1890)

a Mem. Amer. Acad. Arts, Sc., n. ser., vol. VIII, p. 411, pi. XXXIII, fig. 2.
b Gt. Intern. Fisher. Exbib. London 1883, Swed. Spec. CataL, p. 170.
c Girard’s Ammodytes personatus and Cork’s Am. alascanus.

578

SCANDINAVIAN FISHES.

by Mr. A. Svensson, with a Sand-Eel net from Hal-
land, a quantity (several hundreds, if not thousands)
of Sand-Launces about 7 cm. long were taken and ob-
served at Ronnklippa off Runmaro. Lindstrom found
the species oft' Gothland. In the north and east of the
Baltic, however, it is considered at least rarer than the
Sand-Eel, the range of which extends, according to
Mela, in the Gulf of Bothnia to the neighbourhood of
Bjorneborg and in the Gulf of Finland to Cronstadt.
In the Atlantic, on the other hand, the Sand-Eel does
not go so far north, for it is unknown, according to
Collett, north of Trondhjem Fjord. On the English
coast it is common at certain spots, though here and
still more in the Channel on the coast of France the
Sand-Launce is the commoner species. Bonaparte in-
cludes the Sand-Eel among the fishes of Italy, and ac-
cording to Day it occurs, though extremely rarely, in
the Mediterranean; but neither Steindachner, Moreau,
nor Giglioli assigns it to this locality.

These two species, the most common Scandinavian
forms of the genus, seem to have exactly the same
habits, and in most places are found together. In ge-
neral the smaller species, the Sand-Launce, seems to
be the more plentiful. This is evidently the case in
Scania, where Sundevall found the Sand-Eel to form
hardly a tenth of the catch. Off the Danish islands
Kroyer considered the two species to be more evenly
distributed, but on the coast of Jutland the Sand-Launce
was said to be the more common. In the Great Belt
and Sam see Belt Wintrier found the catch to consist al-
most entirely of Sand-Eels. Among some hundred Sand-
Launces forwarded to the Royal Museum from Halmstad
in the month of August, there was only one specimen
of the Sand-Eel. On the coast of Bohuslan, too, ac-
cording to Malm, the Sand-Eel is rarer than the Sand-
Launce, which at suitable spots is common.

The Sand-Eel" generally haunts a sandy bottom, in
which it can bury itself. It works itself with wonder*
ful skill and rapidity into the sand, where it seems to
lie hid, at a depth of lV2 dm. or more, the greater part
of the day, coming up only at intervals. In winter it
lives in deeper water; but in spring, when the water
begins to grow somewhat warmer, both species ascend
together to sandy, shelving spots along the shore and
stay there all the summer. In Scania this takes place
at the end of May, and the fishery begins immediately

afterwards. During the course of October they again
return to deep water.

The most productive Sand-Eel fishery in the whole
of Sweden is carried on in the little inlet just north of
Simrishamn. So much fish is taken here, according to
Nilsson, that- the Rector of Gladsax receives as his tithe
of the fishery 6 barrels (989 litres) of dried Sand-Eels.
The town probably owes its origin to this fishery.
During the whole summer and until October the fishery
is pursued there daily, when not interrupted by storms,
with fine-meshed seines, which are shot at a depth of
3 fathoms and hauled up on shore. The fishing begins
soon after noon and continues until nearly sunset.
Sand-Eels are also especially plentiful at the following
places: south of Ahus, at the fishing-villages of Vik
and Baskemolla (north of Simrishamn) — where in 1880,
according to Mr. Lundberg, Inspector of Fisheries, the
catch was about 50 hectolitres, of a value of 302 crowns
(£16 10s.) — at Kaseberga (between Simrishamn and
Ystad), off Yst-ad, just east of the town and down to
the mouth of Kopinge River, and at the fishing-village
of Abekas (west of Ystad). At these spots some quantity
sometimes comes into the market, but in the rest of
Scania, as well as in Halland, Sand-Eels seem to be used
almost exclusively as bait for Cod. At the south-west
corner of Scania (Trelleborg, Skanor) these fishes are
quite unknown to the fishermen, according to Sunde-
vall; and in the Sound according to Sciiagerstrom,
both species are rare, though large quantities were taken
at the fishing- village of Raa in September, 1837. Further
up the Baltic, as even Linnaeus tells us, some Sand-
Eels are caught off Ottenby, on the south point of
Gland, and used as bait- for Cod. This fishery is carried
on with seines the cod-end (sac) of which is composed
of a sheet which is turned towards the sun and thus
attracts the Sand-Eels by its brightness. A similar
seine is used in ITalland. — At Simrishamn, it- is said,

o

the fishery is carried on in the afternoon. At Ahus
and Ystad the fishermen are out both in the morning
and in the evening, and in dull weather in the middle
of the day as well. It is only at these times that the
Sand-Eels move about in the water. In the island-belt
of Stockholm, in summer, Sundevall pretty often met
with small Sand-Eels, between ®/4 and 1 dm. long,
swimming about freely here and there among the is-
lands where there was no sand at- the bottom, and also

From Ihis point to the end of the description Sand-Eel is used as a general term for both species, except where otherwise stated. Tr.

SAND-EELS.

579

out' in deep water, but not far (5 — 14 metres) from the
surface, at the spots where Herring-fry were abundant.
As he did not see any large specimens at the same time,
he concluded that the young Sand-Eels lead a roving
life, while the older ones seem to be more tied to one spot.

On the shores of Norway the Sand-Eels are not
rare and are known by the name of Siil. When the
bottom is left dry by the ebb-tide, they bury themselves
there, and move about at high water. But they also
occur far out at sea in water of considerable depth,
■where during summer they are caught in large numbers
by several seabirds ( Mormon arcticus , Uria troile). This
can, of course, happen only when the fish is swimming
about- — During winter, when the Sand-Eel keeps to
deep water, we may in all probability assume that like
most other fishes at this season, it leads a still more
quiet life than in summer. At this season, too, it pro-
bably haunts a sandy bottom and lies buried in the
sand. Professor E. W. Areschoug informed Sundevall
that one winter a fisherman who was dragging to find
some Cod-lines that had been lost, took up a Sand-Eel
King (a large Ammodytes lanceolatus ) on a sandy bottom
at a great depth. It appears, however, that even at
this season the Sand-Eel occasionally moves about, from
the fact that during winter it is often found in Cod
caught in deep water.

The food of the Sand-Eel is composed of all kinds
of small marine animals, especially worms, which it is
believed by some to search for in the sand. It is not
for this purpose, however, that it burrows in the ground.
If is impossible for a fish buried in this manner
to seek or seize any prey. The Sand-Eel lies in the
sand to rest and to avoid its many different enemies:
Porpoises — which have been observed even to root up
the bottom to find their victims — Mackerel, Cod-fishes
— especially the Pollack, which has been seen in shoals
chasing the Sand-Eels up towards the surface to seize
them from below and devour them — Garpike, and
other tishes-of-prey, as well as seafowl. The principal
food of the Sand-Eel consists, however, of small fishes,
belonging even to its own genus. We have already
mentioned that the fry swim about where young Her-
rings are plentiful. The middle-sized Sand-Eels devour
their smaller congeners and other fry; while the largest

Sand-Eels seem to live almost exclusively on the middle-
sized ones.

The spaAvning-season of the smaller species, the
Sand-Launce, occurs in August. Sundevall Avas in-
formed, hoAvever, by Mr. Halck of Simrishamn, that
in 1856 the spaAvning of the Sand-Launce lasted at
least until the 15th of September. — Still the spent
fish remain in the shalloAvs together Avith the others.
At many spots in England and Ireland" the Sand-
Launce is caught between the tide-marks even in Avin-
ter, though then, after the spawning-season, it is “so thin
as not to be sought after generally for food”. It is a
remarkable circumstance that the spaAvning does not
begin until the fish has been three months at the spawning-
place. — Less is known of the spawning-season of the
larger species, the Sand-Eel. Nilsson received informa-
tion from Ystad to the effect that the Sand-Eel spawns
there in April; but it is hardly probable that the species
ascends to the shallows so early. On the 12th of Octo-
ber, 1892, our Museum received through Mr. C. A. Lind-
rotii a neAvly caught Sand-Eel 25 cm. long, Avith the
testes just beginning to SAvell, from Stenvik (Ljusteron)
in the island-belt of Stockholm. Malm gives an ob-
servation of a female 28 cm. long, that had fully de-
veloped roe on the 5th of June; but he does not state
Avhether the roe was quite ready to be deposited. At
St. Ives Day found the generative organs of the Sand-
Eel and the Sand-Launce to be equally developed in
August. In Scandinavia both species ascend into shal-
low Avater at the same time, in the month of May, and
it seems most probable that they also spaAvn at about
the same time, the larger perhaps, as is common among
fishes (in the Herring, for example), before the smaller.

Of the growth of the fry Sundevall remarks that
the young specimens, 75 or at most 100 mm. long,
Avhich in summer (from July to September, or perhaps
still longer) rove far and Avide in search of prey, pro-
bably belong to the fry of the previous year, and that
the ordinary specimens 125 mm. or more in length
seem to be a year older. At the end of July, 1880,
off Groto among the Lofoden Islands, Collett also
distinguished between three different generations of the
Sand-Launce, 75, 125, and 170 mm. long and, in his
opinion, respectively 2, 3, and 4 years old6. Whether

a See for example Thompson, Nat. Hist. Irel., vol. IV, p. 238.

b We arrived at quite different results in a consignment of Sand-Launces from Sondrum, off Halmstad, taken at the beginning of
August. Among these specimens a few measured between 50 and 55 mm. in length, while the others, we may almost say, were of all pos-
sible sizes between 95 and 125 mm.

580

SCANDINAVIAN FISHES.

the Sand-Eel (Am. lanceolatus) has reached maturity
at the size last mentioned, is as yet an open question.
It grows, as we have mentioned above, to a length of
80 cm. or more; and both males and females attain
this size. These largest specimens of the Sand-Eel are
known in Scania by the name of Kimgar (Kings). At

o

Alius they are also called Groningar (Greenfish), a
name which is sometimes applied to all specimens of
this species. Some fishermen know that there are two
kinds of Sand-Eels, and that the Kings belong only to
one of them. Nilsson has adopted the names of hvit-
tobis (White Sand-Eel) and blatobis (Blue Sand-Eel)
from Blekinge.

The Kings are rare, probably from a cause easy
of comprehension. It is no doubt difficult for many
specimens to escape for some years the tackle con-
stantly set for them, especially the seine. They are,
therefore, most numerous in proportion to the entire
catch at spots where the two species are not plentiful
enough to make the use of the seine remunerative.
That some few of them do escape, in spite of all, seems
probably to depend on the fact that the older speci-
mens bury themselves in the sand and lead a much
more sluggish life than the young ones, being thus
much less exposed to the risk of being caught. They
are said to be taken only towards evening. The fishery
for Sand-Eels is indeed remarkable, for it shows the
extraordinary hardiness of these species. Off Simris-
hamn Sand-Eels are still taken in quantities only
slightly, if at all, less than in former times, although
a war of extermination has been pursued against them
yearly for several centuries. This war has been car-
ried on with fine-meshed seines long before, during,
and after the spawning-season; and would thus seem
to have been enough entirely to extirpate the spe-
cies in a few years. But it is just this hardiness that
renders the history of the species so much more de-
serving of careful study. The fine and extremely nu-

merous eggs, and perhaps the roving life of the young
specimens, may perhaps compensate to some extent the
numbers destroyed, if only a few large females be
yearly permitted to deposit their roe. Still it is not
unusual to hear the fisherman complain of the decline
of this fishery. Thus we learn now (1890) from Sond-
rum (off Halmstad) that “the Sand-Eel has become
rarer in this neighbourhood than it used to be. For-
merly it was sold and eaten in quantities. Now the
supply is scarcely enough for bait.”

The Sand-Eel is eaten fresh, boiled, or fried, and
is regarded by many as a delicacy. Others assert that
it is too lean, and others again find the smell of the
flesh repulsive. At Simrishamn a great portion of the
catch is dried and preserved for winter use. There is
no distinguishable difference in flavour between the two
species. These fishes are, however, very important to
man in another respect as well. The large shoals in
which they live, entice larger and more valuable fishes
to the fishing-grounds. Mr. Lundberg", Inspector of
Fisheries, remarks that the Sand-Eel is important as
food for the Salmon, which at shelving spots along the
Scanian coast comes close in shore chiefly to hunt Sand-
Eels, and which during certain years thus becomes the
object of a highly lucrative seine-fishery. The Sand-
Eel is also considered excellent bait for Cod and other
large fishes-of-prey, and is used for this purpose wher-
ever it can be procured.

We have already described the seines used for
Sand-Eels. In tidal waters, as for instance on the
coasts of the North Sea, in England, Ireland, and France,
persons may be seen, when the tide is out, scratching
up Sand-Eels with spades and other implements adapted
to this purpose. Similar implements are used in Swe-
den to secure the Sand-Eels that have buried them-
selves in the sand after they have been drawn ashore.

(SuNDEVALL, SmITT.)

Lundberg, Meddelanden rorande Sveriges fiskerier, haft. 2, p. 151.

ONOMORPHS.

581

Fami. MACRUR1DJE.

Body clavate ( more or less tadpole-like), with more or less straight lack and concave ( arcuate ) caudo-ventral
margin , compressed, with elongated, more or less whip-like tail; covered with thin but spiny, carinated or striated
scales. No distinct caudal fin ( the vertical fins confluent behind), but a distinct anterior dorsal fin or at
least a trace thereof in the elongation of the first dorsal rays. Teeth on the intermaxillary bones and in the lower
jaw , but the vomer , palatine bones, and tongue toothless; mouth highly protrusile. Gill-openings large; brancliio-
stegal membranes more or less united to each other, but free from the isthmus. Branchiostegal rays 6 or 7.
Pseudob rand vice wanting. Air-bladder present. Pgloric appendages well-developed.

Here we have a family consisting almost exclusively
of deep-sea fishes"; and besides the characters given
above we observe in the first place those peculiarities
which generally belong to such fishes, especially in the
loose structure of the head, with its ample space for
muciferous canals. Two in particular of the cephalic
bones, the nasal bones, are greatly enlarged, and form
the framework of the highly variable form of the snout in
the Grenadiers, as these fishes are called in Italy6. The
suborbital bones, which form the lower margin of the
orbit, also afford an instance of a systematic peculiarity
rare among the Anacanthini. They sometimes develop a
connexion with the opercular apparatus similar to that
we have seen above in the Gottomorphi and Cyclopteridce.
Another systematic exception might also be used as an
argument in favour of the inclusion of these fishes
among the Acanthopterygians. The first large ray in the
anterior dorsal fin (really the second ray in the fin, the
first ray being extremely short and rudimentary) is a
true spinous ray, without joints. These two points of
resemblance to the Acanthopterygians might well induce
us to range the family Macruridce beside the Gurnards
and the Agonidce, according to SwainsonV proposal.
However, if we trace the form-series back to the least
differentiated Grenadier types'*, we find in the form of
the head the most distinct approximation to the Phycis
group, within which the relationship to the Grenadier-
fishes is also expressed in the form of the snout in the

American Haloporphyrus ( Antimora ) violae. In the cra-
nium, too, we find an evident sign of the relationship
to the Codfishes in the advanced development of the
styloid bone (os opisthoticum) , and the lobate process
erected in an upward and backward direction on the
hind part of the intermaxillary bones is also as well-
developed as in most of the Codfishes. Thus, the most
natural place of the Grenadier-fishes in the system is
beside the Codfishes, as a remarkable variation of the
Anacanthine type, with a characteristic common among
deep-sea fishes in the reversion to or retention of the
original form of the tail without separate caudal fin7.
The primitive (palauchthyic) appearance of the Grenadier-
fishes also depends on the covering of scales. In one
species we find the scales replaced by projecting spines
scattered in the skin. Only in few species and in the
fry of other species do we meet with thin, fully typical
cycloid scales. In the rest of the family the scales are
generally furnished throughout their free surface with
spines or raised carinae, which give the fish a Ganoid
appearance. Bonaparte, therefore, referred the Grena-
dier-fishes to the order Ganoidei, when in 1837 he
established a special family 9, Macrouridce, for these forms.

A juvenile form, Krohnius, is described by Cocco7'
and Emery' as reminding us, by the long rays of the
ventral fins and the position of the first dorsal fin far
forward on the head, of the larvae of the Trachypteroids ,
but as most closely resembling, in the form of the

“ One species ( M acruronus novce-zelandias) is said to live in shallower water.

6 See Risso, Ichthyologie de Nice, p. 201, where the name is supposed to have originated from the resemblance between a soldier’s
helmet and the snout of Macrurus coelorhynclius. The same name occurs in Cuvier ( R'egne Animal, tome II, ed. 1, p. 217; ed. 2, p. 336)
and in Brown-Goode {Fisher., Fish. Industr. U. S., Sect. I, p. 244).

c Nat. Hist. Fish., Amph., Rept., vol. II, pp. 179 and 261.

d See for example Macrurus ( Nematonurus ) longifilis , Gunther, Deep Sea Fishes, Chall. Exped., p. 151, pi. XXXV.

e Brown-Goode, The Fisheries and Fishery Industries of the United States, Sect. I, pi. 64; GOnther, 1. c., p. 94, pi. XV.

I A separate caudal fin may, however, appear to be present. This is due to cicatrization following upon the breaking off of the tail
in an adult Grenadier-fish. Cf. Nilsson, Slcand. Fauna, Fislcarne, p. 606.

g Trans. Linn. Soc. London, vol. XVIII (1838 — 1841), p. 295.

h Giorn. Gabin. Letter. Messina, anno III, tomo V, fasc. XXV (1844), p. 21; reprinted in 11 Naturalista Siciliano, anno VII, No. 4,
l:o Gennajo 1888, p. 101.

1 Mem. d. R. Acad. d. Lincei, scr. 3, vol. Ill, p. 395, figs. 7 and 8.

582

SCANDINAVIAN FISHES.

body, Gunther’s Macrurus eras sleeps a from the deep-
sea fauna of Australia.

In 1884 Professor Leche took a specimen of a Krohnius- form at
the surface off Messina. This specimen (fig. 138) has advanced so
far in development that we need not hesitate in referring it at least
to the genus Macrurus. Including the very fine tip of the tail the
specimen measures 92 mm., the forepart of the body being 12 mm.
long, 9 mm. deep, 6 mm. broad, clumsy, and almost spherical, while
behind this point the strongly compressed tail begins with a depth of
4 ■’A nun. and very gradually tapers to a filamentous, narrow but com-
pressed, and membranous appendage. This appendage, which is I8V2
mm. long, forms a slight break at its insertion in the tip of the tail
and advances straight back, marked with dark reddish brown spots of
pigment, small, but dense, and set in two rows at the base of the
appendage, in one row, more scattered, and larger (finally occupying
the entire breadth of the appendage) towards its tip, which is, how-
ever, colourless. The head does not measure even half of the bulky
forepart, though its length, measured as usual from the tip of the
snout to the hind margin of the gill-cover (obliquely against the lon-
gitudinal axis of the body), is 6'2/3 mm. From the same cause the
length of the snout becomes fairly great, measuring 1 */5 mm., although
the round eyes, with a longitudinal diameter of 2 mm., and set at a
distance of about 2 mm. from each other, evidently lie far forward.
The postorbital part occupies about half the length of the head, or 3Y5

Fig. 138. Krohnius f larnentosus, natural size. From Messina, Feb., 1884,
V. Leche. Original in the possession of the High School of Stockholm.

mm. The mouth is both terminal and lateral, with sharply ascending
cleft. The maxillary bones end almost below the middle of the eyes.

Below the chin there hangs a barbel about equal in length to the
diameter of the eye. The nostrils are set on each side at the upper
anterior corner of the eye, in a common opening, divided into two
parts by a thin partition-wall. The first dorsal fin begins at a distance
of 7 mm. from the tip of the snout, or a little behind the head. It
is high — the longest ray measuring about 7 1('2 mm. — -and pointed,
almost falciform ; but the anterior rays (except the first, which is
rudimentary) are elongated into filaments. The fin contains one rudi-
mentary ray and nine perfect, but simple rays. The length of its
base is 21/,2 mm., or somewhat more than the longitudinal diameter of
the eyes. Behind this fin the dorsal edge still retains a low dermal
flap, a remnant of the embryonic vertical fin, and distinguishable not
quite to the end of the bulky forepart. At this point the flap dis-
appears, but soon returns again, though at first scarcely distinguishable,
at about the end of the swollen forepart of the body, now with true
rays and in the form of a second dorsal fin, which is remarkably low
and follows the dorsal edge back to the base of the caudal filament.
The anal fin begins at a distance of 11 mm. from the tip of
the snout, on the posterior part of that which we have termed the

swollen forepart of the body. Its base thus follows the ventral margin
backwards and upwards, then passing in a rounded obtuse angle to
the ventral margin of the caudal part, and following the latter to its
termination. The rays of the anal fin, which, like those of the dorsal
fins, are all simple, first increase in length uniformly but sharply, the
length of the rays at the beginning of the tail being about equal to
the depth of the latter. Thus the margin of the anal fin is fairly
straight, in spite of the sinus in the margin of the body at this spot.
From this point the rays gradually decrease in length, until at the
beginning of the posterior half of the tail their length increases some-
what, the height of the anal fin being here greater than the depth
of the tail; but towards the tip of the tail the length of the rays
again decreases. At the base of each ray of the anal fin we find a
dark-brown, pigmental spot. The pectoral fins are perhaps the most
characteristic peculiarities of this larva. They are almost semicircular
disks, for the greater part free, attached to the shoulder-girdle only
at the top by a narrow shaft, which is inserted on about a level with
the middle of the eye. The outer part of these disks is extremely
thin and membranous, but with rudiments of the future pectoral rays ;
the inner part is somewhat thicker, in the specimen preserved in spi-
rits opaque, and is of an almost regular, but flattened kidney-shape, the
upper corner of which, situated about one-third of the way up the
pectoral fins, forms the point of origin for the said shaft. If we com-
pare this form of the pectoral fins with their arrangement in certain
Codfishes, where the internal structure is known* 6, it appears extremely
probable that the shaft corresponds to the rudiments of the proximal
parts of the shoulder-blade and the coracoid bone, while the distal parts
of these bones and the basal bones of the pectoral rays are developed
in the kidney-shaped part of the base of the pectoral disk. Shafted
(lobate) pectoral fins are indeed by no means rare in the fry of other
species0; but a shaft so long and so narrow as in this larva — remind-
ing us of the elongation of the basal bones in Lophius — is unparal-
leled within our experience. The ventral fins are set vertically below
the insertion of the pectoral fins, somewhat in front of the middle
point between the tip of the snout and the anal fin — the foremost
point of their insertion lies at a distance of 6 mm. from the beginning
of the anal fin — and are remarkable for the great elongation of the
six middle rays. The first ray is not short, its length being very
nearly as great as that of the head; but the next 6 rays attain a length
of as much as 30 mm., are pigmented in the outer part in the same
way as the caudal filament, and are also flattened throughout their
length like this filament. The innermost two rays are short, the inner-
most ray the shortest of all. The anal aperture lies about half-way
between the insertions of the ventral fins and the beginning of the
anal fin; it has a pointed anal papilla behind it. Half-way between
the ventral fins and the isthmus we find a round depression in the
ventral wall, coasted in front by a semicircular, membranous, dermal
swelling, the two ends of which are continued each by a dermal ridge
which disappears behind. This structure evidently bears the appearance
of having served as an adhesive apparatus by means of which the larva
has been enabled to attach itself to floating objects, and is probably
of importance as an explanation of the bare (scaleless) spot which in
adult examples of some species of the genus occurs on this part of
the body. On the upper portion of the abdominal sides the scales have
apparently begun to develop; but the rest of the body is naked.

It is difficult as yet with certainty to determine the species of
this larva. Among the four Mediterranean species of the genus given
by Giglioli d , it can hardly be referred to any other than Macrurus
Icevis. This opinion is borne out both by the number of rays in the

“ Deep Sea Fish., Chall. Exp., p. 143, pi. XXXVII.

6 See for example Emery, Fierasfer in Fauna und Flora des Golfes von Neapel, II Monogr., p. 29, taf. IV, fig. 40.
c Cf. above, fig. 77, p. 311.

d Espos. Intern, di Pesca, Berlino 1880, Sez. Ital. Cat., p. 98.

GR EN A DIER-FI S H K S .

583

first dorsal (10) and the ventral fins (9) and by the form of the
adhesive ventral disk. The specimen would thus seem to belong to
a species that also occurs within the limits of the Scandinavian fauna.
But Gunther" has described a Mediterranean fish 'that in another publi-
cation by Giglioli'-'' had been named Macrurus ( Hymenocephalus ,
Mystaconurus ) italicus, and to this species, which is distinguished
from Macrurus Icevis by small, villiform teeth in the lower as well
as in the upper jaw and by larger (fewer) scales, our larva may be
more properly referred. The most interesting points in this larva,
however, are the tadpole-like form of the body, a form which is per-
sistent in Macrurus crassieeps, the short snout with the mouth set
in the position normal among the Anacanthini , the shafted pectoral
fins, a trace of the oldest period of the piscine type — though con-
siderably altered by the broad form of the pectoral disks — , the
evanescent adhesive disk on the belly, and the long appendages to
the rays of the ventral fins and to the tip of the tail. These ap-

pendages certainly play their part in assisting the larva to support
itself in the water; but probably serve also to give it a protective
resemblance to the stinging Medusa; c.

Some of the members of this family have long-
been known, at least since the times of Strom d and
Fabricius6; but the great wealth of forms that represent
it in the abysses of the ocean, have first been discovered
and described in recent times by Gunther7 and Vail-
lant6', Brown-Goode and Bean7'. According to the first-
mentioned writer the family contains at least 46 spe-
cies, which he distributes among 4 genera. Only one
of these genera is represented in the Scandinavian fauna.

Genus MACRURUS.

First branchial arch united above and below on the outside by a membrane to the inside of the opercular appa-
ratus, leaving a foramen at the middle which is much smaller than the other gill-slits , of which even the last is
complete. Gill-rakers on the first branchial arch tubercular. Branchiostegal membranes united to each other. Chin

furnished with a barbel. Dorsal fins well-separated from each other.

Within this genus, which received the name it now-
bears of Bloch', Gunther, the most eminent among
those writers who have revised the arrangement of the
genus, in his last treatment of the question has united
several genera which he formerly regarded as distinct,
but which he has now reduced to the rank of sub-
genera. “The dredge of the Challenger,” he writes7,
“secured more than one hundred and forty examples,
referable to thirty species, and proved that this type
of fishes is not only one of the most widely spread in
the depths of all oceans, but also extremely abundant
with regard to species and individuals. These materials
afforded the further evidence that the characters on
which I had relied for the generic groups of Macrurus ,
Coryphcenoides, and Mcdacocephalus, did not possess
the taxonomic value assigned to them, with the ex-
ception of the modifications of the dentition, which,
however, were capable of more precise definition.

“With regard to the form of the snout arid posi-
tion of the mouth, there exists every gradation, from
the most specialized types, such as Macrurus japonicus
and Macrurus parallelus , to Macrurus longifilis, which
may be regarded as representing the original type
whence the others were derived. Its head is com-
pressed, well proportioned, formed by firm bones, the
superficial of which enclose a muciferous system not
more enlarged than we find it in many surface fishes;
its snout is not more tumid or projecting than in the
majority of surface Gadoids, and the wide mouth ter-
minal and lateral. As the muciferous cavities increase
in width, the bones are expanded into thin lamellte
and lose in firmness, those of the infraorbital ring cover
more or less the side of the head, extend backwards
to the angle of the praeoperculum, and push the latter
backwards. The snout k becomes the receptacle of large
or even enormously enlarged cavities, supported by

a Deep Sea Fish , Challeng. Exped., p. 140.

b Giglioli e Issel, Pelagos, p. 228, c. fig. (without description) — cited from Gunther, 1. c. To this publication we have not had access.
c Cf. above, p. 312, note b.
d Sondmors Beskrivelse (1762), vol. I, p. 267.
e Fauna Groenlandica (1780).
f Deep Sea Fishes , Chall. Exped.

g Exped. Scicnt. du Travailleur et du Talisman , Poissons.

h Report on the Fishes, Dredg. 1880 U. S. Survey Steamer “Blake”, Bull. Mus. Comp. Zool., Harv. Coll., vol. X, No. 5.

1 Naturg. Ausl. Fiscli., Tli. 2, p. 150; from ya/pog, long and olyci tail. The oldest generic name is Coryphcenoides (Gunnerus,
1765), but was formed contrary to current rules and suggested for one of the Scandinavian species “until it please the great naturalist
Linnaeus to give the species its proper name.”

j Deep Sea Fish., Chall. Exped., p. 122.

k Chiefly by the extension of the nasal bones, as Reinhardt has already remarked.

Scandinavian Fishes.

74

584

SCANDINAVIAN FISHES.

thin osseous ridges, and projects more or less beyond
the month, which is forced downwards to the lower
surface of the head, like that of a shark. Thus, great
as the dissimilarity is between the extreme forms of
the snout in the species of Macrurus , there is no funda-
mental difference in structure; they merely represent
different degrees of the same line of modification.

“With regard to the scales, there is also every
gradation from the small-scaled Malacocephali to the
large-scaled Macrurus longibarbis. In very young spe-
cimens of all species the scales formed at first are al-
ways thin, without any armature, in fact cycloid. Spines
appear only after some time, generally in the median
line of the scale, simply and not in series; scales with
fully developed armature are generally not found in
specimens under 8 inches in length. In some species
which normally possess strongly spiniferous scales,
individuals may occur (especially such whose skin is
wanting in pigment), in which the spines are much
more feeble and scarcely visible. And finally, there
are species in which the cycloid structure of the scales
remains normally persistent. Thus, neither the size
nor the structure of the scales can be safely used as
a generic character.”

On the other hand, Gunther fully recognises the
validity of a character first suggested for this purpose
by Lutken °, and derived from the presence or absence
of the spines (set in an upward (backward) direction
on the front surface of the ray) that arm the second
(apparently the first) ray of the dorsal fin. Even this
character is subject to changes of growth, for, where
these spines are present, in old specimens they may
become indistinct (tubercular); but they never dis-
appear altogether.

The species hitherto discovered within the limits
of the Scandinavian fauna may be distinguished, ac-
cording to these opinions, in the following manner:

A: Jaw-teeth at least in front cardi-
fonn (or velvet-like); scales middle-
sized or large.

1 : Snout pointed; cheeks carinated :

a: Second ray of the first dor-
sal fin smooth (unarmed). .. J Macrurus (Coelorhynchus)

\ coelorhynchus.

b: Second ray of the first dor-
sal fiu spiny J Macriirus (Macrurus)

2: Snout blunt; cheeks almost^ Fabricii.

even j Macrurus ( Coryphamoides )

B: Jaws furnished with canine teeth ^ rupestris.

set in two rows in the upper jaw,

in one row in the lower I Macrurus ( Malaco -

I ceplialus) Ice vis.

It sometimes happens, however, that the character
derived from the structure and arrangement of the jaw-
teeth, employed in this manner, firstly brings together
forms extremely dissimilar in other respects6, and
secondly is very difficult of employment, for in young
specimens of the Coryphamoid group, even when they
are 12 cm. long, it is hardly possibly to discover more
than two rows of teeth in the upper jaw and one row
in the lower. Again, several species of Gunthers sub-
genera Chalinurus and Optonurus have true canine teeth
in the outermost row of the card in the upper jaw and
only one row of teeth in the lower jaAv. The folloAving
classification appears easier and, therefore, more trust-
worthy.

A : Least breadth of the interorbital
space less than the length of the
snout, than J/4 of the length of
the head, or than 3/5 of the greatest
breadth of the head at the perpen-
dicular from the centre of the eye:
a: Least breadth of the interorbital
space more than or equal to 3/-
of the length of the lower jaw,
which is less than 40 % of the

length of the head Macrurus coelorhynchus.

b: Least breadth of the interorbital
space less than 1/2 the length
of the lower jaw, which is more
than 40 % (44 /?) of the length

of the head Macrurus Fabricii.

B: Least breadth of the interorbital
space greater than the length of the
snout, and greater than or equal to 1/,3
of the length of the head or 7/io °f
its greatest breadth at the perpen-
dicular from the centre of the eye.
a: Least breadth of the interorbital
space more than 4 A of the length

of the lower jaw... Macrurus rupestris.

b: Least breadth of the interorbital
space less than or equal to 3/3
of the length of the lower jaAv Macrurus hvvis.

a Vid. Meddel. Naturh. For. Kbhvn 1872. p. 4.

6 Cf., for example, Macrurus Icevis and Macrurus macrocliir.

GRENADIER-FISHES.

585

MACRURUS COELORHYNCHUS.

Fig. 139.

Snout pointed, flattened, of a broad triangular shape, with sharp sides, which are continued by the carina of the
suborbital ring ( the cheeks ) until they join the angle of the preoperculum ; length of the snout greater than the
least breadth of the interorbital space, which is less than 1/i (— 22*1 — 23*6 % in the specimens examined by us,
which are between 18 and 25 cm. long) of the length of the head, than 4/5 (= 66'4 — 76*8%) of the longitudinal
diameter of the orbits, than '2/3 (= 66*3 — 65 %) of the postorbital length of the head, than 1/i (= 19*4 — 20*4 %)
of the distance between the first dorsal fin and the tip of the snout, than 3/5 (= 56*9 — 54* 1 %) of the breadth
across the cheeks at the centre of the eyes, or than (= 61*1 — 64 %) of the length of the lower jaw, which is
less than or equal to 3/5 (= 35*7 — 39*1 %) of the length of the head, or about 1/3 (= 31*9 — 33*6 %) of the di-
stance between the first dorsal fin and the tip of the snout, but greater than (in young specimens equal to) the
postorbital length of the head. Mouth set on the under surface of the snout, its breadth at the corners less than
2/ 3 (= 52*9 — 60*4 %) of the breadth across the cheeks at the centre of the eyes. Length of the head about 23 — 22 %
(or less) of that of the body, 90 — 85 % of the distance between the first dorsal fin and the tip of the snout,
58 V2 — 57 % of the distance of the second dorsal tin from that same tip, and 7 1 x/2 — 67 % of the distance be-
tween the anal fin and the tip of the snout. Depth of the body at the beginning of the first dorsal fin about
14 — I2V2 % °f the length of the body, at the beginning of the second dorsal fin about 10 % of the same. Pectoral
fins lobate (with brachiate base). Length of the base of the first dorsal fin about 1/s (= 57 — 51 %) of the
distance between the two dorsal fins, which is about equal to the breadth across the cheeks at the centre of
the eyes. 4 or 5 scales in a row from the end of the first dorsal fin to the lateral line. A narrow, oblong,
bare spot in the median line of the belly between the ventral fins. Scales densely set with spines throughout
the free surface , but without ccirince. Second ray of the first dorsal fin ( spinous ray) smooth; its height about
equal to the depth of the body at its base or somewhat less. Length of the barbel under the chin about 1/3

of that of the lower jaw. Jaw-teeth cardiform.

Fig. 139. Macrurus coelorhynchus, natural size. From Messina, through Professor V. Leche. The property of the Museum of

the High School of Stockholm.

R. br. 6; Dv (1)+V7; D.,. 80 — 90; A. 83—98; P. (1) + V,8;
V. '/6; Lin. squ. transu. 19 — 21.

Syn. Lepidoleprus Coelorhynchus. Risso, Ichth. Nice, p. 200, tab.
VII, fig. 22; Ear. Mer., tom. Ill, p. 244; Bonap. {Jtfacrourus'),

586

SCANDINAVIAN FISHES.

Icon. Fn. Ital ., tom. Ill (Pesci), tab. 123, fig. 1; Gthr
( Macrurus ), Cat. Brit. Mus., Fish., vol. IV, p. 392 ; Coll.
Forb. Vid. Selsk. Chrnia 1874, Tillasgsh., p. 129; Mor.,
Hist. Nat. Poiss. Fr., tom. Ill, p. 278; Lillj., Sv., Norg.
Fisk., vol. II, p. 253; Gthr, Deep Sea Fish., Chall. Exped.,

p. 128.

Macrurus atlanticus , Lowe, Proc. Zool. Soc. Lond. 1839, p.
88; Gthr, Cat., 1. c., p. 392.

This species is one of those in which the Macruroicl
type appears in its most developed stage, a point which
we observe principally in the structure and shape of
the head. The species attains a length of 3V2 dm.
The coloration, according to Risso, is grayish with a
reddish violet lustre on the back, silvery with mother-
of-pearl and golden lustre on the sides, bluish black
or black on the belly. The ventral tins, the branchio-
stegal membranes, the branchial cavities, the axil of
the pectoral, and the margins of the vertical tins are
also bluish black or black. The inside of the mouth
and the tongue, on the other hand, are light (whitish
yellow). The snout is transparent as if of cartilage.

To the above diagnosis we shall add merely that
the anterior nostril — the nostrils are set on each side
close to each other, as in most of the members of this

genus — is round, the posterior kidney-shaped, on ac-
count of the semicircular dermal flap that overlaps it
in front.

Macrurus coelorhynchus really belongs to the Medi-
terranean and the neighbouring part of the Atlantic ",
living in from 200 to 300 fathoms of water. In May,
according to Risso, the females repair to rocky shores
to deposit their eggs, which number about 3,000. The
adult specimens are said to feed on “worms and zoo-
phytes.” The dentition tells us distinctly that they
cannot live on large or hard-shelled animals, but that
their food probably consists chiefly of worms and thin-
shelled crustaceans.

Only on one single occasion has this species been
met with in Scandinavia. In February, 1842, the elder
Saks found a specimen that had probably measured
about 29 cm., in the stomach of a Cod that had been
taken off Herlovaer, north of Bergen. This specimen
was so well preserved — only the tip of the tail and
a portion of the scales on one side of the body were
wanting — that Collett justly concluded that the Cod
must have devoured its victim only a short time before
its own capture.

“ Still, it comes extremely near — if it be not identical with — Macrurus caribbceus, which Brown-Goode and Bean have described
(Proc. U. S: Nat. Mus., vol. VIII (1885), p. 594) from a depth of 210 fathoms in the north of the Gulf of Mexico, and which apparently
differs from M. coelorhynchus chiefly in the somewhat smaller breadth of the interorbital space and the longer base of the first dorsal fin.

GRENADIER-FISHES.

587

M ACRURUS FABRIC II.

Fig. 140.

Snout pointed and deep , pyramidal , with the sharp sides continued after a break by the carince of the cheeks
until they join the angle of the preoperculum: length of the snout greater than the least breadth of the inter-
orbital space , ivhich is less than 1/i (20*8 — 17*4 % in the specimens examined by us, which are between about
72 and 78 cm. long) of the length of the head , than 3/- (59*6 — 49*1 %) of the longitudinal diameter of the orbits ,
than ®/5 (54 — 42 %) of the post-orbital length of the head , than 1/8 (17*2 — 15*2 %) of the distance between the

first dorsal fin and the tip of the snout, than 7- (54 %) of the breadth across the cheeks at the centre of the

eyes , or than V2 (47*2 — 37*6 %) of the length of the lower jaw , which is more than */5 (44*1 — 46*4 %) of the
length of the head , than 1/3 (36*3 — 40*5 %) of the distance between the first dorsal fin and the tip of the snout,
and greater than the postorbital length of the head. Mouth set on the under surface of the snout, but its breadth
at the corners more than 2/3 (81 — 85 %) of the breadth across the cheeks at the centre of the eyes. Length of
the head about 23 — 22 % (or less") of that of the body, 90 — 86 % of the distance of the first dorsal fin from
the tip of the snout, about 2/3 (61 — 64*3 %) of that distance of the second dorsal fin, and rather more than 1/g
(56 — 52 %) of the distance between the anal fin and the tip of the snout. Depth of the body at the beginning
of the first dorsal fin about V5 — 7g °f the length of the body, at the beginning of the anal fin about 1/1 of the
same. Base of the pectoral fins only slightly brachiate. Length of the base of the first dorsal fin greater than the
distance between the two dorsal fins, which is considerably less than the breadth across the cheeks at the centre
of the eyes. 5 or 6 scales in a row from the end- of the first dorsal fin to the lateral line. No bare spot (ex-
cept the anal aperture) in the median line of the belly. Scales carinated and spiny. Second- ray of the first

dorsal- fin spiny, or ( in old, specimens ) at least granulated, on the front- surface; its length less than 2/3 of the

depth of the body at its base. Length of the barbel under the chin about ’/3 of that of the lower jaw.

Jaw-teeth car diform.

R. bv. 6; Dy (1) + Y[;D2. 105 — 124; A. 1 13— 148 ; P. 18— 20 ;
V. 1 Lin. squ. tr. 26 — 27 (ad pinn. ventr.).

a According to Collett 21'3 — 1 8 ' 6 % in specimens between 538 and 935 nun. long. Perhaps the rule is that the head is smaller
in the females than in the males.

S>/n. En egen sorte af ji.sk, liig en Brasme , Egede, Grant. Ntje
Per lust rati on, p. 51; Ingminniset , Cranz, Hist. Grunt, (ed.
suec.) pt. I, p. 139.

588

SCANDINAVIAN FISHES.

Coryplicena rupestris , Fabr., Fn. Groenl., p. 154 (exch synon.);
Br„ (Macrounis ; p. p.), Naturg. Ansi. Fisch., pt. II, p. 152,
tab. CLXXVII; Bl., Schn. ( Macrurus ), Ichthyol., p. 103, tab.
26; Reinh., D. Vid. Selsk. Naturv., Math. Afh., vol. V (1832),
p. XVIII; ibid., vol. VII, p. 129; Nilss , Skcind. Fn., Fisk.,
p. 604; Gthr, Cat. Brit. Mus., Fish., vol. IV, p. 390; Esm.,
Forh. Naturf. M. Christ. 1868, p. 527; Ltkn, Vid. Meddel.
Naturhist. For. Kbhvn 1872, p. 4; Br. Goode, Fisher., Fisher.
Industr. U. S ., Sect. I, p. 244, tab. 66.

Macrourus Fabricii, Sundev., Vet.-Akad. Handl. 1840, p. 6;
Mgen, Ofvers. Vet. Akad. Forh. 1867, p. 263; Coll. Forh.
Vid. Selsk. Christ. 1874, Tilleegsh., p. 128; Goode, Bean,
Bull. Essex Inst., vol. XI (1879), p. 7; Jord., Gilb., Bull.
U. S. Nat. Mus., No. 16, p. 811; Coll., N. Mag. Naturv.,
vol. 29 (1884), p. 95; Lillj., Sv., Norg. Fisk., vol. 2, p.
242; Gthr, Deep Sea Fish., Chall. Exped., p. 130.

Macrurus Fabricii, which attains a length of at
least nearly a metre (9x/3 dm.), is only slightly less
typical of the Macruroids than the preceding species,
though the snout is shorter and the mouth broader.
In this species, too, the snout ends in a hard, wartlike
protuberance, but the horizontal lateral carinae are more
concavely curved; and the front exterior corners of the
nasal bones are more distinctly set oft’ from the be-
ginning of the suborbital carina on each side. Besides,
the snout is not so depressed. The carina? on the head
are the same as in the preceding species: one in the
middle line of the upper side of the snout, ending in
a line with the anterior margin of the orbits; one on
each side from the anterior outer corner of the nasal
bones up to the orbital margin, and together with the
latter and the anterior suborbital bone forming a tri-
angular wall round the nasal cavity; one on each side
of the forehead, which is only slightly concave, and
along the side of the occiput, back to the beginning of
the lateral line; one on each side along the temples,
from the upper part of the hind margin of the orbit
to the upper corner of the gill-opening; and lastly one
at the middle of the occiput, very short and of a length
no greater than that of 3 or 4 scales. The eyes are
of a considerable size and oblong, the longitudinal dia-
meter of the orbits occupying more than x/3 (about
35 %) of the length of the head, and the least breadth
of the interorbital space measuring at most 2/3 (60 — 49 %)
of the former. The nostrils are set as in the preceding
species. The intermaxillary bones form only about half
of the margin of the upper jaw, their lobate process
extending as far back as the bones themselves. The
hind extremity of the maxillary bones lies nearly below
the centre of the eye. The lower jaw is longer than
in the preceding species — though the upper jaw pro-

jects beyond it in front — but on this head we must
remark that in the preceding species we find a distinct
alteration due to age, by which the length of the lower
jaw' is increased even relatively during growth, and
young specimens of Macrurus Fabricii corresponding
in size to the adult specimens of M. coelorhynchus are
unknown. The pointed and fairly straight jaw-teeth
are not large, but form broad cards (containing several
rows) both in the lower jaw and on the intermaxillary
bones. The pharyngeal teeth are thicker, strong, and
of conical shape, with the blunt tip marked off as a
distinct crown. The gill-rakers are verrucose. The
operculum, as in most of the Macruroids, forms an al-
most right-angled triangle. The right (slightly obtuse)
angle is turned backwards, the lower posterior side
slightly concave, and the pointed end turned down-
wards, with the point fitted to the lower posterior
corner of the preoperculum, which is elongated in a
backward direction, so that the postorbital part of the
side of the head almost forms a square. The branchio-
stegal membranes are united below into a free but not
very broad fold across the isthmus.

■ The first dorsal fin, vdiich is set in a deep groove
in the dorsal edge, is high and trapeziform, the last
ray ■ — as in most of the Macruroids — being apparently
destitute of membrane behind it, and the posterior rays,
from the second branched ray, decreasing in length
with fair uniformity, so that the hind part of the upper
margin of the fin is straight or slightly convex. In a
specimen 725 mm. long the second branched ray is
77 mm. long, the second simple (the spiny ray) 73 mm.,
and the last ray 18 mm. The first-mentioned ray is
exactly as long as the lower jaw, the length of the
last-mentioned ray only slightly more than half the
distance between the two dorsal fins and somewhat less
than half the base of the first dorsal fin. In the form
of the second dorsal fin Macrurus Fabricii belongs to
a group of the genus that is characterized by the com-
plete development of this fin even in front, its begin-
ning being also situated in front of that of the anal
fin. The first ray is about as long as the last ray of
the first dorsal fin or slightly shorter than it; the next
rays gradually increase in length, though only slightly;
and the height of the fin then decreases extremely slowly
towards the tip of the tail, where it joins the anal fin.

The anal fin is more than twice as high as the
second dorsal fin throughout almost the whole of its
length, with a slightly convex margin. Its first ray is

GRENADIER-FISHES.

589

about as long as the last ray of the first dorsal fin,
and at the middle of the tin the length of the rays is
about 2/3 of the longitudinal diameter of the orbits;
from this point the height of this tin too decreases.
All the rays are articulated, and all, except the first
rays, in the anal tin as well as in the second dorsal
tin are bifid at the tip; all may be easily parted into
their two lateral halves.

The pectoral tins are situated somewhat in front
of (according to Sundevall, vertically below) the be-
ginning of the first dorsal tin. They are obliquely
pointed, and their length is somewhat greater than the
height of the first dorsal fin. The first ray is thick,
but articulated and simple or imperfectly branched at
the tip". The third and fourth rays are equal in length
and the longest in the fin''.

The ventral fins are about 2/3 as long as the pec-
toral, the first ray, simple but articulated, being elong-
ated to a hairlike tip. Among the branched rays the
4th, 5th, and 6th are equal in length and the longest,
thus giving this part of the fins a rounded form.

The scales cover the whole body, except the under
surface of the head, the branchiostegal membranes, and
the fins. On the head the lateral carinse of the snout
and cheeks also mark the boundary of the scales: though
scattered scales may be found below the hind part of
the suborbital carina. The scales on different parts of
the body vary both in size and in form; and they are
armed in various ways not only on different parts of
the body, but also in different individuals. The smallest
scales are set on the head, where they are more firmly
attached to the subjacent bones, especially on the longi-
tudinal carime. In form the scales are quadrangular,
rounded (almost circular), or hexagonal. The anterior
(inserted) part of each scale is smooth, the outer part
is furnished with a high and dentated, longitudinal
carina at the middle and, in most cases, with several
(3 — 5 above and below) small carime, radiating in a
backward direction and sometimes dentated like the
middle carina. The texture of the scale shows nume-
rous, dense and undulating, concentric strias, but no
radiating ones. The lateral line does not pierce any
scales; but its opening pores lie on the outside of a

row of scales in which, in most cases, the middle ca-
rina of the scale is divided (double). In other cases
the pore lies below the simple middle carina.

The body is of a grayish silvery or tin colour;
the fins are brownish black. The cavity of the mouth,
the branchial cavity, and the peritoneum are black.

Macrurus Fabricii is by no means a rare fish off
the south of Greenland and occurs along the east coast
of North America at least as far south as the neigh-
bourhood of Cape Cod. So early a writer as Egede
describes it as “like a Torsk, but with sharp prickles
and spines over the whole skin, and with a narrow
tail or hind part. They are both large and small. The
Esquimaux say that they are of good flavour.” Cranz
mentions a “species of fish that has a large head and
eyes0 like an owl, and which the Esquimaux call Ing-
minniset , because it bellows'* when dying.” On the coast
of Norwegian Fin mark, north of Tromso, the species
has been taken occasionally in recent times on long-
lines, at depths varying between 100 and 300 fathoms.
It was unknown there, however, until fifty years ago
(1839), when Professor S. Loven brought home to the
Royal Museum a specimen, which has been described
by Sundevall, from Hammerfest.

The food is probably composed chiefly of small
fishes, e. g. the Capelin, and crustaceans, even those
with hard shells, to judge by the strength and form
of the pharyngeal teeth. But in this case, as in the
case of other deep-sea fishes, the stomach is generally
turned inside out when the fish is drawn up to the
surface, so that it has been impossible to ascertain its
contents. The fish takes a bait freely, and is a nuis-
ance, says Brown-Goode, to the American fisherman,
who hauls in his line in the hope of a better catch
than an Onion-fish.

The spawning-season of Macrurus Fabricii occurs
on the coast of Finmark at the end of winter or
soon afterwards, for in May Collett has met with
at least two females full of roe. In Greenland Fabri-
cius found that the ovaries of the females were ex-
tremely small in the month of May, and he therefore
assumed that at this season they had generally finished
spawning.

“ On the other hand, it parts easily into its two lateral halves.

6 Gunther remarks a difference between the Norwegian and American forms of this species, consisting, according to his specimens,
in the much shorter pectoral fins of the former. In one of our specimens (stuffed) from Finmark, however, the length of the pectoral fins
is 54 % of that of the head.

e On account of the eyes the American fishermen of the present day call this species Onion-fish.
d When the air-bladder bursts (?)

590

SCANDINAVIAN FISHES.

MACRURUS RUPESTRIS (sw. skolasten”).

Plate XXVII A, fig. 2.

Snout blunt (of a broad pyramidal shape) and like the rest of the head without sharp (with only blunt) carince;
its length less than the least breadth of the interorbital space , which is greater than or equal to J/3 (38’2 — 33*2 %
in specimens examined by us, which are between 121 and 885 mm. long) of the length of the head and greater
than the longitudinal diameter of the orbits (133 — 144 %) or more than 2/3 (81 ’6 — 70‘9 %) of the postorbital length
of the head , than V3 (37'5 — 33‘8 %) of the distance between the first dorsal fin and the tip of the snout , than 7/10
(72 — 93 %) of the breadth across the cheeks at the centre of the eyes , or than i/-0 (89*6 — 88'4 %) of the length of
the lower jaw , which is less them 1/2 (42'6 — -37 %) of the length, of the head , or about. 2/5 (42‘2 — 37*5 %) of the
distance between the first dorsal fin and the tip of the snout, and less than the postorbital length of the head.
Mouth nearly terminal ancl lateral; its breadth at the corners more than 2/3 (70 — 86'5 %) of the breadth across
the cheeks at the centre of the eyes. Length of the head about 1 6 — V5 (18 — 20 %) of that of the body, about
equal to (105 — 95 % of) the distance of the first dorsal fin from the tip of the snout, about l/2 (54 — 45 %) ol that
between the second dorsal and the same point, and about 2/3 (73*2 — 61 '7 %) of the distance between the anal tin
and the tip of the snout. Depth of the body (in adult specimens) at the beginning of the first dorsal fin about */« °f
the length, at the beginning of the second dorsal fin about lj10 or 1/9 of the same. Pectoral fins without brachiate
base. Length of the base of the first dorsal fin about x/3 (28 — 37 %) of the distance between the two dorsal
fins, which is nearly twice the breadth across the cheeks at the centre of the eyes. 6 or 7 scales in a row
from the end of the first dorsal fin to the lateral line. No bare spot (except the anal aperture) in the median
line of the belly. Scales densely covered with spines throughout their free surface, but without carince. Second
ray of the first dorsal fin spiny or (in old specimens) at least granulated on the front, surface; its length in
young specimens greater, in old generally less, than 2/3 of the depth of the body at its base. Length of the
barbel under the chin less than (or about equal to) 1/6 of that of the lower jaw. Jaw-teeth car di form, of uni-
form size or with the teeth in the outer row perceptibly larger than the others.

R. hr. 6; Dr (1) + --- D.r 103—172 (190); A. 104—193;

O 1 0

P. 18 — 23; V. ; Lin. squ. transv. 2G — 27 (ad pinn. ventr.);

6 — 7

Vert. 63 — 89.

Syn. Berglax (Coryphsena dorso dipterygio, pinna dorsi prima retro),
Strom, Sondmors Beskriv., vol. I, p. 267 — 269, not.
Coryphcenoides rupestris, Gunn. Trondhj. Selsk. Skr., vol. Ill
(1765), p. 50, tab. Ill, figg. 1 et 2; Coll., Forh. Vid. Selsk.
Christ. 1874, Tilhvgsh., p. 131; ibid. 1879, No. 1, p. 70;
N. Mag. Naturv. Christ., Bd. 29 (1884), p. 95; Malm, Gbgs,
Boh. Fn., p. 502; Winth., Naturh. Tidskr. Kbhvn, ser. 3,
vol. XII, p. 36; Jord., Gilb., Bull. U. S. Nat. Mus., No.
16, p. 812; Goode, Bean, Bull. Mus. Comp. Zool., Harv.
Coll., vol. X, p. 197; Day, Fish. Gt. Brit., Irel., vol. I,
p. 335, tab. XCIII; Storm, N. Vid. Selsk. Skr. Trondhj.
1883, p. 57: Lillj., Sv., Norg. Fisk., vol. If, p. 259;
Gthr, Deep Sea Fish., Cliall. Exped ., p. 138.

Macrourns Stroemii, Beinh., D. Vid. Selsk. Naturv., Math. Afb.,
vol. V, p. XIX; Sundev., Vet.-Akad. Handl. 1840, p. 11;
Kr., Voy. Scand., Lapp., cett., Gaim., Zool., IJoiss., tab. 11.
Lepidoleprns norvegicus, Nilss., Prodr. Ichth. Scand., p. 51;
Id. ( Macrourus ), Skand. Fn., Fisk., p. 600; Gti-ir ( Cory -
pheenoides ), Cat. Brit. Mus., Fish., vol. IV, p. 396; Ltkn,
Vid. Meddel. Naturhist. For. Kbhvn 1872, p. 5 (sep.).

Macrurus rupestris is one of the Macruroids that
have the second dorsal fin extremely little developed

in front, while behind the fin rises somewhat higher
than usual, so that a little in front of the tip of the
tail it is only slightly lower than the anal fin at the
same point. The eyes are rather small, the longitudinal
diameter of the orbits being less than 3/io (— 27 — 23 %)
of the length of the head, a sign that the deep-sea
character is not so pronounced as in the preceding
species. The carinse on the head are blunt, though
they occupy the same position as in the preceding
species and in dried specimens or those which have
long been preserved in spirits, are distinctly prominent.
These three characteristics are enough always to render
this species easily recognisable among its congeners in
our fauna. However, the interspinal bones (the sup-
ports of the dorsal rays) are perfectly developed be-
tween the muscles of the back even in the space be-
tween the two dorsal fins, where no rays are present;
and in a young specimen, 121 mm. in length, we ob-
serve not only that the second dorsal fin begins with
distinct rays considerably further forward than in old
specimens and only slightly behind the first third of

= Shoemaker’s last.

GRENADIER-FISHES.

591

the body, but also that in the dorsal margin, in front
of the first distinct ray of the second dorsal fin, there
lies a row of soft, whitish, dermal protuberances, while
from each of the posterior among these protuberances,
forward to a point 3 /jo of the way along the body from
the tip of the snout, there projects the tip of a ray,
extremely small and scarcely perceptible. There can
be no other explanation of this than that the second
dorsal fin originally extended much further forward
even than it does in adult specimens, but that its an-
terior part undergoes a reduction. Still, it is a rule
— though with considerable individual variations —
as Collett has pointed out, that the number of rays
in this fin as well as in the anal fin increases with acre
during the latter period of growth, even while the length
of the body increases from half a metre to a metre.
This increase — from about 100 to about 200 in the
former fin" — is in its magnitude an extremely un-
common phenomenon in the class of fishes. Collett
also connects it with another great abnormity, the in-
crease in the number of the vertebras — 13 — 14 ab-
dominal and 49 — 75 caudal vertebras — which proceeds
according to the same rule and with the same exceptions6.

The first dorsal fin is set further forward than in
the rest of the Scandinavian Macruroids, its beginning
lying in front of the end of the first fifth of the body.
It moves somewhat further back, however, with age.
The length of its base is less than i/5 (79 — 58 %) of
the least breadth of the interorbital space. The first
ray is as usual rudimentary, the second ray both the
longest and the thickest, nnarticulated, and measuring
rather more than half (50 — 55 %) the length of the
head. The other rays — except the last, which is also
simple — decrease in length uniformly, but sharply,
the fin thus • acquiring a more triangular form than in
Macrurus Fabricii.

The anal fin begins at a distance from the tip of
the snout which in young specimens measures about J/4
of the length of the body, in older ones about 1/3 thereof.
It is of fairly uniform height, with slightly concave
margin: the first ray is slightly shorter than the next
ones, the length of the rays increases or decreases ac-
cording to the curvature of the belly, and behind the
middle of its length the height of the fin is greater than

the depth of the body at the^same point. Round the
tip of the tail the anal fin coalesces with the second
dorsal fin into a symmetrical tip. The vent is situated
just in front of the beginning of the anal fin.

The pectoral fins are set just in front of the per-
pendicular from the beginning of the first dorsal fin.
They are evenly rounded at the tip, and their length
is perceptibly more than l/3 (58 — 6l72 %) of that of
the head. The first ray is extremely short, and the
ninth (8 — 10) is the longest.

■ The ventral fins are situated in old specimens in
front of the perpendicular from the beginning of the
first dorsal fin; but during youth they lie considerably
behind it. The usual removal in a forward direction of
these fins thus takes place in this species too, the distance
between the foremost point in the insertion of the ventral
fins and the beginning of the anal fin in young speci-
mens (12 cm. long) being only slightly more than Vs
(22'8 %) of the distance between the anal fin and the
tip of the snout, but in old specimens nearly 1/3 (30 or
31 %) thereof. The first ray is elongated to a long tip,
hairlike at the extreme end, which extends to the lie-
ginning of the anal fin; the second ray is about V3 as
long; and the length of the other rays is such that the
margin of the fin is somewhat convex.

The jaw-teeth form a kind of transition from the
preceding species to the following one. On the inter-
maxillary bones (in adult specimens) they are set in a
card containing 5 or 6 rows, but behind this card, for
the greater part of the length of the bones, they are set
in only 2 rows. In the lower jaw the case is similar,
but here there is only one row behind. In front the two
intermaxillary cards have an empty space between them.
The pharyngeal teeth — 2 or 3 rows on the two anterior
upper pharyugeals, 4 rows on the posterior upper pha-
ryngeal; 6 rows on the oblong, curved lower pharyugeals,
with the largest teeth in the innermost (hindmost) row
— are straight and conical, without the sharp break at
the apex which characterizes them in Macrurus Fabricii.

The anterior nostrils are as usual round, the posterior
oblong and set transversely just in front of the orbital
margin; but the latter are generally remarkably narrow,
this being due to the projection throughout the whole
of their length of the marginal membrane in front.

O o

° Still, it is not a superfluous remark that mistakes may easily be made in the counting, partly on account of the magnitude of the
numbers, and partly because the two lateral halves of the otherwise simple rays easily fall apart, and the rays are thus counted twice over.

b Both the rule and the exceptions should be kept in mind, when one has to judge of the systematic significance of the varieties or
races which, especially among the Salmonidce and Clupeidce , have been based upon the variations in the number of the vertebra;.

Scandinavian Fishes.

75

592

SCANDINAVIAN FISHES.

In this species the scales extend over the whole of
the head, even below the suborbital ring, on the branches
of the lower jaw, and in a large patch on the space
between the branches of the lower jaw ( nientum ); only
the branchiostegal membrane and the isthmus are as
usual bare. The scales" are of essentially the same form
as in the preceding species, but show no longitudinal
carime or curves, and are densely covered throughout
their free surface with short spines, which are arranged
in a quincunx of oblique, curved rows. These spines
give the surface of the body a velvety appearance. The
lateral line does not pierce any scales in this species
either, its opening pores lying in the form of grooves
without spines on each scale.

Among the internal organs the liver and the pyloric
appendages are especially remarkable for their advanced
development. The abdominal cavity, with the peritoneum
coloured black in front and chocolate behind, extends to
a line with about the ninth ray of the anal fin, and
the liver extends equally far back on the left side.
The ovary is simple and triangular, with the oviduct in
the lower, blunt corner.

The body is plain grayish violet shading into yel-
low. The tins are of a dark brownish violet, with a
dash of the blue tinge of dew. The iris is whitish
yellow, shading into silver and rose-pink. The mouth,
the pharynx, and the branchial cavity are black.

Macrurus rupestris, which attains a length of at least
a metre, has been found not so very seldom — for a deep-
sea fish — on the south coast of Norway, up to Helgo-
land, at depths between 150 and 500 fathoms. Accord-
ing to Storm it is taken very often on long-lines in
Trondhjem Fjord, even at the very head of the fjord,
but seldom in less than 150 fathoms of water. In a
single day in 1880, according to Collett, 20 specimens
of this species were caught oft’ Bjornor (in the south
of the Government of Trondhjem, lat. 64° 10 N.). Off
the extreme south of Norway, young specimens between
10 and 12 cm. long were taken off Arendal during the
expedition of the Swedish gunboat Gunhild in July, 1879,
at a depth of 335 — 370 fathoms and on a bottom of fine,
brown clay. It is thus quite at home in these regions.
It also penetrates into Christiania Fjord, at least to the
latitude of Horten. On the coast of Bohuslan it was
first found by Fries, who has left in the Royal Museum

“ See Iyr0YEr’s plate in Gaimard’s voyage.
b Rink, Gronland, Bd. 2, Naturh. Tiling, p. 25.
c Arctic Manual and Instructions , 1875, p. 120.

a specimen 78 cm. long, taken in May, 1838. Malm
found a stuffed specimen in Uddevalla Museum that had
been purchased in the fishmarket there in March, 1872.
In recent years Mr. C. A. Hansson has forwarded to the
Royal Museum two specimens. The first, which has
been about 83 cm. long, but is now broken off short
46 cm. from the snout (the caudal part is lost), was
found floating dead at the surface, between the Foster
and Weather Islands, on the 20th of June, 1882. The
second specimen, the original of our figure, is a female
885 mm. long, and was taken on a long-line in about
80 fathoms of water, off Svangen in South Ramso Fjord,
twelve miles south-west of Stromstad, on the 5th of
November, 1889. In March, 1877, according to Wintiier,
two specimens were cast ashore near the Skaw and
forwarded to Copenhagen Museum. This is all that is
known of the occurrence of the species in the Skager Rack.
In the Cattegat (south of the line from the Skaw to Mar-
strand) it has not yet been found. To the west, according
to Gunther, it has been met with by the English Por-
cupine and Knight Errant expeditions in 200 — 500 fa-
thoms of water between the Shetland and Faroe Islands.
In 1880, during the Blake expedition, according to
Brown-Goode and Bean, A. Agassiz obtained a specimen
804 mm. long, at a depth of 524 fathoms, in lat. 41°33' N.
and long. 65° 55' W. It is included among the fishes
of Greenland by the younger Reinhardt6 and LutkenS
Macrurus rupestris in all probability leads the same
life as other deep-sea fishes, though it seems not to be
of so pronounced character as our other Macruri , for,
as we have mentioned above, it has been taken in 80
fathoms of water. Its food seems to be composed chiefly
of crustaceans: Decapods, at least as large as shrimps,
and Hyperioids have been found in its stomach. The
spawning-season seems to occur in winter, perhaps even
in autumn. On the 12th of November Collett met
with a gravid female off Langesund. The female secured
by Mr. C. A. Hansson from Svangen and the original
of our figure had eggs 1 millimetre in diameter in its
ovary on the 5th of November; and as, according to
Collett, the diameter of the ripe eggs is 2 millimetres,
it had still some time to wait before depositing its
spawn. According to Collett the number of the ripe
eggs when the roe is deposited is about 12,000 or hardly
more than that number.

GRENAI ) I ER-FISH ES.

593

MACRURUS LJEYIS.

Fig. 141.

Snout Hunt but high , only slightly projecting, and like the rest of the head without sharp carince; its length less
than the least breadth of the interorbital space, which is about equal to x/3 (33*1 % in the specimen examined by
us, which has probably — when perfect — been about 41 cm. long) of the length of the head, slightly less than
(937 % of) the longitudinal diameter of the orbits, about 2/5 (41 ‘1 %) of the postorbital length of the head, and
more than either x/4 (28'5 %) of the distance between the first dorsal fin and the tip of the snout or than.\/l0
(86*5 %) of the breadth across the cheeks at the centre of the eyes, but less than 2/3 (59'8 %) of the length of the
lower jaw, which is more than 1j2 (55*3 %) of the length of the head, nearly 1/2 (477 %) of the distance between
the first dorsal fin and the tip of the snout, and greater than the postorbital length of the head. Mouth, almost
terminal and lateral; its breadth at the corners more than 2/3 (80'8 %) of the breadth across the cheeks at the
centre of the eyes. Length of the head somewhat less than X/G of that of the body", about 4/5 (86‘3 %) of the
distance of the first dorsal fin from the tip of the snout, nearly x/2 (47*8%) of that between the second dorsal fin
and the same point, and nearly 3/4 (73* 1 %) of the distance between the anal fin and the tip of the snout. Depth of
the body at the beginning of the first dorsal fin about x/7 (15 %) of the length of the bod}', at the beginning of

the second dorsal fin about x/10 thereof. Pectoral fins with brachiate base. Length of the base of the first dorsal

fin about 3/s °f the distance between the two dorsal fins, which is nearly twice (185 % of) the breadth across

the cheeks at the centre of the eyes. About 14. — 16 scales in a row from the end of the first dorsal fin to the

lateral line. In the median line of the belly, just in front of the insertions of the ventral fins, we find a trans-
verse, oblong (kidney-shaped) patch without scales and just in front of the vent a similar, but round patch.
Scales spiny throughout their free surface, but without carince. Second ray of the first dorsal fin smooth {without
spines ); its height about 2/3 (69 %) of the depth of the body at its base. Length of the barbel under the chin
more than x/3 of that of the lower jaw. Jaw-teeth set in two rows on the intermaxillary bones ( the outer row
with larger, canine-like teeth), in one row of canine-like teeth in the lower jaw.

R. br. 7 ; Z>, .

(D +

D2 ■ 200 (ca);

A. 200 (ca);

p. m + f r.

1/

/S'

Syn. Macrounis ler.vis, Lowe, Proc. Zool. Soc. Lond. 1843, p. 92;

Gthp. ( Malacoceplialus ), Cat. Brit. Mas., Fish., vol. IV, p.
397; Ltkn, Vid. Meddel. Naturh. For. Kblivn 1872, p. 1
(sep.); Malm, Gbgs, Boh. Fu., p. 503; Winth., Nat. Tidskr.
Kblivn, ser. 3, vol. XII, p. 36; Lillj., Sv., Norrj. Fisk.,
vol. II, p. 273; Gthr ( Macrurus ), Deep Sea Fish. Chall.
Expect., p. 148, tab. XXXIX, fig. B.

Macrurus Iceris, which attains a length of about
half a metre, with eyes relatively equal in size to those
of M. Fabricii (35 % of the length of the head) is
evidently a much more pronounced deep-sea fish than
M. rupestris; and its more scattered, but stronger
(canine-like) jaw-teeth and relatively longer jaws are
signs of its more predatory habits.

The body is remarkably elongated, this being due,
according to Gunther, to the fact that the tail “tapers
to a very long and narrow band.” But the forepart

externally corresponds fairly closely to that of the two
preceding species, the depth at the beginning of the
first dorsal fin being about !'/10 (897 %) of the length
of the head, and at the beginning of the second dorsal
fin about 2/3 (647 %) of the same. The breadth of
the head straight across the cheeks at the centre of
the eyes is less than 2/5 (38'2 %) of its length.

The smooth spinous ray in the first dorsal fin and
the small scales render this species easily recognisable
among our Macruri. We must remark, however, some
(5) raised, but only slightly marked stria; (osseous
ridges under the skin) on the operculum and suboper-
culum, radiating backwards and downwards from the
articulation of the former.

The coloration, according to Gunther, is brownish
on the head, especially on the gill-cover, with a lustre
of silver and gold; the axil of the pectorals, the ventral
fins and the belly between them, and the vent are black;
the dorsal and pectoral fins are blackish; the anal fin

a According to Gunther and Lutken.

594

SCANDINAVIAN FISHES.

is black at the margin. The inside of the month is
white, the branchial cavity black.

Macrurus Icevis lias a wide geographical range, but
is still extremely rare in the museums. It was disco-
vered off Madeira by Lowe, and the British Museum

at the beginning of December, 1871. Long before this
time, however, the Museum of Gothenburg had received
a specimen through Mr. Deyenbeeg. This specimen
was dropped on the shore near Lysekil by a seagull on
the 10th of November, 1852; and has since been de-

Fig. 141. Macrurus Icevis , seen from the side, together with the forepart, seen from below. 2/3 of the natural size. Found at Lysekil, on
the 10th of November, 1852, by Deyenberg. The property of the Natural History Museum of Gothenburg.

has acquired another specimen from the same locality ,
through Johnson. But the species was not recognised
again until Lutken described a specimen that was cast
ashore on the north of Jutland near the Skaw and found

scribed at length by Malm in his Goteborgs och Bolius-
Icins Fauna*. Finally the species has again been found
off the coast of Pernambuco, where the Challenger Ex-
pedition took a- specimen at a depth of 350 fathoms b.

Fam. OPMIBIIDiE.

Body of a compressed tadpole-like or Eel-like form, ivith elongated, more or less whip-like tail , and covered with
thin, cycloid scales or naked. All the vertical fins confluent (no distinct caudal, no anterior dorsal fin). Teeth as
a rule present both, in the jaws and on the vomer and, palatine bones. Gill-openings large: the branchiostegal mem-
branes more or less united to each, other, but free from the isthmus. Branchiostegal rays usually S or 7C. Pseudo-
branchiae and air-bladder generally present11. Pyloric appendages only slightly developed or wanting.

This family consists
fishes; and the reductions
nature appear here too.

in great part of deep-sea
characteristic of a life of this
But the same reductions in

the complete structure of
affects those fishes of this
terranean chasms or lead

the Anacanthine type also
family which live in sub-
a kind of commensual life

“ By the kindness of Dr. Stuxberg, the Curator of the Museum, we have been enabled to make use of this specimen as the original
of our figure.

b Whether it occurs in the Mediterranean, is still somewhat uncertain, though probable. In his list of the Mediterranean fishes, how-
ever, Giglioli gives only a name that might be applied to it; and it is evidently another species that Moreau has described under this name.
c Exceptionally 6 or even 5.
d Often wanting, however.

OPHIDIOID ONOMORPHS.

595

within the body of other animals. Even true surface-
fishes or shore-fishes also occur within this family. In
the great majority of cases we may rely upon the
family-character that the caudal fin is not separated
from the other vertical fins; but it very often appears,
at least by a closer arrangement of the rays, that here
we have the rudiment of a distinct caudal fin, and in
some exotic species the caudal fin is free or united to
the dorsal and anal fins merely by a short fin-mem-
brane. In most cases the ventral fins are reduced,
filamentous, and removed to a greater or less distance
in a forward direction, even in front of the shonlder-
girdle and below the tongue, as if they were barbels.
In other cases these fins are wanting. Some of these
fishes in form come very near the Macruri, others
remind us of the Plvycis- type, and others again resemble
the Blennomorphs and Gobiomorphs.

In general these fishes are small and worthless;
but Genypterus capensis, which sometimes attains a,
length of a metre, bears among the inhabitants of Cape
Colony the significant name of Koning van Klipvischen ,
and Brotula multibarbata is a favourite and expensive
dish in Japan.

About 75 species are at present known with
more or less certainty and distributed among 33
genera.

The family as now defined and ranged most
nearly corresponds to Muller’s" family Ophidini, with
the addition (made by Gunther* * 6) of the Cuvierian
Gadoid genus Brotula and its relatives (the sub-
family Brotulince), of Richardson’s genus Maclicerium
(subsequently altered for reasons of priority to Con-
grogadus ), and of Ruppel’s Haliophis (the subfamily
Gongrogadince).

Subfamily OPHiDIIN IE.

Filamentous ventral jins inserted in front of the shoulder- girdle.

The majority of the members of this subfamily
are shore-fishes; but the only one that can be of any
faunistic interest to us, is a deep-sea fish.

The genus Ophidium , which has given its name
to the family as well as to the subfamily, is fairly
common in the Mediterranean and on the coast of
Brazil. In the former locality it has long been known:
even Pliny probably referred to it when he mentioned
a fish of this name that was employed in the medicine
of the ancients, while in Belon’s time0 the fish was
much esteemed by the prelates of Rome as an excellent
dish. In more recent times the genus has gained ce-

lebrity, through Muller’s d and Costa’s6 researches, for
the singular manner in which the air-bladder is united
in front to the spinal column, in the nearest resem-
blance to the corresponding structure in Fierasfer.
Muller imagined that within this genus he could make
use of anatomical differences as specific distinctions be-
tween forms in which no external differences could be
detected; but Costa showed that a, great proportion of
the former appear as differences dependent on age and
sex. One species has a range extending from the Medi-
terranean out into the Atlantic and up to England, but
has not been found in Scandinavian waters.

a Abli. K. Akad. Wiss. Berl. 1844, pp. 177 and 202. Bonaparte had already established a family Ophidiidce in 1831, but he ranged

it among the Apodes and included in it the Sand-Eels.

6 Cat. Brit. Mus., Fish., vol. IV, p. 370.

0 La nature et diversite des poissons, Paris 1555, p. 126.
d Abh. K. Akad. Wiss. Berl. 1843, p. 150.

c Fauna del regno di Napoli , Pesci , part. 1, Ophidium , tav. XX, ter. August, 1843.

596

SCANDINAVIAN FISHES.

Genus RHODICHTHYS.

Body tadpole-like , with fairly deep and broad head and strongly compressed tail ,
the great median height of the vertical fins , the margins of which evenly but sharply converge
fin , which is united at the base to the dorsal and anal fins, but for the greater part of its length free,
fins filamentous, bifid in the outer part, and inserted
Anal aperture situated, in front of the pectoral fins,
able distance from the beginning of the anal fin.
lower jaw; vomer and palatine bones toothless.

the naked skin transparent.

This genus is one of the most handsome discoveries
made in the depths of the sea by the Norwegian Arctic
Expedition. The form of the body reminds us strongly
enough of the Great Forked Beard; but the loose, al-
most gelatinous, and naked, almost transparent body
most nearly resembles that of the other true deep-sea
fishes within the Ophidioid family. According to Col-
lett the ventral fins are “firmly united to the hyoid
bone'' — the genus is thus referred to this subfamily —
but according to the figure these fins are set compara-
tively far back, considerably further back than in a true
Ophidium, and Lilljeborg says they are “attached to
the shoulder-girdle. " In the latter case the genus ought

which is increased in depth by
towards the caudal
Ventral

beloiv the posterior part of the copular row of the hyoid bone,
between the lower corners of the gill-openings, at a consider-
Small, cardiform teeth on the intermaxillary bones and in the
Branchiostegal rays 6. Pyloric appendages 10. No scales;

No spines on the head.

rather to be ranged among the subfamily Brotulince.
In the form of the body too, as Lilljeborg has already
remarked, it comes very near Bathyonus, one of the
Brotulince: but the opercular spine and the scales are
wanting, and in Bathyonus the vent is situated close
to the beginning of the anal fin. The limit between
these subfamilies is, however, rather uncertain, fixed
as it is according to a character which is an expres-
sion for the general rule among the Physoclysts that
the ventral fins move forward with increasing age and
advancing development of form.

Only one species of the genus is known.

THE ROSE FISH (sw. kosenfisken).

EIIODIC HTH YS REGINA .

Fig. 142.

Length of the head {in a male nearly 3 dm. long ) about 1/i of that of the body. Greatest depth of the body about

1/5 of its length and almost equal to the distance between the vent and the tip of the snout. Breadth straight

across the cheeks greater than the postorbital length of the head and about 3/5 of the entire length of the head.
Breadth of the interorbital space about s/5 of the entire length of the head and about 3/i of the postorbital length
thereof. Outer diameter of the iris 1/7 of the entire length of the head and somewhat more than 1/4 of the post-
orbital length of the head or than i/9 of the length of the snout. Mouth broad, both terminal and lateral; the

maxillary bones extend somewhat behind the perpendicular from the hind margin of the iris. Lower jaw shorter

in front than the upper jaw. Nostrils set far apart, the posterior pair separated from the eyes by a distance equal
to the diameter of the latter, the anterior pair about, half-way between, the posterior and the tip of the snout. Height
of the gill-openings equal to the breadth across the cheeks. Distance between the dorsal fin and the tip of the snout
slightly greater than the length of the head and about 4/5 of the distance between the anal fin and the same point.
Greatest, height of the dorsal fin 3/10 of the depth of the body at the beginning of the fin; greatest height of the

anal fin 3/10 of the depth of the body at the beginning of the fin. Depth, of the base, of the caudal fin rather

more than l/3 of the diameter of the iris. Length of the caudal fin equal to the breadth of the interorbital space.
Length of the pectoral fins about equal, to the postorbital length of the head. Length of the ventral fins about equal
to the distance between the anal fin and the tip of the snout. Coloration dark carnation, with a few lighter, clouded
spots; branchiostegal membranes deep crimson; peritoneum, branchial cavity, and pharynx black.

R. hr. 6; I). GO; A. 57; P. 11 1. 12; C. 10. 154, tab. V, figg. 37 — 39; Lillj., Sv., Norg. Fisk., vol.

Syn. Rhodichthys regina, Coll., Forh. Vid. Selsk. Christ. 1878, II, p. 238; Gthr, Deep Sea Fish., Oh all. Exped., p.

No. 14, p. 99; N. Nordh. Exped., Zool. , Fiske, p. 121.

ROSE FISH.

59 7

The above description and the appended figure are
derived from Collett’s work on the deep-sea fishes
taken by the Norwegian Arctic Expedition. Only one
specimen of this species is as yet known. It was taken
on the 16th July, 1878, at a depth of 1,280 fathoms, 465

board, although the hauling of the trawl had taken se-
veral hours; and it was kept alive for some time, but
appeared to lie extremely sluggish in its movements.
The transparency of the body rendered the internal
organs visible, so that in the living fish one could see

A" 'ON

mmmm i

Fig. 142.

Rhocliehthys regina , <p, 2/3 of the natural size. From a depth of 1,280
perature of — 1’4° Celsius), 465 kilom. N. E. of Jan Mayen I.,

fathoms, on a clayey bottom ( Biloculina-chxy with a tem-
19th July, 1878. After Collett.

kilometres north east of Jan Mayen Island, about mid-
way between this island, Norwegian Finmark, and Bear
Island. The bottom at this depth consisted of Bilocu-
lina- clay, and the bottom temperature was — 1 '4° Cel-
sius. The fish was still alive when it was drawn on

the spinal column, the several divisions of the brain,
the labyrinth, the gills, and most of the viscera. In
the stomach were found still determinable remains of
three crustaceans, Bythocaris leucopis, Pseudomysis
abyssi, and a Iiyperioid.

Subamily FIERASFIN M\

No ventral fins. Body Eel-like , with pointed tail.

“Fishermen say that of this sea-cucumber (Cazzo | red, ribbon-like fishes which are called Cipollefi wrote
reale — Holothuria tubulosa) are generated the oblong, | Columna6 at the beginning of the 17th century. So

a Gthr, Cat. Brit. Mas., Fish., vol. IV, p. 370.

b De aquatilibas aliisque nonnullis animalibus, Romae 1616, p. XXXVII.

598

SCANDINAVIAN FISHES.

long has it been known that a singular and intimate
relation exists between sea-cucumbers and certain fishes
of this subfamily which in Cuvier" formed the sub-
genus Fierasfer of the genus OpMdiiim. These fishes
have the habit of seeking shelter and, to some extent,
food inside sea-cucumbers and bivalves, where they lead
a parasitic life, feeding upon the minute animals drawn by
the current into the respiratory tree of the sea-cucumber
or towards the mouth of the bivalve. Now and then
they make sallies in search of food, but soon return
to their hiding-place.

whose habits are best known, the Mediterranean F. acus ,
usually makes its way in the same manner, according
to Emery, tail foremost into the respiratory tree and
body cavity of the Holothurice. Soft, almost transparent,
and helpless, without weapons of defence, not even with
scales to protect its body, and with powers of sight
certainly feeble during the daytime, as shown by the
contracted pupils, the fish feels its way, with head
turned down, until with its snout and Avith the system
of the lateral line it feels the current that passes into
the hind part of a Holothuria. Noav the fish bends

Fig.

143.

Fierasfer acus , in the act of entering a Holothuria. 2/3 of the natural size. After Emery.

Hoav this habit has arisen, Ave can at least con-
jecture from Verrill’s observation'' of a species of the
preceding subfamily, Ophidium marginatum, in Great
Egg Harbour on the coast of NeAv Jersey. “We dug
tAvo specimens,” he Avrites, “out of the sand near Ioav-
Avater mark, Avhere they burroAved to the depth of a
foot or more. When placed upon moist sand they
burroAved into it tail foremost Avith surprising rapidity,
disappearing in an instant.” That species of Fierasfer

the tip of its tail doAvmvards and inserts it into the
cloaca of the sea-cucumber. In adult specimens of this
species the tip of the tail is Avithout fin-rays, but the
last rays of the dorsal and anal fins lie close to the
body at this spot, so that the end of the tail forms a
comparatively stiff point, Avhich can be inserted Avithout
difficulty into the opening. By the breathing of the
sea-cucumber and the undulating movements of the
fish itself the tail is iioav forced farther and farther

“ Regne Animal , ed. 2, tome II, p. 359. In Risso Fierasfer was the specific name of Fierasfer acus ( Ophiclium imberbe , Lin., p. p.).
1 American Naturalist, vol. \r (1871), p. 399.

OPHIDIOIDS.

599

into the respiratory tree, until only the forepart remains
outside. The fish now entirely enters the body of the
sea-cucumber by means of a few powerful, undulating
movements. Now and then it peeps out to breathe, to
seize some small crustacean that has come too near,
or to evacuate the contents of its intestine through
the vent, which is situated far forward. The fish must
also, in all probability, leave its place of refuge and
lead an independent life during the spawning-season.
After the first fish a second, a third, or even a seventh
may take up its abode in the same sea-cucumber. One
Fierasfer seems generally to do no harm to the sea-
cucumber; but if several come, it dies after ejecting
its entrails, and the fish peep out from the empty skin.
Thus, the Fierasfer lives originally in the ramified
breathing duct (respiratory tree) of the sea-cucumber;
but by the bursting of the walls of this duct it easily
gains a, passage into the large cavity of the body,
where it is frequently found. Some species of the ge-
nus lead a similar life in the body cavity of the Indian
t repangs, and in all probability gain entrance in the
same manner; but the Fierasfers that live within the
body of the large starfishes (genus Culcita), must cer-
tainly pass through the mouth, for in these animals
there is no anal aperture (cloaca) to admit of the pas-
sage of any fish.

Even though these fishes may derive some nourish-
ment from the fluids in the body of the sea-cucumber
in which they live, still they are no true parasites, and
seek their food chiefly outside, thus being frequently
found swimming freely about. The case may be
different in the other genus of this subfamily, Enclie-
Hophis, which is a native of the waters round the Phi-
lippine Islands, and whose parasitic nature seems to be
more pronounced and to have involved the loss not
only of the ventral, but also of the pectoral fins.

From the Gulf of Panama, on the west coast of
Central America*, Putnam has described a species, Fie-
rasfer dubius, that hospitates in pearl-mussels, be-
tween the mantle and the shell. Here it leads a life
so quiet and inoffensive that the mussel sometimes
succeeds in enveloping it in a layer of nacre.

The weak sight possessed by these fishes, at least
by daylight, is compensated by a special development of
the 'senses of touch and hearing. The system of the

lateral line (fig. 144) includes large passages on the
head, from the snout round the eyes (the suborbital
and supraorbital branches), along the forehead and the
top of the head on each side to the temples, and from
the under surface of the lower jaw, along the margin
of the preoperculum, to the same spot, where the lateral
line proper begins. The two lateral halves of the sys-
tem are united by transverse canals between the eyes
and between the temples. The sensory organs lie in
large numbers along these canals as well as in a row
along each side of the back and in another denser row
along each side of the base of the anal fin. The laby-
rinth is remarkable for its comparatively large otoliths.
The powers of hearing are probably strengthened —
and sounds perhaps produced, though we have no ob-
servations on this head — principally by the connexion

Fig. 144. Cephalic system of the lateral line (“muciferous canals’’)
in Fierasfer cleutatus. A. from above; B. from the left.

After Emery.

(probably common to all the Ophidioids) between the
air-bladder, the first vertebrae, and the base of the
occiput. In the Cods we have seen an internal con-
nexion between the air-bladder and the transverse
processes of the middle and posterior abdominal verte-
brae. Here this connexion is removed farther forward;
and the first four (or, as in Fierasfer dentatus, only
the first three) vertebrae with their ribs are modified
accordingly. The first vertebra is as usual close! v
united to the occipital bone, the second is shortened.
The first two pairs of ribs retain a. greater or less amount
of mobility, the third (and in Fierasfer acus even the
fourth) pair coalesces with the body of the vertebra
like ordinary transverse processes. In Fierasfer acits
the third pair is so much enlarged that it coalesces

a According to Putnam (Proc. Soc. Nat. Hist. Boston, vol. XVI (1874), p. 343) the same species inhabits Holothurians on the east
coast of Central America, in Florida Bay.

Scandinavian Fishes.

76

600

SCANDINAVIAN FISHES.

not only with its own vertebra, but also with the
fourth. The ribs of the latter have assumed the form
of disks united to its back. In Fierasfer dentatus the
third pair is only slightly extended, the second and
first pairs all the more so, though far less than the
third pair in the former species. To these bones is
attached the anterior part of the air-bladder — which
extends along- the whole dorsal wall of the abdominal
cavity, which is much broader in acus than in dentatus
— and here are extended muscles to draw the air-
bladder forward, among which the median are especially
large and powerful, originating from and coasting the
parasphenoid bone and attached to the anterior end of
the air-bladder.

The eggs of these fishes are developed floating-
in the sea, united within a slimy mass, like those of
the Angler. Of the spawning (deposition of the eggs)
we know only that it takes place at night. The deve-
lopment advances quickly. By the sixth day the yolk
is absorbed, and the larva begins to eat. It is now
only 3 or 4 mm. long, but already Eel-shaped, and,
in Fierasfer acus , has a singular appendage on the
back, almost as long as itself. This is the first- ray
of the future dorsal fin, which has early grown into a
long, stiff stem, with foliate growths set alternately.
Mere we again meet with an example of the long-

appendages which we have already observed on several
occasions in the larvae of pelagic fishes, and which help
the larvae to support themselves as they float in the
water and drift with the current. These appendages
probably give the larvae a protective likeness to small
Medusae, and perhaps, as Emery supposes, serve as a
lure to entice the small crustaceans ( Entomostraca ) on
which the larvae live". This evanescent growth does
not appear in the larva of Fierasfer dentatus , in which,
on the other hand, the body is still more elongated
and resembles a long, narrow ribbon''. In the case of
this larva Emery has also made the observation that if-
possesses a considerably greater number of vertebrae
than the full-grown fish, which generally has only 95
vertebras (26 abdominal), while in one larva he found
175 vertebras. In adult specimens of Fierasfer dentatus
too, the tip of the tail is often broken off short, and
bears a secondary growth, corresponding to a caudal
fin, such as we have above seen to be common among
the Macruroids. Emery was therefore quite justified
in his suspicion c that Fierasfer dentatus in later years
discards its shelter within the sea-cucumber, or at- all
events cannot gain entrance in the same manner as its
congener, although he subsequently^ met- with a small,
but full-grown specimen (with 115 vertebras) in a
Holotharia tubulosa.

Genus FIERASFER.

Pectoral fins present e. Branchiostegal rags 7. Pseud obranchice wanting. Branchiostegal membranes united under-
neath, but free from the isthmus. The upper jaw projects slightly beyond the lower. (Jar diform 1 teeth, in the
jaws , on the vomer , and on the palatine bones. Anal aperture nearly vertically below the insertions of the

pectoral fins. Pyloric appendages rudimentary .

This genus seems t-o attain the highest point of its
development in Indo-Aust-ralian Avafers, Avhere 5 or 6
species are knoAvn. One species has been described
from the east coast of Africa, one from Central Ame-
rica-, and t-wo from Europe. In 1853, during the voyage

round the world of the frigate Eugenie , Professor H.
Kinberg took near the Keeling Islands specimens of
Fierasfer parvipinnis 31 cm. in length and Avit-h a-
depth of 23 mm. just behind the head. The other
species of the genus are considerably smaller.

a In Kaup (Ann. Mag. Nat. Hist., ser. 3, vol. VI (1860), p. 272, tab. Ill, fig. D) this larva has borne the name of Porobranchus
linearis , in Gasco and subsequently in Costa (Anuario del Museo Zoologico di Napoli, Anno V, tav. I, fig. 1) that of Vexillifer de Filippi.

b Another characteristic, as yet observed only in the larva of Fierasfer acus, is that the tip of the tail ends, as in the Macruroid
larvae ( Krohnius ), in a long, filamentous prolongation.

0 Fn., FI. Golf. Neap. II, Monogr. Fierasf., p. 18.
d Mitt-h. Zool. Stat. Neap. Ill, p. 282.
e Though sometimes small.
f Sometimes set in one row in tire jaws.

OPHIDIOIDS.

601

FJERASFER DENTATUS.

Fig. 145.

Among the anterior jaw-teeth we find a pair or two of curved canines both in the upper jaw and the tower.
Length of the pectoral fins 1/2 that of the head , which in adult specimens measures about 101/2 % of that of the
body. Greatest depth of the body ( just behind the head) somewhat less than 1/2 (46 — 48 %) of the length of the
head , which is less than the distance between the vent and the tip of the snout ( during youth % t° °h> °ld
specimens 8/9 thereof). Longitudinal diameter of the orbits about V3 of the length of the lower jaw , which is
more than half ( about 56 %) of the length of the head, but less than half (about 45 %) of the distance between
the vent and the tip of the snout. Postorbital length of the head about 70 % of the entire length, of the same.
Height of the anal fin, which begins just behind the vent and somewhat behind, or vertically below the beginning
of the dorsal fin, greater at the middle of its length than either the height of the dorsal fin or the depth of the
body at this point, but somewhat less than the greatest depth of the body. The abdominal cavity extends behind
the vent for a distance about half as great again as that between the vent and the tip of the snout. Body trans-
parent, during life colourless , save for the whitish, brassy iris and the silvery abdominal cavity.

Fig. 145. Fierasfer dentatus, natural size. From 100 — 200 f thins, of water on the Jutland Reef.

Taken in 1880 by Fisherman B. Westergren.

E. hr. 7; I). ca 144 — 180; A. ca 165—180; P. 16; V. 0; C. 0.

a Syn. Ophidium dentatum, Cuv. (subgen. Fierasfer ), Regne Anirn.,
ed. 2, tome II, p. 359; Gtur ( Fierasfer ), Cat. Brit. Mus.,
Fish., vol. IV, p. 383; Couch, Fish . Brit. Isl., vol. Ill, p.
133, tab. CL VI; Putnam ( Eehiodon ), Proa. Boston Soc. Nat.
Hist., vol. XVI, p. 346; Gigl. ( Fierasfer ), Espos. Intern.
Pesoa, Berl. 1880, Sez. 1 1 al . , Cat., p. 97; Emery, Atti Accad.
Line., ser. Ill, Mem. Cl. Sc. Fis., Math., Natur. vol. VII,
p. 183, tab. I, cett. ; Ip., Fn., FI. Golf. Neapel, II, p. 16,
tab. I, cett.; Day, Fish. Gt. Brit., Irel., vol. I, p. 328;
Coll., Fork. Vid. Selsk. Christ. 1882, No. 19, p. 3 cum tab.;
Lillj., Sr., Norg. Fisk., vol. II, p. 234.

Eclnodon Drummondii, Thomps., Trans. Zool. Soc. Lond., vol.

II, p. 207, tab. XXXVIII.

Encheliophis tenuis, Putnam (larva) 1. c., p. 347.

In Scandinavia as everywhere Fierasfer dentatus
is an extremely rare fish ; and its habits, at least- when
adult, are almost unknown. It was originally described
as a Mediterranean form, but even there it is seldom ob-
served. Emery saw only one adult specimen at Naples,
but he mentions at least three larvae which he procured
from Holothurhe. Giglioli assigns this species to Naples,
Messina, and Venice. On the other hand, neither
Canestrini, Moreau, nor Steindachner mentions its
occurrence in the Mediterranean. In the Atlantic it
was first met with by Thompson, who received through
Drummond a specimen ‘28 cm. (11 in.) long that had

602

SCANDINAVIAN FISHES.

been found oil the south coast of Ireland lying dead
on the beach after a storm. Through professor S.
Loven the Royal Museum received in 1880 a specimen
that had been caught by Fisherman B. Westergren
“at a depth of from 100 to 200 fathoms on the Jut-
land Reef,” west of the Skaw. This specimen seems
to have been taken from the stomach of some larger
fish (probably a Cod or Ling) where it had some time
undergone the process of digestion, for the skin together
with all the fins and a great portion of the tail has
been digested, the hindmost of the remaining vertebrae
being quite bare. In its present condition the specimen
is 188 mm. long, the greatest depth of the body 11
mm., the length of the head 23 mm., the length of the
lower jaw 13 mm., the longitudinal diameter of the
orbit 4'5 mm., the length of the eye-ball 3‘6 mm., the
postorbital length of the head 16 mm., and the distance

between the tip of the snout and the vent 29 mm.
To judge by the length of the head the specimen, when
perfect, has been only slightly smaller than the one
described by Collett. The latter specimen was 225
mm. long; it was found in March, 1881, floating at the
surface off Rovaer, outside Stavanger Fjord, and for-
warded by Dr. Jensen to Bergen Museum. These two
specimens are up to the present the only finds of the
species within the limits of the Scandinavian fauna.
According to Couch (1. c.) the distinguished collector
and observer Thomas Edwards found 6 small examples
of this species, between 6 and 7 cm. long, in March,
1863, on a sandy bottom off the coast of Banff. They
moved through the water, he wrote, like Pipe-fishes, but
never attempted to hide among seaweed or under stones,
preferring to keep to the sand, where they would lie for
hours with the body in an undulating or curved position.

Fam. LYCODIDiE.

Body elongated , more or less anguilliform, in front terete, behind compressed, naked or covered with thin cycloid
scales. All the vertical fins confluent; no separate caudal fin. Jaws (. sometimes the palate as ivell) armed with
teeth. Gill-openings vertical and small, the branchiostegal membranes coalescing below with the shin and forming
a broad isthmus. Pseudobranchice distinct. Air-bladder wanting. Pyloric appendages usually wanting,

but sometimes present in a rudimentary form.

This family, which we here range last among the
Anacanthini, occupies a singular intermediate position
between other types. Artedi" and Linnaeus6 referred
the only species that they knew, the Eelpout or Vivi-
parous Blenny, to the genus Blennius; Gisler6, the
first to discover the occurrence of this species in Scan-
dinavia, regarded it as an intermediate form between
Blennius and Ophidion. Cuvier^ adhered to Artedi’s
opinion, and Reinhardt6, the first describer of the
types of the second division of the family, acknowledged
that Lycodes ought to be referred to Cuvier’s Mala-
copterygii thoracici, but still ranged the genus beside the
Blennies, on the grounds that it was a link between
Anarrhichas and Zoar evens ( Enchelyopus ), which latter

a Syn. Pise., p. 45.

b Fn. Suec., ed. II, p. 113.

c Vet.-Akad. Hand!. 1748, p. 42.

d R'egne Animal , ed. II, tome II, p. 240.

e Danske Vid. Selsk. Naturv., Math. Afh., Dee] 7, p. 147.

t Abh. Akad. Wiss. Berlin 1844, p. 165.

,J Sveriges och Norges fislcar, part II, p. 17.

genus “evidently was an offshoot of the Gadoid family.”
The same reasoning recurs in Muller7, who united the
Lycodidce to the Blenniidce on account of the pseudo-
branchite. Lilljeborg9, has grouped together the osteo-
logical characters that separate the Lycodidce from the
Gadidce. We notice in particular the absence in the
former of the two characteristics of the latter that we
have mentioned above, namely the great extent of the
styloid bone (a character which is also wanting, how-
ever, in the Sand-Eels) and the lobate process on the
hind part of the upper margin of the intermaxillary
bones (which is also wanting in the Ophidioids). By
the majority of their characters, however, the Lycodoids
are joined to the other Onomorphi as Malacopterygian

LYCODOIDS.

603

fishes (cf. above, p. 211): they seem to be especially
closely allied to an Australian genus among the Ophi-
dioids" that has received of Gunther6 the name of
Gongrogadus and resembles the Lycodoids in the form

of the body, the form of the fins, and the absence of
the air-bladder.

Gilt/ was the first to give the family its present
definition. About 20 or 30 species are recognised and
distributed among 8 genera.

Genus ENCHELYOPUS.

Dorsal fin f urnished behind with a depression , the rays within which are short and, spinous. Ventral fins jugular
and rudimentary, with 3 or 4 rays. Palatine and vomerine teeth wanting.

The genus of the Eelpouts, known in its European
species since the time of Schonevelde (1624), first
excited attention on account of its reproducing itself
by the birth of living, viable young. It is equally re-
markable, however, for the development of spinous rays
at so unusual a spot, corresponding to the posterior part
of the dorsal fin in the Sea-cat (cf. above, p. 231) or
to the upper margin of the caudal fin in other fishes,
a spot where spinous rays also occur.

The oldest post-Linnaean generic name, Enchely-
opusd , was adopted by Gronovius in 1763e from Klein,

in whose works it contained a most heterogeneous
medley of fishes. The name given by Gronovius was
however forgotten, though Valenciennes gave a refe-
rence to it/, until it was restored in 1863 by Gill5'.
The genus has been best known by the name of Zo-
arcesh , which was given it in 1829 by Cuvier* 1, or
Zoarcceus, as Nilsson tvrote the word/

The genus belongs exclusively to the seas of the
Northern Hemisphere and contains only two species,
which are very closely related to each other, one from
Europe and one from North America.

THE EELPOUT (s\v. tanglaken).
ENCHELYOPUS VIVIPARUS.

Pi. XII, fig. l.

Bays of the anal fin less than 100. Body slimy and covered with thin, oblong, rounded, cycloid scales, de-
pressed in the skin. Coloration greenish yellow or yellowish brown (lighter on the belly and the under surface
of the head) marked across the dorsal fin and the back with dark brown transverse bands, which are more or less
distinctly branched below and meet or alternate with a row of similarly coloured spots on the lower part of the sides.

R. hr. 6; D. (72— 80) + (6—10) + (16— 21); A. 80—88/
P. 19; V. 3; Vert. 108—111.

Syn. Tertia Mustelarum species, Schonev., Ichthyol. Slesv. Hols.,
p. 49; Mustela Lumpen Antverpice dicta, Willugb., Hist.
Pise., p. 120; Art., {Mustela), Gen. Pise., Append., p. 83;
( Blennius ), Syn. Pise., p. 45; Gronov., J. F. ( Blennius ),
Act. Upsal. 1742, p. 87; Lin., It. Wyoth., p. 182; Gisl.,
L c. ; Gronov., L. T., Mus. Ichthyol., p. 65.

Blennius viviparus, Lin., Mus. Ad. Frid., p. 69, tab. 32, fig.
3; Syst. Nat., ed. X, tom. I, p. 258; Fa. Suec., ed. II, p.
113; Mull., Zool. Dan., tab. LVII; Retz., Fn., Suec. Lin.,
p. 325; Cuv. ( Zoarces ), L c. ; Nilss. ( Zoarcceus ), 1. c.; Ekstr.,
Vet.-Akad. Handl. 1834, p. 48; Flmng ( Gunnellus ), Brit.
Anim., p. 207; Cuv., Val. {Zoarces), Hist. Nat. Poiss., vol.
XI, p. 454; Fit., Ekstr., v. We. {Zoarcceus), Skand. Fisk.,
ed. I, p. 36, tab. 8, fig. 1; Parn. {Zoarces), Mem. Wern.
Soc., vol. VII, p. 337; Kr., Damn. Fiske, vol. 1, p. 355;

“ Machcerium, Richardson, Voy. Ereb., Terr., Ichthyology, p. 72, tab. 44, figs. 1 — 6.
b Cat. Brit. Mus., Fish., vol. IV, p. 388; Intr. Stud. Fish., p. 550.
c Proc. Acad. Nat. Sc. Pliilad. 1862, p. 501 and 1863, p. 254.
d Eel-like, from I'yyfiAvg and cop.

e Zoophyl., p. 77. As early as 1760 (Act. Helvet., IV, p. 259) Gronovius applied it to the Eelpout, but then without characterizing
the species.

I Cov., Val., Hist. Nat. Poiss., vol. XI, p. 452.

9 Proc. Acad. Nat. Sc. Philad. 1863, p. 256.
h ZcociQyfig, life-preserving.

1 Regne Anim., 1. c.
i Prodr. Ichth. Scctnd. (1832), p. 104.
k Sometimes 89, according to Moreau.

604

SCANDINAVIAN FISHES.

Nilss. ( ZoarccBus ), Skand. Fn., Fish., p. 203; Sund., Stockh.
L. Hush. Sallsk. Hand]., H. 6 (1855) p. 80; Gthr ( Zoarces ),
Cat. Brit. Mus., Fisli., vol. Ill, p. 295; Mgrn, Finl. Fiskfn.
(disp. Helsingf. 1863), p. 21; Lindstr. ( Zoarcceus ), Gotl.
Fish., Got]. L. Hush. Sallsk. Arsber. 1866, p. 15 (sep.);
Steind., Stzber. Akad. Wiss. Wien. Naturw. Math. Cl. LVII, i
(1868) p. 676; Coll. (Zoarces), Forh. Vid. Selsk. Christ.
1874, Tilhegsh., p. 78; Malm, Gbgs, Boll. Fn., p. 473;
Winth., Naturh. Tidskr. Kbhvn, ser. 3, vol. XII. p. 23;
Bncke, Fisch., Fischer., Fischz. 0., W. Breuss., p. 80;
Day, Fish. Gt. Brit., Irel ., vol. I, p. 211, tab. LXI, fig. 2;
Mor., Hist. Nat. Poiss. Fr., vol. II, p. 156; Mela, Vert.
Fenn ., p. 292, tab. IX; Mob., Hcke, Fisch. Osts., p. 61;
Lillj., S'v., Norg. Fish., vol. I, p. 552; Hansen, Zool. Dan.,
Fishe, p. 49, tab VIII, fig. 2; Bncke, Handb. Fischz.,
Fischer. (M. v. u. Borne) p. 92.

Blennius Dump mus, Lin., Syst. Nat., 1. c., — ex Art. et

WlLLUGHB.

Obs. The only constant character we have been able to discover,
to distinguish between the European and American Eelpouts, lies in

the number of the fin-rays. This character is evidently connected with
a greater elongation of the hind part of the body in the American
form (Enchelyopus anguillaris ), especially as this form appears in a
specimen 40 cm. long which the Boyal Museum has received through
the Smithsonian Institution. Todd’s figure in Brown-Goode (rFhe Fishe-
ries and Fishery Industries of the United States, Section I, Plate 67)
shows, however, that in this respect the variations may be great in
both forms. Even among our Scandinavian Eelpouts it seems to be
possible to distinguish between two forms. The first of these forms
by the more slender form of the body represents the most advanced
development of the male characters, like the American species, has the
beginning of the dorsal fin situated further forward (its distance from
the tip of the snout less than 16 % of the length of the body), and
in several other respects shows traces of the persistency of the juve-
nile characters. This form occurs, according to the collections of the
Royal Museum, on the coast of Finmark and otf Kola Peninsula. In the
second form, a southern one, which the Museum possesses both from
Bohuslan and the Baltic, the region of the anal fin is generally shor-
ter, the head larger, and the beginning of the dorsal fin situated
further back. For the sake of brevity we refer the reader to the
following table of averages.

Length of the body expressed in millimetres

Distance between the dorsal tin and the tip of the snout in % of the length of the body

„ ,, „ „ „ ,, ,, ,, „ „ „ „ „ „ ,, distance between the beginning of the

anal tin and the tip of the caudal tin

Postorbital length of the head in % of the length of the body

,, „ „ „ ,, ,, „ „ „ distance between the beginning of the anal fin and the tip of

the caudal fin

Distance between the beginning of the anal fin and the tip of the caudal fin in % of the length of the body

A 3 ts

399

14.1

21.1

9.5

14.2
67.o

Average in

5. c
5'

e

E ^ rz
3 5“

a

99

14.2

21.8
9 3

14.4

64.9

O' Pf «
9 2-3'
a 5s re

qb 4
- a |

1 s-*

0 w a

c" e

164

15.i

24.4
9.0

15.5
62.3

l A
16.7

26. g

10.3

16.3
63.o

BBS.
£3- O a
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The ordinary length of our common Eelpout is
about 30 cm., but we have received specimens from the
island-belt of Stockholm that were nearly 40 cm. long.
Further south the Eelpout seems to attain a greater
size; Parnell mentions specimens taken off Berwick
(at the mouth of the Tweed) that were nearly 2 ft.
(61 cm.) in length". The females attain a greater size
than the males.

In the form of the body the Eelpout is most like
the common Burbot. The depth of the body at the
beginning of the anal tin in young specimens (less than
1 dm. long) measures about 87s' — 97s 0//° °f the length
of the body, in older specimens about 97 2 — 1 1 1/2 %
thereof. The head is small, its length in adult Eelpouts

being about 18 or 19 % of that of the body. Under-
neath it is flat, above rounded, with tumid cheeks and
with forehead somewhat compressed laterally and slop-
ing in a steep curve towards the snout. The mouth is
of moderate size, with rounded jaws and very tumid
lips, which are wrapped round the jaw-bones and entirely
cover them. The upper jaw is protrusile to some ex-
tent, though only slightly. Both the jaws are of the
same length and furnished with conical, small but strong
teeth, with rather blunt tip''. I11 the upper jaw these
teeth are set in two rows, the anterior (outer) fairly
regular and containing about 12 teeth on each of the
intermaxillary bones, the posterior (inner) less regular,
with shorter teeth, and ending usually at the root of

Amer.

a The American form sometimes attains a length of 372 (107 cm.) and a weight of 12 lbs. (5 kgm.), according to Storer (Mem.

Acad. Arts, Sc., N. Ser., vol. V, p. 264).

h The teeth of the American Enchelyopus are green, coarser, and still blunter, reminding us strongly of those of the Sea-cat.

EELPOUT.

605

the 7th or 8th tooth in the anterior row. In the lower
jaw the form of the teeth is the same, and they
are set. in almost the same manner; but the posterior
row is somewhat longer and ends only a. little sooner
than the anterior row. There are no teeth on the pa-
late or the tongue, which is flat, rounded, and fleshy,
but scarcely free, at all. 1 he palatal folds within the
jaws are well-developed, but not at all deep. In the
pharynx we find above on each side three small, trans-
versely-set pharyngeals, set with short, conical teeth in
more or less curved rows, convex in front, the middle
pair of pharyngeals with two rows of teeth, the anterior"
and the posterior pairs with only one. The lower pha-
ryngeals are longer, pointed at both ends, and furnished
with three longitudinal rows of short and blunt teeth.
The gill-rakers are scattered — 16 on each of the first,
pair of branchial arches — and resemble short, pointed
papillae, each with a small, sharp, osseous spine within
it. The branchiostegal membrane is united underneath
to the corresponding membrane on the other side, but
coalesces entirely with the skin of the isthmus (the
breast), leaving no free margin behind, though the
margin may be distinguished. The gill-openings extend
downward somewhat beyond the lower end of the in-
sertions of the pectoral fins and are separated from it.
by a distance about equal to the length of the snout;
while above they reach over a part of the upper margin
of the operculum, the skin of which is prolonged behind
into a blunt point. The eyes are comparatively small,
rather prominent, and set in front of the middle of the
head and high up, the distance between them being
about equal to the breadth of either eye. They lie in
planes that converge somewhat in a forward and upward
direction. The nostrils, one on each side, are set. as in
the Sea-cat at. about the middle of the length of the
snout, and resemble fairly high, flexible, dermal tubes
with even margin. The eye is surrounded as usual by
a ring of muciferous pores, belonging to the cephalic
system of the lateral line. One of these pores, the
second in the frontorostral branch of this system of
ducts, has been regarded as a posterior nostril. The
first pore of this branch lies just in front of the nostril.
Through both these pores a fine hair may be passed
into the duct beneath the skin, far back and above the
eyes. The lower jaw is also coasted by a row of si-
milar and distinct pores.

The body proper (the trunk) is cylindrical, with
pendent bell}’, especially in gravid females. The Eel-
like tail, which ends in a point, is strongly compressed
laterally. During life the fish is coated with a thick
layer of mucus, partly concealing the thin, scattered
scales, which lie depressed in the skin without touching
each other. These scales are almost circular or oblong,
and smooth-margined, with fine, concentric striae, in
the outer part dense, in the inner part, (nearer the
nucleus) scattered, and grooves radiating in all direc-
tions. The lateral line is not very distinct. The head
is scaleless.

The vent lies just in front of the beginning of the
anal fin, a little in front of the middle of the body
(cf. the above table), and further forward in young
specimens, as in the American form, than in old. Be-
hind it. is furnished with a papilla, most distinct in
the males, similar to that we have above remarked in
the Cottoids.

The dorsal fin is low, begins just behind the occi-
put. (cf. the above table), and extends, imperceptibly
diminishing in height, back to the extreme end of the
tail; but a little in front of this point it shows a more
or less broad depression or incision, which is supported
by spinous rays, an extremely singular variation of the
other soft rays, and characteristic of this genus. The
number of these spinous rays varies between 6 and 10".
In front of the depression the dorsal fin contains about
80 rays, all weak and articulated, and all, with the
exception of the first rays, branched at the tip. These
rays are united by a fairly thick, slimy skin, which
is strewn at the base of the fin with scales similar to
those which clothe the body. These scales extend
higher up in the posterior part of the fin than in the
anterior. The anal fin is of exactly the same structure,
form, and extent in a backward direction as the dorsal;
but is without, any incision, coalescing completely and
without the least break with the caudal fin. At the
base the anal fin is strewn with scales in the same
manner as the dorsal. From 70 to 80 rays may be
counted with tolerable ease; but behind these the re-
maining rays lie so close to each other that, their num-
ber can hardly be fixed with certainty.

The pectoral fins are fairly large — their length
from the upper angle of the fin varying between 1 1
and 13 % of that of the body — and broad, with rounded

a Ekstrom and Fries have seen two rows of teeth on the anterior pair as well.

6 In the American Enchelyopus anguillaris their number may rise to at least 16

606

SCANDINAVIAN FISHES.

tip. They are made up of 19 branched rays and a
thick membrane. The ventral fins are rather small
and lie close to each other, in front of the pectoral
tins. They contain 3 rays, which ai*e branched at the
tip and very difficult to distinguish.

The only external difference between the sexes
that we have been able to discover, is that the males
are generally smaller and more slender than the females.

The coloration of the Eelpont is not very diver-
sified. The ground-colour of the entire fish is yellowish
brown, the under surface of the head and belly being
grayish yellow. On the sides of the body, back to the
depression in the dorsal fin, we find two rows of more
or less distinct transverse spots, 13 — 15 in each row.
The spots of the lower row alternate more or less re-
gularly with those of the upper and at the upper cor-
ner touch the latter, which advance some way over the
dorsal fin. Behind the incision in this fin the spots
grow gradually less distinct towards the end of the tail.
The outer rim of the dorsal fin is black. The outer
margin of the anal fin is flame-yellow, like the tips of
the ventral fins. The pectoral fins are darker than the
others and edged with more dirty yellow. Several
blackish spots occur on the upper part of the head,
between the eyes and the tip of the snout, and on the
cheeks, growing rather more indistinct on the gill-cover.
The iris is dark brown, with a fine, light yellow ring
round the pupil.

The internal organs remind us more of the Cotti
than of the Cods. The oesophagus and stomach are
short, the bottom of the latter extending hardly beyond
a line with the middle of the pectoral fins when folded,
and bending downwards to form a short pyloric part,
turned in a forward direction, at the end of which we
find two short, saccate bulbs or rudimentary pyloric
appendages, one on each side. At the middle of its
length the intestine forms a winding bend, a curved
double coil. At first sight the liver seems to consist
of one single lobe, which curves upwards and surrounds
the small gall-bladder, the stomach, and the oesophagus;
but this lobe is divided distinctly enough into three
parts, with the middle (outermost) lobe shortest and the
right lobe longest. The spleen lies just behind the
bottom of the stomach and is small and flat, at the
upper (dorsal and also left) end pointed, at the lower
end rounded. The air-bladder is wanting. The testes

of the male are smooth and oblong, lie close to each
other, and at the spawning season grow far forward,
close to the kidneys. The most remarkable of the in-
ternal organs of the Eelpout is the ovary of the female.
On opening a female which is not gravid we find the
ovary, which is simple and of oval form, lying right
under the kidneys and along the spinal column. It is
of thin texture and partly filled with yellow eggs of
the size of a pin’s head. On examining the ovary after
the beginning of the period of gestation, when the ovary
serves as a uterus, we find the walls firmer and pene-
trated by a great number of bloodvessels, while to the
inner surface are attached small, oblong, transparent
vesicles, which contain a clear fluid and a tiny embryo
floating in the fluid. At the last stage we find the sac
enormously distended and full of closely packed young
specimens lying free within the ovary itself. The ve-
sicles have collapsed, but are still attached to the walls.
During this period we generally find a great number
of the young expelled into the abdominal cavity itself
by the bursting of the ovary. In a female 31 cm. long
we have found altogether 196 young, each 37 mm. in
length. Seventy-five of them lay in the abdominal
cavity, the rest within the ovary. In another female,
322 mm. in length, we have counted 262 young. Other
writers state the number of the young at over 300.

The Eelpout seems to arrive at maturity early, for
distinct eggs have been found in specimens only 15 cm.
long. Its faculty of giving birth to living young pre-
supposes an actual copulation between the sexes, as well
as the fertilization of the eggs within the body of the
mother. However, no such copulation has yet been ob-
served; nor caii any certain time of year positively be
given as the spawning- season of the Eelpout, for at
almost all seasons we may find gravid and parturient
females and at the same time others that contain only
small eggs. The former are most often met with, how-
ever, in December and January. This agrees with
Benecke’s statement (1. c.) that during the spring and
summer months — from March to August — the males
have bright orange fins and wear a kind of festal dress.

Brehm has observed in a salt-water aquarium the
manner in which the Eelpout gives birth to its young.
“The fish,” he says", “which even at other times is slug-
gish, seeks a certain spot in the aquarium several hours
before parturition and stays there motionless, until all

Bremh’s Thierleben , 2:te Aufl., Die Fische , p. 138.

EELPOUT.

607

or at least most of the young have been excluded. The
fry creep forth, head first, one after another, and sink
down to the right and left of the tail of the mother-
fish, which she keeps in a somewhat elevated position.
Now that they have reached the bottom, they lie there
several hours, perhaps a whole day, without perceptibly
moving or rising. If there are several Eelpouts in the
same aquarium, one may observe, to one’s surprise,
two or more of them swim (dose up to the mother-fish
and press her on both sides, apparently to assist the
operation, but in reality only to devour the young as
soon as they appear. The mother does not hesitate to
folloAv their example if she is not sufficiently supplied
with other food. In most cases the Eel pout brings forth
all her young at a birth; but it sometimes happens
that only a part of them are born at first and the rest,
or even only some of the rest, one or more days later".

The food of the Eelpout consists chiefly of mol-
lusks, crustaceans, and worms, but also of small fishes.
It lives in water of moderate depth on shores where it
finds a stony bottom overgrown with seaweed. Hence
it is known as Tdnglake or Stenlake {tcnig, tang, Jake,
Burbot), which are its ordinary names on the coast of
the Baltic, where its outward resemblance to the Burbot
has thus attracted most attention. In other localities
its slight likeness to the Eel and the long-known fact
that, unlike most of the Teleosts, it brings forth its
young alive, have given rise to the ancient and popular
belief that the Eelpout is the mother of the Eel, and

have bestowed upon it various names. Thus, in German
it is still called Aalmutter, in Danish Aalekone, Aale-
moder , or Aalekus, and in Bohuslan generally Alkussa
or simply Kussa. Its manner of life has also contri-
buted in a high degree to its comparison with the Eel.
It lives in scattered and solitary specimens, is found
everywhere, but seldom in numbers, and conceals itself
under stones, among seaweed, and in crevices in the
bottom. All round the coasts of Scandinavia it is com-
mon, from Varan ger Fjord" along Norway and Sweden
into the Baltic and up to the island-belt of Tornea.
On the Danish coast and on all the coasts and banks
of the North Sea the Eelpout is a common fish; but
west and south of the English Channel it is rare,
though Steindacbner states (1. c.) that it has been met
with in the neighbourhood of Cadiz. The Eelpout also
makes its way occasionally into fresh water. It has
been taken for instance, according to Brehm, in the
Havel off Spandau.

The flesh of the Eelpout is firm, white, and in
flavour not unlike that of the Eel. Still it is not eaten
in many places, a circumstance which seems to be due
entirely or at least most usually to the green colour
of the bones, which becomes deeper and deeper during
the process of boiling and excites the groundless suspi-
cion that the flesh is poisonous. There is no special
fishery for the Eelpout: it is taken only by accident
and generally while the fisherman is drawing the seine
for other fish. (Ekstrom, Fries, Smitt.)

Genus LYCQDES.

Vertical fins similar in structure throughout their length. Ventral fins jugular and rudimentary, with 2 — 6b rays.

Jaws, palatine hones, and the head of the vomer furnished with teeth.

This genus, which was founded in 1831 by Y.
Reinhardt0, is extremely closely allied to the preceding
one in form and the changes of form as well as in the
distribution of colour on the body. Externally the
absence of the depression in the margin of the dor-

sal fin is a readily perceivable difference from the Eel-
pouts, but both genera are identical in all other respects,
if we except the more back ward beginning of the dorsal
fin in Lycodes, where the distance between the begin-
ning of this fin and the tip of the snout is always

a The Royal Museum has received from the White Sea through Lieutenant LI. Sandeberg an Eelpout 19 cm. long which is remarkable
for the extraordinary height of the anterior part of the dorsal fin.

6 The latter number in Lycodes macvops , according to Vaillant, who, however, describes these rays as simple ( Exped . Scient. du Tra-
vailleur et du Talisman, Poiss., p. 30 7), wh ile in other species they are known to be branched.

c Overs. Vid. Selsk. Forh. Kb/tvn 1830 — 31, p. LXXIV. It was not until 1838 — Nature., Math. Afli., Deel VII, p. 147, published
by the same society — that the genus was completely determined and characterized. The name of Lycodes (wolflike, Gr. Lv'/.og) is derived
from the resemblance of the fish to Anavrliiclias lupus.

7 1

Scan cl in a v i a n Fis lies.

608

SCANDINAVIAN FISHES.

perceptibly more than half (at least 55 %) of the distance
between the anal fin and the latter.

The changes caused by age in the coloration of
Lycodes are extraordinarily great. All the fry — with
the exception of one species — are adorned with a hand-
some, regular marking (see for example tigs. 148 and
150) of selliform, dark-margined spots across the dorsal
lin and the back. On the hindmost part of the tail
these spots are generally prolonged downwards, forming
transverse bands across the dorsal fin, the body, and
the anal fin. The spaces between the spots grow lighter
and lighter, being sometimes milk-white on the dorsal
fin and the back, and the same light colour sometimes
appears in the form of an ocellated spot within one or
another of the selliform spots. With age, however,
these selliform spots coalesce below, and the boundary
between them and the coloration of the rest of the
body is effaced, being sometimes replaced by a net-
work of darker colour, starting from the original dark-
coloured margin of the spots. These changes of colo-
ration are also common to other kindred genera within
the family, and strongly remind us of the distribution
of the spots in the Eelpouts, in all its irregularity.

The dentition of the palatine bones and of the head
of the vomer in Lycodes — a character which is wanting
in the Eelpouts — is here counterbalanced by a nega-
tive character, the absence of transverse palatal folds
behind the rows of teeth in the jaws. The nostrils are
simple and tubular in this genus also, but the snout
is more elongated and of a looser structure, this being
due to the still greater development of the cavities be-
longing to the cephalic system of the lateral line. The
extension of this system over the body is singular
enough. No less than three lateral lines may appear
on each side of the body — but, as far as we know,
no more than two in the same individual. These lines
are generally wanting in young specimens and are again
effaced in old. In most cases two lateral lines start

from each temporal region, one — corresponding to the
ordinary mediolateral line ■ — in a slight curve down-
wards to the middle of the side, or even down to the
anal region and then along the base of the anal fin,
the other — a dorsal line — usually with larger but
more scattered pores and seldom extending farther than
to a line with the vent or a little behind it. Sometimes
again a ventral branch starts from the anterior part of
the mediolateral line, its structure being' the same as
that of the latter. Lutken, who was the first" to draw
attention to all these differences, based upon them a
system of determining the species within the genus;
but Gunther has pointed out-6 the systematic difficulties
involved in a strict adherence to characters derived
from this relation, characters which at different ages
and in different individuals show variations, the cause
of which is as yet unknown, and which also in many
cases defy observation. Gunther passes the same judg-
ment upon the characters which have been drawn from
the extent of the scales in these fishes.

Ever since Richardson described his Lycodes rnu-
cosus from Northumberland Sound", we have known
that scales may be wanting within this genus in indi-
viduals up to a length of 28 cm. In 1874<z Sleeker
founded the genus Lycodalepis to emphasise this cha-
racter, which, according to Bean", may be persistent-
in the species just mentioned even at a length of 43
cm. However, to the best of our judgment and with
the knowledge we now possess of the changes of growth
and sexual differences within the genus, this species
can scarcely be distinguished, except perhaps as a local
variety, from Lycodes Turneri, the form described by
Bean7 and Turner5' from Alaska, or from the form
brought home by the Vega Expedition6 from the en-
trance of Chatanga Bay (Siberia, E. of Cape Tschelju-
skin). The latter form is in all probability identical
with the form subsequently described by Lutken' under
Collett’s name of Lycodes Lutkenii, which Gunther

a Vid. Meddel. Naturh. For. Kbhvn 1879 — 80, p. 329. Kroyer, however, had already remarked a dorsal and a mediolateral line in
his Lycodes perspicillum (Naturh. Tidskr. Kbhvn, ser. Ill, vol. I, (1862), p. 292).
b Deep Sea Fish., Cliall. Exped., p. 79.
c The Last of the Arctic Voyages, vol. II, p. 362.
d Vers]., Med. Akad. Wet. Amster., 2 R., 8 Del, p. 369.
c Bull. U. S. Nat. Mus., No. 15, p. 113.
f Proc. U. S. Nat. Mus., vol. I, p. 463.

■' Contrib. Nat. Hist. Alaska , Arct. Ser. Publ. Sign. Serv. U. S. Army, No. II, p. 93, PI. 4.
h Gr. Intern. Fisher. Exhib. London 1883, Swed. Catal., p. 176.

' Kara-Havets Fiske , Dijmphna-Togtets Zool.-bot. Udb., p. 14, Tab. XVI. In the largest of these specimens Lutken remarked a
considerably greater number (13 — 15) of palatine teeth than in Lycodes mucosus (7, both according to Richardson’s figure and in the speci-
men from Chatanga Bay). But the great resemblance in other respects between these nominal species forces us to the conclusion that this
difference is individual and may depend on circumstances of age or sex, unless indeed it belongs to a local variety.

LYCODO IDS.

had unhesitatingly referred to the old Rein hard t.ian
species Lycodes reticulatus. This form, which belongs
to Kara Sea, is furnished when 225 mm. long, accord-
ing to Luticen, with scales over the whole tail and the
greater part of the back behind the beginning of the
dorsal tin; but the above-mentioned specimen from
Chatanga Bay, which we proposed in 1882 to refer to
Lycodes , mucosus, is entirely scaleless at a. length of
24(1 mm. The individual variations in this respect
seem to be so great that we must abandon the attempt
to base the determination of the species upon the pre-
sence or absence of scales. No more reliance can be
placed on the proposed determination — here as in the
Gobies — based on the greater or less extent of the
scaly covering in a forward direction, over the trunk
and head, in those forms which are furnished with
scales at an early age and retain them at all sub-
sequent ages.

The differentiation of the species within this genus
from each other is a more advanced development of the
specific difference which seems to be present, but can
scarcely be regarded as fixed, within the preceding
genus. A trace of the most essential difference between
the European and the American Eelpout may be found,
as we have seen above, in the two forms that can be
distinguished with tolerable ease among the European
Eelpouts, and may be reduced to a greater or less
elongation of the caudal part of the body, in combina-
tion with a greater or less number of vertebra?. So
also within the genus Lycodes we can distinguish be-
tween two main groups of forms, the first with com-
paratively shorter body, like that of the Eelpout or
even deeper than it, the second with exceedingly elong-
ated, more Anguilliform body. Within these groups
the most general specific difference lies in the same
relation, conjoined with the varying size of the head,
originally a distinction of sex and age, for the relative

609

length of the head generally increases with age and is
always greater in the males than in the females.

The genus is composed exclusively of deep-sea
fishes, most of them living at depths of from 300 to
500 fathoms or more, and at least one species taken
in 1,333 fathoms of water. During youth, however,
they are met with nearer land and in shallower water,
sometimes at a depth of only 50 or 30 fathoms. For
the scientific elucidation of these forms we have prin-
cipally to thank Collett and Lutken. To the gene-
rality of fishermen the genus is unknown, and it has
no popular name; Lilljeborg has proposed the Swedish
names of dlbrosmar or dllubbar (Eel-Torsks).

The species that belong to the Scandinavian fauna
may be distinguished in the following manner:

A: Depth of the body at the beginning of
the anal fin more than 8 % of its length.

Branchiostegal rays 6.

a: Body plain, without spots Lycodes frigidus.

b: Coloration light with dark-edged, sel-
liform spots in the fry, in adult spe-
cimens dark with light transverse
spots or light with dark transverse
spots.

a: Length of the head more than
39 % of that of the tail from the
beginning of the anal fin to the

tip of the last fin-rays Lycode, s reticulatus.

fj : Length of the head less than 39
% of that of the tail from the
beginning of the anal fin to the

tip of the last fin-rays Lycodes Valdii.

B: Depth of the body at the beginning of
.the anal fin less than 8 % of its length.

Branchiostegal rays 5.
a: Length of the head at least about
*/4 (more than 23 %) of that of the
tail from the beginning of the anal
fin to the tip of the last fin-rays... Lycodes Sarsii.
fj: Length of the head at most about
'/- (scarcely more than 20%) of that
of the tail from the beginning of the
anal fin to the tip of the last fin-rays Lycodes murcena.

610

SCANDINAVIAN FISHES.

LYCODES FRIGID LIS.

Fig. 146.

Coloration plain, reddish gray-brown. Body in quite full-grown specimens entirely covered with scales forward
to the head and a little way over the bases of the vertical fins, in younger specimens naked on the middle of
the belly, on the fins, and on a strip along their bases. Depth of the body at the beginning of the anal fin in
adult specimens 10 V2 — 11 % of the length of the body. One ventro-lateral line, continuous and with dense pores;
one dorso-lateral line, with scattered pores ( both often indistinct). Head depressed ( flattened ) in front; its length
more than V2 the distance between the anal fin and the tip of the snout, and in quite full-grown specimens ( more
than 3 dm. in length) 42 — 47 %, in younger ones 35 — 42 %, of the length of the tail from the beginning of

the anal fin to the end of the last fin-rays.

Fig. 146. Lycodes frigidus , from the Norwegian Arctic Expedition. the natural size.
•The property of the Museum of Christiania University.

R. br. 6; D. 99—104; A. 85—90; P. 20—21; V. 2 (4?).
Syn. Lycodes Vahlii, Coll., Forh. Vid. Selsk. Christ. 1878. No. 4.

p. 11.

Lycodes frigidus , Id., ibid., No. 14, p. 45; Norsk. Nordh.

Exped., Fislce, p. 96, tab. Ill, figg. 23 et 24; Lillj., Sv.,

Norg. Fisk., vol. II, p. 19; Gthr, Deep Sea Fish., Challeng.

Exped., part. LVII, p. 79.

This species is known almost exclusively from the
collections of the Norwegian Arctic Expedition and
Collett’s description of them. It is said to attain a
length of almost 51 cm. Its principal characteristic
lies in the coloration, for in other respects it partly
coincides with Lycodes reticulatus and partly ranks as
an intermediate form between the latter and Lycodes
Vahlii. To judge from our knowledge of the other

species only male specimens, sterile or with the organs
of generation at rest, have been found. In the other
forms of this group of the genus the length of the
head in the males is greater, in the females less, than
half the distance between the anal tin and the tip of
the snout. Our knowledge of Lycodes frigidus does
not, therefore, preclude the possibility that by this
specific name are designated a number of sterile and
perhaps hybrid individuals of one or other of the two
following species. Among the fifteen more or less
adult individuals described by Collett all the younger
ones, with the exception of the youngest (118 mm.
long), are identical, according to the measurements giv-
en, in their most essential characters with Lycodes
Vahlii ; but the older specimens (more than 3 dm. long)

l

LYCODOIIXS.

(11

with L. reticulatus'1. It also seems no easy task to
distinguish from L. frig id us the specimen 56 cm. long
that Gunther (1. c., p. 77 and Plate XIII) has referred
to the last-mentioned species.

On the coast of Scandinavia only three specimens,
all young, have been found. At first two specimens,
respectively 37 and 62 mm. in length, were taken in
the month of June by the Norwegian Expedition of
1887, on the banks off Helgeland and Lofoden, at depths
of 350 and 457 fathoms and on a clayey bottom. A
third specimen, 148 mm. in length, which is referred
by the coloration to this species, but in other respects
entirely agrees with the young of Lycodes Vahlii, was
taken in 1889 by Dr. Carl Aurivillius, in between 10
and 40 fathoms of water, on a clayey bottom at Orsta
and Borsas in the inner part of Gullmar Fjord (Bohuslan).
The other known specimens have been found in 260 —
1,333 fathoms of water, off the extreme north of Nor-

way, Bear Island, and Spitzbergen. Collett gives a
list of a number of crustaceans and some parts of a
cuttle-fish as having been found in the stomach of this
species, and remarks that several of the former belonged
to such species as generally live in the surface-regions
of the open sea, but have thus been shown also to in-
habit the deepest abysses of the ocean. All the spe-
cimens taken by the Norwegian Expedition, except one,
were caught in ice-cold water; and it was this circum-
stance that suggested to Collett the name of frigidus.
The expedition kept some of them alive for a, time in
a vessel on board. These specimens were extremely
sluggish, preferring to lie still at the bottom of the
vessel and in a bent position, like that usually assumed
by the Eelpout and the Sea-snails. When thrown
into alcohol, however, they became extremely violent
and proved, like most bottom-fishes, to be extraordi-
narily tenacious of life.

LYCODES RETICULATUS.

Figs. 147 and 148.

Coloration in the fry more or less light , brownish yellow , with selliform spots on the trunk and tail and a scuti-
form figure, containing two light, fronted, spots , on the top of the head, in older specimens orange or paler, with
light or even white transverse bands between the darker ones on the trunk, and tail and light or even 'white spots
on the top of the head, in the oldest specimens of a more undiversified grayish brown , until the traces of the dark
transverse bands on the sides and cdso the head, crossed by a network of black streaks. The covering of scales
appears at different ages and occupies different extents of the body, but 'when complete extends over the greater
part of the trunk and tail, and of the base of the dorsal fin, leaving the head, the belly, the base of the anal fin,
and the anterior part of the base of the dorsal fin naked. One medio-lateral and one dor so -lateral line. ( both often
indistinct). Depth of the body at the beginning of the anal fin about 9 — 14 % of its own length. Head depressed
in front: its length more than 39 % (as a rule 42 — • 58 %) of the length of the tail from the beginning of the

anal fin to the tip of the last fin-rays.

R. hr. 6; D. (80) 91—95; .4. (63) 70—78; P. (16) 19—23;
V. 4; Vert. 95—97 (= 23— 25 -Kr).

Syn. Lycodes reticulatus, Rhdt, D. Vid. Seisk. Naturv., Math. Afh.

Kbhvn, vol. 7, p. 167, tab. VI; Gthr, Cat. Brit. Mas., Fish.,

vol. IV, p. 320; Coll., Forh. Vid. Seisk. Christ. 1878, No.
14, p. 59; Ltkn, Vid. Meddel. Naturli. For. Kbhvn 1879 — -
1880, pp. 318 et 330; Gthr, Deep 6'ea Fish., Chalk Exped.,
part. LVII (Zook, vol. XXII), p. 77, tab. XIII; Ltkn, Dijmphna-
Togt. Zook Bot. Udb., p. 136, tab. XVII, figs. 4 — 5.

An estimates of the averages, according to Collett’s measurements of these specimens, gives the following results:

Average in:

Length of the
body varying
between 118
and 238 min.

Length of the
body varying
between 270
and 570 mm.

6 specimens.

9 specimens.

Length of the head in % of the length of the body

22.2

24.8

Depth of the body at the beginning of the dorsal fin „ ,, „ ., ., „ „ ,,

10.7

14.0

Distance between the dorsal fin and t he tip of the snout ,, ,, ,, .. ,, .,

25. «

30.2

., „ „ vent ,, „ ,, ,, „ „ „ „ „ ,, „ ,, ,, ,,

39.5

44.5

Length of the head in % of the distance between the vent and the tip of the last fin-rays

37.o

44.7

612

SCANDINAVIAN FISHES.

Lycodes seminudus, Rhdt, I. c., p. 223; Ltkn, Vid. Meddel.,
1. c., p. 325; Coll., N. Nordh. Exped. , Zool., Fiske , p.
113, tab. IV, fig. 28.

Lycodes perspicillum , Kr., Overs. Vid. Selsk. Forh. Kbhvn
1844, p. 140; Voy, Scand., Lap. cett. (Gaim.) tab. 7;
Naturh. Tidskr. Kbhvn, ser. Ill, vol. I, p. 289.

Lycodes mucosus, Richards., Last , Arct. Voy. (Belcher), vol. 2,
p. 362, tab. XXVI; Bean, Proc. U. S. Nat. Mus., vol. I,
p. 465 (c??); Id., Bull. U. S. Nat. Mus., No. 15, p. 112;
Smitt, Gt. Intern. Fish. Exhib. London, 1883, Swed. Catal.,
p. 17 6.

Lycodes Rossi, Mgrn, Ofvers. Vet. Akad. Forh. 1864, p. 516.
Lycodes Turned, Bean, Proc. U. S. Nat. Mus., vol. I, p. 463
(9?); ibid., vol. IV, p. 244 ( Tuvnerir ); Turner, Contrib.
Nat. Hist. Alaska, Arct. Ser. Pub]. Sign. Serv. U. S. Army,
No. II, p. 93, tab. 4.

by the Knight Errant Expedition at a depth of 608
fathoms in Faroe Channel; and as the other finds extend
all round the world in the extreme north, it is quite

Fig. 148. Lycodes reticulatus, Rossii, twice the natural size. Taken
in 1861, in Treurenberg Bay, Spitsbergen, at a depth of 5 fathoms.

probable that some similar find may once be made in
Scandinavian waters. The species seems to be capable
of ascending fairly high towards the coast, for the Swed-

Fig. 147. Lycodes reticulatus, Tumerii, '/s of the natural size. From the entrance of Chatanga Bay (lat. 75° N.; long. 113“ 30’ E. Gr.),

24th Aug., 1878; Vega Expedition.

Lycodes Lutkenii , Coll., N. Nordh. Expecl., Fiske, p. 103,
tab. Ill, fig. 25; Ltkn, Dijmph.-Togt. Zool. Bot. Udb., p.
128, tab. XVI.

? Lycodes pallidus , Ltkn, 1. c., p. 134, tab. XVII, figg. 1 — 3
(nec Coll.).

? Lycodes coccineus , Bean, Proc. U. S. Nat. Mus., vol. 4, p.
144 (= L. seminudus).

From different regions in all parts of the Arctic Ocean
a number of more or less diverging forms of this species
have been described under different specific names. On
the grounds already given, and if we take into account
the great variations to which the majority of the specific
characters hitherto adopted are demonstrably subject,
these forms may well be comprised within one species,
even if it should prove in the future that some one
or other of them be constant. According to Gunther
(Dee}) Sea Fishes , 1. c.), one of these forms was found

ish Spitsbergen Expedition of 1861 took a young
specimen 33 mm. long in 5 fathoms of water in Treu-
renberg Bay, and the Vega Expedition found their
specimen on a bottom of clay mixed with stones, at a
depth of 15 fathoms in Chatanga Bay, where the bottom-
temperature, however, was — 0‘8° Cels. (30‘5° Fahr.).
The numerous specimens found by the Dijmphna Ex-
pedition in Kara Sea were taken at depths of 46 — 106
fathoms. One of the two specimens which Kroyer re-
ceived from Greenland, and which were taken with a
dredge by Captain IIolboll, was 67 mm. long and was
caught in 90 fathoms of water, the other 40 mm. long
and caught in 30 fathoms of water. Of the older spe-
cimens which have come into the possession of Copen-
hagen Museum at different periods since the time of
Reinhardt, Kroyer writes: “They are generally found

LYCODOIDS.

613

in the stomach of Greenland Sharks" and other fishes
that live at great depths. We may remark, as an ex-
ception, that one of the specimens was accompanied by
the information that it had been caught at the surface,
where it had been found in an exhausted state. It had
probably been driven up from the bottom by some
accident or other. The contents of the stomach of these
specimens consisted chiefly of crustaceans. In the gape
of the larger of his specimens Kroye'r found a Sipun-

culus, an Annelid that lives in the loose bottom of the
sea. It seems probable that the larger specimens —
for the species may attain a, length of at least 56 cm.
— in general live on larger prey and more substantial
food. Collett found in the stomach of his speci-
men, a female 37 cm. long, a partly digested Cottun-
culus microps 1 1 cm- in length and fragments of
smaller fish, the species of which could not lie deter-
mined.

LYCODES V AH LII.

Figs. 149 and 150.

Coloration, which changes with age essentially as in the preceding species , brownish, black , with the light transverse
bands crossed in the older specimens with black streaks and spots or finally disappearing . The covering of scales
varying as in the preceding species , but extending, when complete , forward over the occiput (, sometimes with, scat-
tered scales on the forehead as well), on the belly to the ventral fins, on the vertical fins out, to their margin, and
over a third of the inner surface of the pectoral fins. One mediolateral and one ventro-lateral line ( both often
indistinct). Depth of the body at the beginning of the anal fin about 81/ 2 — 13l/2 % of its own length. Head
convex even in front; its length less than 39 % (as a rule 30 — 38 %) of that, of the tail from the beginning

of the anal fin to the tip of the last fin-rays.

Fig. 149. Lycodes Vahlii, Esmarkii, 1/3 of the natural size. Taken in Varanger Fjord on the 30th of May, 1882.

The property of the Museum of Christiania University.

R. hr. G; D. (96) 100—118; A. (82) 90—102; P. 18—23;
V. (3) 4—5; Vert. 112 — 118 (= 23 -25 +,r).

Syn. Lycodes Vahlii, Rhdt, D. Vid. Selsk. Naturv. Math. Afh., vol.

V, Overs, p. LXXV ; Id., ibid., vol. VII, p. 153, tab. V;
Gthr, Cat. Brit. Mas., Fish., vol. IV, p. 319; Esm., Skand.
Naturf. M.. Christ. 1868, Forh., p. 524; Coll., Forh. Vid.
Selsk. Christ. 1878, No. 4, p. 11 et No. 14, p. 54; Ltkn,
Vid. Meddel. Naturh. For. Kbhvn 1879 — 80, pp. 311 et 329;
Bu. G. et Bean,. Proe. U. S. Nat. Mus., vol. 2, p. 209; Br.
Goode, Fisher., Fisher. Industr. U. 8., Sect. I, pi. 67.
Lycodes gracilis, Sars, Forh. Vid. Selsk. Christ. 1866, p. 40,
tab. I, figg. 1 — 3; Coll., N. Nordh. Exped., Zool ., Fislce,

p. 106.

Lycodes Esmarkii , Coll., Forh. Vid. Selsk. Christ. 1874, Til-
hegsh., p. 95; Ltkn, 1. c., p. 316; Coll., N. Nordh. Exped.,
Zool., Fiske, p. 84, tab. II, figg. 19 — 21, tab. Ill, fig. 22;
N. Mag. Naturv. Christ., Bd. 29, p. 73; Lillj., 8v., Norg.

Fiskar, vol. II, p. 6; Gthr, Deep Sea Fish., Chall. Exped.,
part. LVII (Zool., vol. XXII), p. 77.

? Lycodes pallidas, Coll., Forh. Vid. Selsk. Christ. 1878, No.
14, p. 70: N. Nordh. Exped., Zool., Fislce, p. 110, tab.
Ill, figg. 26, 27.

? Lycodes lugubris , Ltkn, Vid. Meddel. Naturh. For. Kbhvn
1879—80, pp. 315 et 330.

The character that essentially distinguishes Es-
mark’s Lycodes — as this species has been called in re-
cent times — from L. reticulatus, lies in the greater
elongation of the tail. With this character is connected
the greater number of the vertebra? and also, in general,
of the rays of the vertical fins. Here, as in the preced-
ing species, however, the latter number is difficult to fix

Lcemargus borealis.

614

SCANDINAVIAN FISHES.

with certainty; and a more trustworthy expression of
the character may always be found in the proportion
between the length of the head and that of the tail.
In its adult state this species is also better equipped
with scales, though the variations seem to be quite as
great as in the preceding species. On the grounds
given above, and as constant characters of form are an
indispensable condition of systematic clearness, we com-
bine under one specific name the two forms, Lycodes
Vahlii and L. Esmarkii, which have recently been
distinguished on account of the difference in the den-
tition as stated by Lutken.

In form the body resembles that of the Eelpout,
with the characteristics common to all the Lycodes.
The head, which with its flat under surface presents in
the adult fishes the form of a cone longitudinally bi-
sected and somewhat obtuse at the tip, generally mea-
sures in young specimens and in the females rather less,
in the males rather more, than 22 % of the length of
the body, varying in this manner between 1 8 1/2 and
23 V2 % thereof. The only noteworthy irregularity in its
conical shape is formed by the more or less tumid
cheeks (the masticatory muscles). In young specimens
the large muciferous cavities along the upper and lower
jaAvs are especially distinct, generally in the form of
holloAv depressions; but in the older specimens these
depressions disappear. On each side of the snout, at
about the middle of its depth and nearer to the tip of
the snout than to the eye, lies a single tubular nostril.
The eye is oblong and not very sharply marked off from
the surrounding skin, Avhich coalesces Avith the trans-
parent cornea. The length of the eye varies between
about 7 % (in the fry) and 2 % of that of the body
(the latter proportion in the most developed individuals).
Its position is such that in the fry the postorbital length
of the head measures about 44 % of its entire length,
in adult specimens about 55 % thereof, with all the
variations lying between these two percentages. The
sides of the occiput end each in an obtusely pointed
and triangular, opercular flap, Avhich in old specimens
lies a little above and behind the upper end of the root
of the pectoral fin and sometimes measures at the upper,
free margin nearly half the length of the remainder of
the postorbital part. The loAver angle of the gill-opening
lies on a level Avith the loAver angle of the pectoral fin
and in a line Avith the preoperculum, the breadth of

" According to Goode and Bean (1. c.) the length of the. lowi
in a male 632 mm. long 1 Of ,, % thereof.

the isthmus between the two gill-openings being about
equal to the length of the nose between the nostril and
the eye. The length of the loAver jatv varies, according
to our measurements, between about 10 and 11 1/2 % of
that of the body", and is generally someAvhat less than
in the preceding species. The cleft of the mouth is
horizontal; it extends in young specimens hardly any
distance, if at all, behind the perpendicular from the
anterior margin of the eye, but in old specimens di-
stinctly beyond this line. In old specimens the hind
extremity of the maxillary bones lies in a line Avith the
posterior part of the eye, the distance betAveen the tip
of the snout and this extremity measuring in the fry
about 7 % of the length of the body, in old specimens
up to at least 1 13/8 % thereof. This elongation of the
labial parts involves a more advanced development of the
teeth. The teeth in the jaAvs and on the palate are strong,
almost cylindrical, more less curved, and rather scattered.
They are largest in the anterior part of the intermaxil-
lary bones and of the loAver jaw, and are here set in
old specimens in three or even four toavs, in young in
only two, the number of the toavs decreasing behind
until at last there is onty one roAv or sometimes, in
the loAver jaw, two. The foremost teeth in the jaws
are directed straight forward. The i’oavs of intermaxil-
lary teeth do not extend further back, even in old spe-
cimens, than to about a line Avith the middle of the
snout, Avhile in the lower jaw the roAvs extend beyond
the corners of the mouth. On the other hand, the fore-
most intermaxillary teeth are set under the projecting
part of the snout, in front of the tip of the loAver jaw.
On the head of the vomer Ave find from 5 to 8 teeth
of more conical form, arranged transversely in a trian-
gular group. On the palatine bones, according to Col-
lett, Lycodes Esmarkii ahvays has 9 — 11 teeth in a
single or irregularly double toav; Avhile in L. Vahlii,
according to Lutken, these bones are set Avith 11 — 14
teeth in the females and Avith someAvhat feAver than 20
in the males, ahvays in a double roAv. Such is the
tenour of the proposed specific distinction, or in other
Avords: “The toav of palatine teeth is never longer than
the intermaxillary toav, but in most cases (especially in
adult specimens) considerably shorter, sometimes scarcely
half as long” (Coll., of Lycodes Esmarkii ), and: “The
roAv of palatine teeth is generally longer than the in-
termaxillary roAv, at least only slightly, if at all, shorter

er jaw in a female 540 mm. long was 9 1 2 % of ihat of Hie body and

LYCODOJDS.

615

than it (Lutk., of Ly codes Vahlii). In the first place,
however, this character is subject, according to Collett’s
measurements, to considerable variations — even in the
same specimen, a male 58 cm. long, which has been
kindly lent me for examination by Professor Collett,
the row of palatine teeth on one side differs from that
on the other, the right containing 8 teeth, the left 12.
In the second place, we have here to deal with a sexual
character, as both Lutken and Collett have pointed
out: the males in general have longer rows of teeth
than the females, both on the intermaxillaries and on the
palatine bones. Thus, when we remember that in Ly-
codes Esmarldi the four rows of teeth, or at least three '
of them, according to Collett’s measurements (. Nordh .
Exped., 1. c., p. 90), may be equal in length to each
other even in specimens 3 dm. long, the character can
scarcely be regarded as a universally valid distinction
between the two proposed species.

The body, which just behind the head is terete,
about as broad as deep, is gradually compressed later-

Fig. 150. Ly codes Vahlii , gracilis , twice Ihe natural size. Taken
at Drobak in 1866 by M. Sars. The property of the Zoological
Museum of Christiania University.

ally from this point. The depth at the beginning of
the anal fin is 10 — 1 3 1 /2 % of the length of the body
in old specimens, but only about 8 Vs * thereof in the
fry, assuming that we are right in referring to this
species, the fry of which are otherwise unknown, the
Ly codes gracilis of M. Saks (fig. 150). The type-spe-
cimen of this proposed species has been placed at our
disposal for purposes of examination by the kindness of
Professor Collett. In all essential characters of form
the relation of this specimen to Ly codes perspicillum and
L. Rossi — the assumed fry of the preceding species —
is the same as that of Lycodes Vahlii to L. reticulatus
in their adult state. The distance between the dorsal
fin and the tip of the snout is 23 %c\ the distance be-
tween the anal fin and the latter somewhat more than
38 %b , and the length of the tail 61 %e of the length
of the body. The length of the head is only 34 %d
of that of the tail.

“ In L. perspicillum and L. Rossi 261/2 — 29 %.
b !> H >> !) )! >> 431/3 — 46 /.

c ,, ,, ,, ,, ,, ,, 55’/ 2 53 %.

rl „ „ „ „ „ „ 46—40 %.

e Sars found the same uumber in bis Lycodes gracilis.

In this species as in the preceding one, the pectoral
fins are broad and rounded, the membrane being espe-
cially thick below. Their length from the upper angle
of the insertion varies between about 14 and 12 1/s %
of that of the body. The ventral fins are set just behind
the line between the lower angles of the gill-openings,
and so near each other that the distance between them
is scarcely equal to the breadth of their base. They
are of an oblong, triangular shape, and their length in
the male is about equal to the longitudinal diameter of
the eyes, in the female, according to Lill.jeborg, only
% thereof. The skin is so thick that it is difficult to
count the rays without dissection: Brown-Goode and
Bean state the number at 3 ", Reinhardt and Collett
at 4, Lilljeborg at 5.

The vertical fins are of fairly uniform height, but
ascend anteriorly in a rounded slope and coalesce behind
into a sharp (in young specimens) or rather obtuse (in
older ones) point. The anal fin is somewhat lower than
the dorsal, and its length about i/5 of that of the latter.
The longest rays of the dorsal fin, which are situated
above the anterior part of the anal tin, measure about
half the depth of the body at the beginning of the latter.
In the dorsal fin, according to Reinhardt, only the first
ray is simple, with the exception of the rays that occupy
the extreme tip of the tail; in the anal fin, according
to Lilljeborg, the first three rays are simple.

The scales resemble those of the Eelpout in form
and structure. They are thin, flexible, and in general
elliptical, with extremely thin, central nucleus of the
same form and dense, fine, concentric strhe (ridges),
which are prettily broken by numerous, straight, radiat-
ing grooves over the whole surface. When they are
allowed to dry under the thin skin that covers them,
we therefore see, in old specimens, small, fine notches
all round the margin of each scale, the traces of the
ends of the grooves at the edge. The scales of the
lateral line are more circular, with the nucleus thickened
and with a canaliculate duct, contracted and almost
closed at the middle by the growth of the margins
towards each other. Scales of this structure appear even
where the lateral line is externally invisible. The longest
axis of the scale generally lies transversely across the
body. In a male 58 cm. long one of the largest scales

Scandinavian Fishes.

78

616

SCANDINAVIAN FISHES.

is 3 mm. long (deep) and 2 mm. broad, one of the scales
of the lower lateral line 2YS mm. deep and 2 mm. broad.

The internal organs, which are described at
length by Reinhardt, resemble those of the Eel-
pout in all essential respects, only that here the stomach
is somewhat longer. The peritoneum, as well as the
pharynx and palate, is black. In the preceding species
it is white. The abdominal cavity occupies Vs of the
length of the body, and the length of the intestine is
3/5 of the latter. The pyloric appendages of this spe-
cies are very rudimentary. In the preceding species
they are about as large as in the Eelpout. The testes,
are, as usual, two, but the ovary is simple. In adult
females the ripe eggs are as large as those of the Sal-
mon, according to Collett, or about 6 mm. in diameter,
but comparatively few in number, “hardly more than
1,200 in each individual. ’ The spawning-season occurs,
according to the same writer, probably during the first
months of spring. In the intestine Reinhardt found
entire shells of digested mussels, Collett crushed sea-
urchins, starfish, and sea-anemones, as well as frag-
ments of worms and crustaceans.

When this species has passed through the early sta-
ges of growth, it is of a deep brownish black, with from
5 to 8 whitish yellow, transverse bands across the dorsal
fin and down the sides, the posterior bands extending
farthest down, and the hindmost ones of all advancing
even over the anal fin. Two spots of the same light
colour as the transverse bands are set beside each other
on the occiput, just as in the preceding species. The
head is grayish brown, with the opercular flap shading
into black. The white scales shine through the thin

skin that covers them. Such is the appearance of the
fish, according to Collett, until it attains a length of
about 4 dm. The black ground-colour now begins to
encroach upon the transverse bands of whitish yellow,
which are thus broken up, on the dorsal fin into narrow
stripes, on the back into confluent, ocellated spots, and
often joined in pairs in their lower parts so as to form
garland-like figures. In the same way the occipital spots
may be gathered into a transverse band between the
gill-openings. Lmodes VahJii, it is said, may finally
assume an almost plain, brownish black tint.

Esmark’s Lycodes seems to be fairly common in the
deep water off the extreme north of Norway, where its
presence was first discovered in 18,64 by Esmark. Col-
lett enumerates 22 large specimens, between 58 and
70 V2 cm. long, as having been taken in recent times,
principally during the years 1881 — 83, in most cases
on long-lines in Varanger Fjord and at depths of about
100 — 250 fathoms. The species has also been found by
the Norwegian Arctic Expedition in 260 — 459 fathoms
of water, west of the northern part of Spitzbergen, and
at a depth of 350 fathoms off’ Bodo. Gunther states
that it has been met with in Faroe Channel; and the
small specimen from Drobaksund in Christiania Fjord,
that Avas taken in 1866 by M. Sars in 50 or 60 fathoms
of Avater, in all probability belongs to this species.
Brown-Goode and Bean have described it from La
Have Bank (42° 43’ N. lat.) and the Grand Banks on
the east coast of North America. Reinhardt’s type-
specimens Avere from Greenland. It is thus an Arctic
deep-sea fish with a fairly extensive range to the
south.

LYCODES SARS1I.

Fig. 151.

Coloration grayish brown , with regular , brownish black, transverse spots on the back and the upper half of the sides
and also lower down on the hind part of the tail. A blackish brown , longitudinal band forward along the snout
from each of the eyes. Scales scattered, and extending from the occiput along the dorsal side to the beginning of
the last third of the tail. Depth of the body at the beginning of the anal fin about 6 or 7 % of its own length.
Tip of the tail blunt. Length of the head more than 22 % (23 — 26 %) of the length of the tail from the

beginning of the anal fin to the tip of the last fin-rays.

Fig. 151. Lycodes Sarsii, twice the natural size. Taken off Beian in Trondbjem Fjord, on the 31st of August, 1882, in between 80 and
200 ftlmis. of water, by G. 0. Saks. The property of the Zoological Museum of Christiania University.

R. br. 5; R. 15—16; V. 2 (?).

Syn. Lycodes Sarsii , Coll., Forh. Vid. Selsk. Christ. 1871, p. 62; ibid.

1874, Tillcegsh., p. 102; N. Nordh. Kosped ., Zool., Fislce , p.

117; N. Mag. Naturv. Christ., Bd. 29 (1884), p. 78, tab. I, figg.
3 — 4; Ltllj., Sv., Norg. Fish., vol. II, p. 23; G-thr, Deep
Sea Fish., Chalk Exped., part. LVII, (Zool., vol. XXII), p. 80.

LY CODOIDS.

617

Anguilla kieneri, Gthr, Ann., Mag. Nat. Hist., ser. TV, vol.

XIII, (1874), p. 139; Day ( Zoarces ), Proc. Zoo). Soc. Lend.

1882, p. 536.

Lycodes Sarsii would seem to be the smallest Scan-
dinavian species of the genus, for the largest specimen
among those found by Sars in Norway was only 62
mm. long, and the specimen described by Gunther from
the Porcupine Expedition measured 85 mm. As yet,
however, it may be doubtful whether the species is
really distinct from Lycodes Verrillii, which has been
described by Brown-Goodk and Bean" from the deep
water off Nova Scotia and New England, and which
attains a length of at least 18 cm. According to Col-
lett, however, the latter form is distinguished by the
extension of the scaly covering over the lower part of
the body as well, the collection of the spots into more
regular transverse bands across the sides, the pointed
tail, and the stronger teeth, peculiarities all of which, as
we have seen in the preceding forms of the genus, may
appear as differences dependent on age and sex.

The slender, Anguilliform body distinguishes Ly-
codes Sarsii from the preceding forms of the genus.
The greatest depth of the body, just behind the head,
is about 7i2 (8’4 — 8'1 %), and its depth at the begin-
ning of the anal tin 1/15 or Vie (about 6’/2 0/o) of its
length. Its breadth in front is equal to, behind only
slightly less than its depth, the terete shape being dis-
tinctly persistent back to the hindmost part of the tail.
The length of the head is about l/6 (16 or 17 %) of
that of the body. It is of a rounded quadrilateral, pa-
rallelopiped form, reminding us of the head of the
common snake, with comparatively short and roundish
snout. In a specimen 62 mm. long the length of the
eye is 3/10, and that of the postorbital part a little

more than 1/2 of the length of the head. The large
muciferous cavities along the upper and lower jaws —
the row of the latter being continued back to the inferior
corner of the preoperculum — suggest that in this spe-
cimen the characters of youth have been persistent,
fhe beginning of the dorsal fin lies at the end of the
first quarter of the body, that of the anal fin just in
front of the end of its first third, the length of the tail
thus measuring not quite 70 % of that of the body.
The length of the pectoral fins is about equal to the
postorbital length of the head: these fins are blackish
brown at the upper margin. The ventral fins are co-
nical and of about the same length as the eye. They are
set in front of the line between the lower angles of the
gill-openings. Without dissection it is difficult to de-
termine the number of their rays; but they seem to
contain at least three rays each.

Lycodes Sarsii was discovered in 1869 by Prof.
G. 0. Sars, and was described in 1871 by Collett
from a young specimen 43 mm. long, taken at a depth
of between 100 and 150 fathoms in Hardanger Fjord.
In 1882 Sars again met with the species during his
dredging operations at a depth of between 80 and 200
fathoms off Beian outside Trondhjem Fjord, where he
took three specimens between 57 and 62 mm. long.
During the Porcupine Expedition of 1869 Wyville-
Thompson took a specimen 85 mm. long “in the North
Atlantic at a depth of 1 80 fathoms.” Whether it is
identical in species with Lycodes Verrillii, a form quite
common, according to Brown-Goode and Bean, on
the east coast of North America, may still remain an
open question. In any case it is up to the present
one of the rarest forms in the Scandinavian fauna.

LYCODES ML ILEX A.
Fig. 152.

Body plain grayish brown. Scales scattered and irregularly distributed, but extending , when complete , over the
whole body except the head and’ fins. Depth of the body at the beginning of the anal fin about 4 % of its own
length. Tail pointed. Length of the head less than 21 % {IT /, — 16 %) of that of the tail from the beginning

of the anal fin to the tip of the last fin-rays.

R. br. 5; D. 101 — 118; A. 87—103; P. 13—17; V. 3 (?).

Syn. Lycodes murcena, Coll., Forh. Vid. Selsk. Christ. 1878, No. 4,
p. 15 et No. 14, p. 74; N. Nordli. Eecped., Zool., Fiske,
p. 116, tab. IV, ligg. 29 — 31; Lillj., Sv., Norg. Fisk., vol.

II, p. 25; Gthr, Deep Sea Fish., Cballeng. Exped., part.
LVII (Zool., vol. XXII), p. 79, tab. XII, fig. A.

This species is still more elongated than the pre-
ceding one, the caudal part being prolonged to such

Amer. Journ. Sc., Arts, vol. XVI (1877) p. 474; Bull. Ess. Inst., vol. XI (1879) p. 9; Proc. U. S. Nat. Mus., vol. 3 (1880), p. 477.

618

SCANDINAVIAN FISHES.

an extent that the distance between the anal fin and
the tip of the snont is only about 28 % of the length
of the body. The form of the body is otherwise the
same, almost terete throughout, evenly tapering back
from the occiput, and with no distinct lateral compres-
sion before the end of the tail. According to Collett,
the depth at the beginning of the dorsal fin is il/2 —
5 %, and at the beginning of the anal fin 3 — 4 % of
the length of the body. The head is more depressed
and broader, with longer snout, than in the preceding
species: its length is 14 — 12 % of that, of the body.
Half of its length is occupied by the postorbital part;
and in a specimen 2 dm. long the eye occupies 1/5 (in
younger specimens nearly 74) of the length of the head.

to that of the eye. The same remark applies to the
number of their rays as in the preceding species.
Sometimes, according to Collett, we may trace two
lateral lines, one ventral and one median, though the
pores in the latter are rather scattered.

Of Lycodes murcena as of L. Sarsii only four spe-
cimens are yet known in the North, and only one of
these can lay claim to a place within the Scandinavian
fauna. This specimen was found by the Norwegian
Arctic Expedition in June, 1877, at a depth of 350
fathoms off Helgeland. In 1878 the same expedition
took two specimens in 459 fathoms of water, west
of the northern part of Spitsbergen, and one spe-
cimen in 658 fathoms, west of Bear Island. In Faroe

Fig. 152. Lycodes murcena , natural size, from the Norwegian Arctic Expedition. The property of the Museum of Christiania University.

Even in specimens 2 dm. long the muciferous cavities
along the jaws and the lower margin of the preoper-
culum are as distinct as in the preceding species. The
dorsal fin begins further forward than in the preceding
species, the distance between it and the tip of the snout-
being at most about 1/5 (less than 21 %) of the length
of the body or 3/i (in the preceding species at least 4/5)
of the distance between the anal fin and the tip of the
snout, which distance here measures about 26 — 28 %a
of the length of the body. The pectoral fins are ob-
tusely rounded, and their length is about 9 — 873 % of
that of the body. The ventral fins are of the same
form and position as in the preceding species, and their
length too, in a specimen 2 dm. long, is about equal

Channel, however, the English Knigld-Errant Expe-
dition of 1882 took several examples of this spe-
cies at depths of between 540 and 608 fathoms. Lyco-
des murcena thus seems to be one of those Arctic species
which in deep water have a range extending to the
south through the cold layers of the ocean. Its manner
of life is otherwise unknown, but in all probability
resembles that of the other bottom-fishes, among which
it passes its time resting on the cold clay or buried
in its shelter. In the stomach Collett has found frag-
ments of crustaceans ( Themisto and Nannoniscus).

The largest specimens known measure nearly 23
cm. (9 in.), according to Gunther.

PHYSOCLYSTI PLECTOGNATHI.

Physoclysts in which the intermaxillary hones are usually firmly united to the maxillaries, and in which the dental

and articular parts of the lower jaiv are also more or

their lamellce pectinated. No

We now approach those groups among the Phy-
soclysts that, each in its own manner, remind us in a
higher degree than the fishes we have already examined,
of the piscine types of primeval times, the Ganoids.

' less perfectly confluent. Branchial arches complete , and
externally visible interclavicles.

One universal feature that strikes us in the following
Physoclysts, is the marked ossification of the dermal
system as opposed to the weaker ossification of the
endoskeleton. Those parts of the endoskeleton that

According to Gunther’s figure (1. c.) 30 %

PLECTOGNATES.

619

have originally started from the dermal system, as for
example the clavicles, here in many cases have the
character of Ganoid scaly or scutate growths in the
skin. This fairly prominent touch of resemblance to
the Ganoids has induced scientists long to retain the
majority of the following Physoclysts among the lowest,
least modern Teleosts, th'ough the characters by which
both the Plectognates and the Lophobranchii have been
distinguished ever since the time of Cuvier, are evi-
dently later modifications (higher metamorphosic degrees)
of the Teleostean type.

The Plectognates compose an order fairly rich in
forms, varying both in the form of the body and, still
more, in the texture of its covering. Some of them,
most of the Balistoids, are of a fairly regular piscine
form, others of a. polygonal form, which has given them
the name of Coffer-fishes, others again, the Gymnodonts,
of an elongated saccate or globular form. Some have
hard, granulated scales, densely imbricated, juxtaposed
to each other like a mosaic, or firmly united to a ca-
rapace that may cover the greater part of the body,
others small, spiny scales or, in their stead, loose, mo-
bile spines, sometimes of considerable size, which are !
erected in self-defence. All the Plectognates have a
comparatively small, but well-armed mouth, with strong,
but few teeth, the strength of which is increased by
the firm support afforded them by the coalescence and
reduction of the component parts of the jaws. The
short jaw bones form the tip of the facial part of the
head, which part is generally considerably elongated
and also deepened. This elongation chiefly affects the
ethmoid bone, the extent of which on the upper surface
of the cranium may rival that of the frontal bones, and
the parasphenoid bone (the sphenoid bone of fishes),
but is accompanied by the elongation of the preoper-
culum and interoperculuinh The last-mentioned bone
has a narrow, terete shaft, which extends from the hind
inferior angle of the lower jaw along the inside of the
lower anterior prong of the pr<?operculum, just as the
latter extends back from the outer surface of the pro-
cess Avith Avhich the loAver jaw articulates, on the qua-
drate bone. The deepening of the facial part depends
chiefly on the great vertical extension of the parasphe-
noid bone and the pterygoid arch. Sometimes, it is
true, as in the Balistoids, the palatine bone on each side
may be a small, terete, and more or less mobile bone,

bifid at the top and united to the ethmoid bone by
ligaments. But in other cases, as in the genus Tetrodon ,
it resembles the pterygoid bone in form and is firmly
united to the loAver surface of the ethmoid bone. Both
the palatine bones and the vomer are toothless; and the
pharyngeals, which are free, may, it is true, be armed
with teeth, as in the Balistoids and Tetrodonts, but may
also be destitute of teeth, as in the Coffer-fishes. The
shoulder-girdle is generally Avell-developed. The post-
temporal bone is indeed insignificant, being confluent
Avith the mastoid bone, and the supraclavicular bone
sIioavs scarcely anything unusual in its development; but
the clavicular bone proper, though sometimes of ordi-
nary form, may extend so far imvards in some of the
Plectognates, as in the Coffer-fishes, that the bone of
one side is contiguous to that of the other throughout
the depth of the median plane of the abdominal cavity,
thus forming a kind of diaphragm of bone Avith only a
round, more or less tubular hole in the middle for the
passage of the oesophagus. The shoulder-girdle of the
Coffer-fishes is still further extended, beloAv the coracoid
bones, by a pair of large ‘interclavicles,’ of Avhich in the
preceding fishes Ave have found scarcely a trace, but
which Ave shall again meet in the Sticklebacks in the form
of Ganoid plates. In Batistes the clavicle extends so
high up that Avith its upper extremity it touches the
cranium, at the styloid bone, just within and under the
articular surface of the supraclavicular bone.

In Artedi the Plectognates formed the framework
of the order which he called Brancliiostegi (i. e. with
hidden gills , a reference to the narroAV gill-openings just
above or in front of the insertions of the pectoral fins);
and the principal character of the order, according to
its founder, lay in the absence of special branchiostegal
membranes. In general, however, the exact opposite is
the case; the opercular apparatus is by no means im-
perfect in these fishes. The operculum and suboper-
culum are small, it is true, in the Balistoids and Coffer-
fishes, but of normal size in the Tetrodonts. The ad-
vanced development of the preoperculum has been
mentioned above. The hyoid arches, to Avhich the
branchiostegal membranes with their 4* — 6 rays are
attached, have gained in breadth Avhat they have lost
in length; and in most cases one of the rays in each
membrane, usually the innermost, is considerably stronger
than the others, sometimes, as in Tetrodon , several times

a This elongation and deepening is most marked in the Scleroderms (see below); in the Gymnodonts it is less prominent.
6 Sometimes 3, according to Bleeker.

620

SCANDINAVIAN FISHES.

as broad and expanded like a wing. But the branchio-
stegal membranes are entirely hidden under the common
dermal covering, so that the Artedian character of the
order holds good as an external one. The pelvic bones
are generally wanting, but sometimes present, as in the
Balistoids, in which case they are united into a large,
scythe-shaped bone, inferiorly bounding and supporting
the abdominal cavity, even when the pectoral tins have
disappeared or have been reduced to a spiny, osseous
projection at the hind extremity of this bone. The
remainder of the skeleton is marked not only by its
weak ossification, but also by the small number of the
vertebrae, which may sometimes (in the Coffer-fishes)
sink to 14 and never exceeds 30. Sometimes, as in the
Tetrodonts, the upper spinous processes (neural spines)
of the first 5 or 6 vertebrae are longitudinally cleft in
two, with the halves laterally extended. Ribs are want-
ing in most cases, but present in the Balistoids. As
the caudal fin is frequently of extraordinary size — though
with few, less than 13, branched rays — , its insertion
in the skeleton has also undergone proportionate increase
in strength and firmness, partly by means of large
hypural bones, which are firmly united in the Coffer-
fishes into a compressed tip, hard as ivory, partly by
high and longitudinally extended, neural and haemal
spines on the last caudal vertebrae. Sometimes however,
as in the Sunfishes, the caudal fin is a secondary growth
and finds an extremely feeble support in the more or
less atrophied tip of the tail.

In spite of the considerable differences that occur
both in the form and covering and also in the internal
structure, the Plectognates together form a natural group,
with persistent traces of the Ganoid type, which give
them an appearance unusual in the fauna of our day.
The nearest approach to the preceding forms we find,
as Cope has already pointed out", in a comparison be-
tween the Balistoids (especially the genus Triacanthus,
with its free maxillaries) and the Leather-fishes ( Acan -
thuridce, a family within the comprehensive series of
Mackerel-fishes), with the same form of body, the same
elongation and deepening of the forepart of the head
and shortening of the jawbones, the same union of the

posttemporal and mastoid bones, and the same length
of the pelvic bones, which in the Acanthuroids, however,
are not confluent. Set side by side with the skeleton of
an Acanthurus the skeleton of a Batistes seems at the
first glance to claim a position within the same family.
The systematic separation between the Plectognates and
the other Physoclysts is thus by no means too great to
admit of their being ranged side by side with each other.

About 200 species of Plectognates are known. The
greatest wealth of forms belongs to the tropical seas;
but one or two species penetrate into rivers, or even
pass their whole life there. To man these fishes are of
only slight value. Their flesh is in general not eaten* 6,
and many of them are even poisonous. They attract
attention, however, both by their extraordinary form
and often by their beauty of colour, as well as by their
singular manner of life. Most of them are shore-fishes,
but some belong to the surface-regions of the ocean;
they do not include any deep-sea forms of pronounced
character, although two species have been met with
below the 100-fathoms lineb Most of them are ren-
dered incapable of any vigorous motion by their hard
and stiff dermal covering or by their clumsy form.
By the small size of the gape most of them are com-
pelled to find subsistence in small animals or in pieces
torn from larger ones. Their food consists chiefly of
corallines, jelly-fishes, worms, or mollusks. The lively
colours that adorn many of them are reminiscences of
their life in the coral-groves, where they associate with
scaly-finned Chsetodonts of no less varied hues, or among
the seaweeds with their shifting colours. The species of
plainer dress resemble the sand in colour, or are attired
for a life in the open sea. Their powerful teeth may
well serve to protect them from attack, but as a rule
they have quite other weapons of defence. Many of
them have been famed from ancient times for their
faculty of filling the oesophagus, which is capable of
extraordinary distension, with air. In this manner the
body is expanded, sometimes to a perfect globe, and
the dermal spines are erected in all directions into a
front of pointed spikes. “One day,” wrote Darwin from
Bahia in 1832 d, “I was amused by watching the habits

" Trans. Amer. Philos. Soc., Philad., vol. XIV, n. set., p. 458.

6 According to Schleqel (Sieb., Fn. Japan.'), however, several of these fishes are eaten by the Japanese: one Tetrodon is even stated
to compose the principal food of the poorer classes during winter, while the fisherman is forbidden, under pain of heavy penalties, to expose
for sale other species of the same genus.

c Both these finds, however (see Gunther, Deep /Sea Fish. Chall. Exped., p. 266), may have entered the trawl while it was being
sunk or hauled in.

d A Naturalist' s Voyage (London, ed. 1884) p. 13.

PLECTOGNATES.

of the Diodon ( Ckilomycterus ) antennatus, which was
caught swimming near the shore. This fish is well
known to possess the singular power of distending itself
into a nearly spherical form. After having been taken
out of water for a short time, and then again immersed
in it, a considerable quantity both of water and air is
absorbed by the mouth, and perhaps likewise by the
branchial orifices. This process is effected by two meth-
ods; the air is swallowed, and is then forced into the
cavity of the body, its return being prevented by a
muscular contraction which is externally visible; but
the water, I observed, enters in a gentle stream through
the mouth, which is kept wide open and motionless:
this latter action must, therefore, depend on suction.
The skin about the abdomen is much looser than that
on the back; hence during the inflation, the lower sur-
face becomes far more distended than the upper; and
the fish, in consequence, floats with its back downwards.
Cuvier doubts whether the Diodon in this position, is
able to swim; but not only can it thus move forward
in a straight line, but it can turn round to either side.
This latter movement is effected solely by the aid of
the pectoral fins, the tail being collapsed, and not used.
From the body being buoyed up 'with so much air, the
branchial openings were out of water; but a stream
drawn in by the mouth constantly flows through them.

“The fish, having remained in this distended state
for a, short time, generally expelled the air and water
with considerable force from the branchial apertures and
month. It could emit, at will, a certain portion of the
water; and it appears, therefore, probable that this
fluid is taken in partly for the sake of regulating its
specific gravity.

“This Diodon possessed several means of defence.
It could give a severe bite, and could eject water from
its mouth to some distance, at the same time making a
curious noise by the movement of its jaws. By the
inflation of its body, the papillae, with which the skin
is covered, become erect and pointed. But the most
curious circumstance is, that it secretes from the skin
of its belly, when handled, a most beautiful carmine-red
fibrous matter, which stains ivory and paper in so per-
manent a manner, that the tint is retained with all its
brightness to the present day. I am quite ignorant of
the nature and use of this secretion. I have heard

(>21

from Dr. Allan of Forres, that he has frequently found
a Diodon, floating alive and distended, in the stomach
of the Shark; and that on several occasions he has known
it eat its way, not only through the coats of the sto-
mach, but through the sides of the monster, which has
thus been killed. Who would ever have imagined that
a little soft fish could have destroyed the great and
savage Shark?”

The power here mentioned by Darwin, that of pro-
ducing sounds, is one of the faculties highly characteristic
of these fishes. The Plectognates have long been known
as the most assiduous musicians of the ocean. Now it
is the jaws, now the articular surfaces of the spinous
rays in the dorsal or ventral fins, where these fins con-
tain such rays, that make a snapping, ringing, grating,
or laughing noise"; now it is the hissing sound of water
and air driven out of the body; now it is the air-
bladder whose vibrations may be distinctly heard like
the beating of a drum4. For this purpose the air-bladder
is attached, in the drumming Balistoids, to the shoulder-
girdle on each side and to special, large dermal plates
just above the upper angle of the pectoral tin. By the
quivering movements that accompany the violent con-
traction (ef. above, p. 193) of the ventral parts of the
lateral muscles, the postclavicular bones, which are in-
serted in these muscles, are set in a vibrating motion
which is transferred to the wall of the air-bladder and
to the air contained therein, from which the sound is
transmitted to the said osseous plates in the skin above
the angle of the pectoral fin, and through their medium
to the outer world. All the snapping, ringing, grating,
laughing, or hissing sounds are probably intended, here
as in other fishes which have this power, in the first
place to frighten away some approaching foe. The
notes or drumming noises of the air-bladder may also
conduce to the same result, or may be produced by
the death-throes of the fish. But, as Sorensen has
remarked, the sonorous tones that compose the song
of these fishes, undoubtedly serve in other cases, espe-
cially during the spawning-season, as call-notes.

Cuvier1' divided the Plectognates into two families,
the first with free teeth in the jaws and usually with
hard (plate-like) dermal covering — hence the name of
les Sclerodermes, from Gxfajpos, hard and deQ/ucc, skin
— the second with the jaw-teeth confluent and forming

“ Cf. Sorensen, Om Lydorganer hos Fishe , Kjabenhavn, 1884.

6 Cf. Mobius, Balistes aculeatus , ein trommelnder Fisch , Stzber. Akad. Wiss. Berk 18 89, p. 999.
c Regne Animal !, ed. 1, tom. II, pp. 145 and 149; ed. 2, tom. II, pp. 365 and 371.

622

SCANDINAVIAN FISHES.

composite dental growths, one or two in each jaw,
which project like a beak beyond the lips — hence the
name of les Gymnodontes , from yvurog, naked and bd'ovg ,
tooth. Of these families the latter is distinctly more
perfectly Plectognate in its characters. At the present

time we generally follow Bonaparte" and divide these
fishes into four families, two within each of the Cu-
vierian families b. Only two species, each representing
a distinct family, have been found within the limits
of the Scandinavian fauna.

Fam. 0 R T H A G 0 R I S 0 I S) M.

Gymnodonts with stunted trunk , body compressed laterally { round or elliptical) and not distensive, and- contiguous
vertical fins. Dental disks of the jaws undivided. Pseudobranchice large. Air-bladder0 , pelvic bones ,

and ventral fins wanting.

With an appearance as though they were of the
normal piscine form but with truncate tail, these Lagg-
bukar {Tun-bellies) as Retzius called them together

After Wellenberg.

with the rest of the Gymnodonts — deservedly excite
attention even by their external characters. In form
and appearance they remind us somewhat of the Opah;
but merely a casual examination is enough to show

that they are of an entirely different type. They really
rank among the most singular piscine forms. The
stunted appearance of the body depends, it is true, on
the comparatively small number of the vertebrae (16
or 17); but this number may be still smaller in the
Coffer-fishes. The short form of the body is really
caused by its depth posteriorly, this being due to a
prominent development of the upper and lower arches
of the vertebrae, with their spinous processes, in the
caudal part of the body, and of the interneural spines
of the dorsal fin and the in ter haemal spines of the anal
fin (fig. 153). This deviation from the other Plecto-
gnates is connected with an extremely unusual course
of development.

In the middle of the last century KolreuterC gave
an excellent description and a figure of no small merit,
considering the period of its execution, of a fish 16
mm. long and 19 1/2 mm. deep, “which at a casual
glance might easily be taken for a head cut off from
the trunk, and nothing more.” Closer examination
soon showed, however, that this specimen was a young
form of Orthagoriscus ( Tet-rodon ) mola, as Linnaeus e
interpreted this find. But the most singular circum-
stance was that it had no caudal fin whatever. In
another specimen, 40V2 mm. long and 49 1/2 mm. deep,
on the other hand, Kolreuter found a distinct, cre-
nulated caudal lobe along the hind margin of the fish,
uniting the dorsal fin to the anal. Since that time
these small fishes have been met with on numerous

“ Catalogo metodico dei pesci europei , p. 87.

b Bleeker ( Atlas Ichthyologique, tome V) divided the Plectognates into three orders and these again into 6 families corresponding to
Gunther’s subfamilies: Ostraciontina ; Triacanthina , Balistina; Triodontina , Tetrodontina , Molina.

c Costa ( Fn . d. Regn. di Napoli , tav. 63 e 64) describes and figures a rudimentary air-bladder in Orthagoriscus mola.
d Nov. Comment. Acad. Sc. Petrop., vol. X (1766), p. 337, tab. VIII, figs. 2 and 3. Mola aculeata , limbo abdominis producto,
attenuate , carnoso.

e Syst. nat., ed. XII, tom. 1, part. 2, Add. (in fine).

SUNFISHES.

623

occasions, and the late development of the caudal fin
has been fully demonstrated (fig. 154). According to
Lutken®, however, we have a still earlier developmental

Fia;. 154.

Larvae of Ortliagoriscus mola; A. at a length of 18 mm.,
B. at a length of 32 min. After Gunther.

stage of Ortliagoriscus in the form named by Richard-
son Ostracion loops (fig. 155), after the larval speci-
mens taken at the surface of the South Atlantic and

Fig. 155.

Ostracion hoops, Richardson.

After GUnther.

figured by Hooker. These larva; are of a still more
curious form, with their large eyes and their equipment
of large spines, such as those we have seen above on the
heads of several Acanthopterygian larvae, notably within
the great series of the Scombromorphs, and with small
spines besides on the dorsal side. The difference be-
tween them and Kolreuter’s larvae, however, is not so
great as to prevent us from easily recognising the cor-
responding position of the large spines. As in the
younger of Kolreuter’s larvae we here find one large
spine at the middle of the forehead between the eyes;
three at the sharp dorsal margin, the hindmost of which,
just in front of the dorsal tin, is the largest; one6 on
each side above the eyes, as well as on each side of
the hind part of the body between the dorsal and anal
fins. Whether the other spines correspond exactly in
both larvae, is a more difficult point to decide. The

4

caudal tin is still less developed than in the younger
of Kolreuter’s larvae. As the development advances,
the large spines are said to become comparatively
smaller, but the small spines, the papillae with radiat-
ing striae, to extend densely and evenly over the tvhole
body. The transition to the Ortliagoriscus form is thus
not far distant0.

If we compare these youngest forms of Ortliago-
riscus with the normal forms of piscine larvae, the
greatest difference lies in the fact that here the caudal
tin, which otherwise is the first fin to appear — as it
also is the original organ of motion in all vertebrates — ,
is developed last of all the fins. This difference has
left its mark not only on the structure of the skeleton,
but also on the development of the muscles. Here the
original and, in later life, the most powerful organs of
motion are the dorsal and anal fins. Their supporting
bones (the interneural and interhsemal bones) and their
motory muscles exercise a determinant influence on
the development of the whole trunk-

In dissecting an Ortliagoriscus d we have first to cut
through the enormously thick skin, which is lined with
a strong tendinous membrane ( aponeuros ). Within the
latter, on the sides of the body, w6 make the remark-
able observation (fig. 156) that the muscles most highly
developed in other fishes, the large lateral muscles of
the body, are here apparently wanting. Between the
spinal column and the skin runs, as usual, in a hori-
zontal direction, a fibrous membrane (sf), which here
is especially strong, and divides the visible muscular
mass, just as in other cases it divides the large lateral
muscles of the body, into two halves, an upper and a
lower. But by far the greater part of this muscular
mass is here composed of the flexors of the dorsal ( mpd )
and anal ( mpa ) fins. The flexors of the dorsal fin,
divided into two layers of different colour and firmness
— a surface-layer and an under-layer — run from the
occiput, from the spinal column with its superior pro-
cesses and interneural bones, and from the upper sur-
face of the horizontal fibrous membrane (sf), and are
attached by a special sinew to each side of the basal
vol. VIII (1871), p. 320, note.

Powerfully magnified.

u See GOnther, Ann., Mag. Nat. Hist., ser.

6 In K6lreuter’s larva two.

c According to Lutken’s opinion of KOlreuter’s larvae. Still we should observe that the larvte of Diodon correspond just as closely
to Richardson’s larvae, especially in the position of the large spines; but in the former the end of the notochord projects between the dorsal
and anal fins. A larva of this description, 2 inm. long and of ellipsoidal form, which was taken at the surface “between Java and the Cape of
Good Hope,” has been obtained by the Royal Museum through Captain P&hl of Hamburg. In the same jar (and therefore taken probably at
the same time) lay another larva, 4 mm. long and of more distinct Diodon form, but still, like the preceding one, with the dorsal and anal
fins and the tip of the tail between them all situated on the ventral side.

d Cf. Wahlgren, Nagra anteckningar om en stor klumpjisk , Lunds Univ. Arsskrift, tom. IV.

Scandinavian Fishes.

79

624

SCANDINAVIAN FISHES.

part of each ray in the dorsal tin. The flexors of the
anal tin, also divided into a surface-layer and an under-
layer, originate from the bodies of the hindmost ab-
dominal vertebrae, from the caudal vertebrae with their
inferior processes and interhaemal bones, and from the
under surface of the horizontal fibrous membrane, and
are attached to the rays of the anal tin in the same
manner as we have just described in the case of the
flexors of the dorsal tin. The flexors of the caudal tin

sf mpd
mpa , \

Fig. 156. Position of the muscles and viscera in Orthagoriscus mold,
on a reduced scale. After Cleland.
mpa , the flexors of the anal fin ( musculi pinnce analis ); s/, the
fibrous dividing membrane ( septum fibrosum ) ; mpd, the flexors of the
dorsal fin ( musculi pinnce dorsalis ) ; maa, the anterior abdominal muscle
(m. abdominalis anterior ); h , the liver; l , the ligament between the
intestinal sac and the liver; i, the peritoneal investment of the in-
testines (intestinal sac); p, the peritoneum, opened and partly removed;
map, the posterior abdominal muscle (in. abd. posterior)-, a, the vent;
mpc, the flexors of the caudal fin (m. pinnce caudalis).

(mpc) form no compact muscular mass; each of the rays
in the caudal tin is furnished on each side with a spe-
cial, ventricose muscle, and these muscles start partly
from the bodies of the hindmost vertebra:', partly from
the supporting bones of the caudal tin (the hindmost
interneural and interhtemal bones), partly from the
superior and inferior folds of the horizontal, inter-

muscular, fibrous membrane (sf). Of the system of the
large lateral muscles there remain only two abdominal
muscles, belonging to the under-layer of the system,
the first (map) starting from the top of the clavicle, the
second (maa) from the coracoid bone, and both attached
to the outer surface of the strong peritoneum (p). So
great is the modification both of the skeleton and of
the musculature that has accompanied the late develop-
ment of the caudal fin, and rendered the dorsal and anal
fins the principal organs of locomotion. These two fins
have attached to themselves, from the very beginning,
the great mass of muscles and supporting bones; while
the caudal fin has become a secondary organ, which has
been forced to adapt its development to pre-existent
circumstances, and which has thus sunk almost to an
adipose fin, though thin osseous rods have been deve-
loped to rays within it. The influences of this revolu-
tion in the development of the organs of motion have
also affected the central nervous system. The family
is remarkable, from an anatomical point of view, not
only for the insignificant size of the brain", which is
extraordinarily small, even within the class of fishes,
but also for the fact that the spinal marrow, soon after
its passage through the occipital foramen, ramifies in
the well-known “horsetail form” (cauda equina), without
being continued any further in the form of a continuous
chord within the spinal canal* * 6.

Whether more than two species may be distinguished
with certainty within this family, must still be regarded
as a doubtful question. Ranzani’s attempt0 to establish
16 species, distributed among 6 genera, has long ago
been abandoned as futile; and Putnam’s evidence in
favour of the opinion that the young specimens described
by Kolreuter, Pallas, and others belong to a distinct
species of a separate genus seems to require further
confirmation. It has also been proposed0 to refer the
two established species to two distinct genera, one of
which, Ranzania, with more elongated body and with
the dermal covering smooth but divided into hexagonal
plates, has been met with on the English coast, but
never, up to the present time, within the limits of the
I Scandinavian fauna.

“ In large specimens of Orthagoriscus mola, according to Haktjng, the brain may weigh no more than '/718,,0, e- n°t quite 0‘000014,

of the total weight of the body.

6 Arsaky, De piscium cerebro et medulla spinali, Halle 1813, p. 4, tab. Ill, fig. 10.
c Nov. Comm. Acad. Scient. Inst. Bonon. Ill (1839).
d Proc. Amer. Assoc. Adv. Sc., 19th Meeting (1870), p. 255.
e Steenstrup and LOtken. Overs. Vid. Selsk. Forh. Kbhvn 1863, p. 42.

S UNFISH.

625

Genus ORTHAGORISCUS,

Body round or of a short elliptical form. Skin rough ivith dense, fine spines and papillae. Distance between the
dorsal fin and the tip of the snout (in adult specimens ) less than 3/4 of the length of the body.

The name of this genus", according to Rondelet’s*
interpretation of Pliny, is of classical origin and derived
from its power of producing grunting sounds. The
name was introduced by Schneider0 into the post-
Li nnaean nomenclature. Rondelet’s contemporary Sal-
vianus did not approve of his interpretation, and gave
the Sunfishes another name, Mola (millstone), with re-
ference to the round form of the body and derived from
the popular language of Marseilles. This generic name
was adopted by Johnston d and Willughby6, but in
Linnaeus it passed into a specific designation7.

Orthagoriscus is distinguished from the other genus
of the family not only by the different covering of the
body, but also by a special direction of development,
which brings about a secondary increase of compara-

tively considerable extent in the region of the caudal
tin, and involves an elongation of the body that causes
the apparent removal in a forward direction with age
of the dorsal and anal tins. The above-mentioned un-
certainty with regard to the distinction of the species
within this genus depends partly on our imperfect know-
ledge of the earlier stages of the development, partly
on a singular form of the caudal tin, with a pointed
upper lobe, which has been observed both in younger
individuals (fig. 154, B) and in older ones6', without
enabling us as yet to decide whether it is of incidental
(individual) description or of some other signification.
In the Scandinavian fauna, however, this uncertainty
is of no importance.

THE SHORT SUNFISH (sw. klumpfisken).
ORTHAGORISCUS MOLA.

Plate XXVII, fig. 4.

Pectoral fins rounded in old specimens; their length at most about x/7 (14 or 15 %) of that of the body. Colo-
ration bluish gray, on the back darker and brownish, growing lighter ivith a silvery lustre down the sides,

on the belly whitish.

R. br. 6; D. 17 — 18*; A. 15 — 17 ; P. 11—13*; V. 0; V.
11—14; Vert. 16 l. 17.

Syn.' Ostracion cathetoplateus subrotundus inermis asper, pinnis pec-
toralibus horizontalibus, foraminibus quatuor in capite, Art.,
Gen., p. 61; Syn., p. 83.

Tetraodon Mola, Lin., Syst. Nat., ed. X, tom. I, p. 334;
Pall. ( Diodon ), Spicil. Zool. , torn. I, fasc. VIII, p. 39, tab.
IV, fig. 7; Retz. {Tetrodon),Ye t. Akad. Handl. 1785, p. 115,
tab. IV; Bl. {Diodon), Naturg. Ausl. Fisch., pt. I, p. 75,
tab. CXXVIII; Retz. {Tetrodon), Fn. Suec , Lin., p. 310;
Schn. {Orthagoriscus), 1. c.; Nilss., Prodr. Ichtli. Scand ., p.

Fins brownish.

Ill; Wellenbergh, Observ. Anat. de 0. mola, disp. Lngd.
Bat. 1840; Schleg., Fn. Japon., Poiss., p. 288, tab. CXXVII;
Costa, Fii. Regn. Nap., Pesci, pt. 2, Plettogn. Gymnod., tab.
LXIII et LXIV ; Ekstr., Gbgs. Vet., Vitt. Samh. Handl. 1850,
p. 40; Kr., Danin. Fiske, vol. Ill, p. 732, Nilss., Skand.
Fn., Fisk., p. 697; Cleland, Nat. Hist. Review 1862, p.
170, tab. V et VI; Gthr, Cat. Brit. Mas., Fish., vol. VIII,
p. 317; Coll., Vid. Selsk. Forli. Christ. 1874, Tilliegsh.,
p. 203; ibid. 1879, No. 1, p. 101; N. Mag. Naturv. Christ.,
Bd. 29 (1884), p. 114; Malm, Gbgs, Boh. Fn., p. 599;
Winth., Naturh. Tidskr. Kbhvn, ser. 3, vol. XII, p. 54; Mor.,

° ’ ()o!)ayooi (JY.og, sucking-pig.
b De Piscibus, lib. XV, cap. VII, p. 425.

Syst. Ichthyol. Blochii, posth., pp. LVII and 510.
d Historia naturalis de piscibus et cetis , lib. 1, tit. 1, cap. Ill, art. II, punct. VIII.
e Historia Piscium. lib. 4, sect. Ill, cap. VI, p. 151.

f Covier is stated, it is true, in his anatomical lectures — in 1798, according to Steenstrup and LiItken, in 1800, according to
Gill; thus prior to the appearances of Schneider’s work quoted above — to have advocated the resumption of Mola as a generic name {Mola
rotunda ); but in his systematic works he recognised Orthagoriscus.
g Monaco, Bull. Soc. Zool. Fr., tome XIV (1889), p. 17.

h Sometimes 16, according to Kroyer and Moreau; sometimes as many as 20, according to Lilljeborg.
i Sometimes 14, according to Steenstrup and Lutken; sometimes 18, according to Lilljeborg.

* Sometimes 14, according to Retzius.

626

SCANDINAVIAN FISHES.

Hist. Nat. Poiss. Fr., tom. II, p. 74; Mob., Hcke, Fisch.
Osts., p. 101; Day, Fish. Gt. Brit., Irel., vol. II, p. 272,
tab. CXLVIII; Be. Goode, Fisher., Fisher Industr. U. S., sect.
I, p. 169, tab. 35; Lillj., Sv., Norg. Fisk., vol. 3, p- 425.

Orthagoriscus hispidus, Schn., 1. c., p. 511 (juv., ex Pall., 1. c.);
Cuv., R. anim., ed. I, tom. 2, p. 149 = 0. spinosus , Id.,
ed. 2, tom. 2, p. 370; Rich., Voy Sulph., Zool. , vol. I,
p. 125, tab. LXII, figg. 10—12.

Cephalus brevis , Shaw, Gen. Zool., vol. V, p. 437, tab. 175.

Diplanchias nasus , Rafin., Caratt. Ale. N. Gen., p. 17; Stp,
Ltkn ( Mola ), Overs. Vid. Selsk. Forh. Kbhvn 1863, p. 36;
Ltkn, Forh. Skand. Naturf. M. Sthlm 1863, p. 378; Wahlgk.
N. Ant. Mola nasus, Lunds Univ. Arsskr., tom. IV (1867)
cum tab.

Ozodura orsini, Oz. ursini , Tympanomium planci, Diplanchias
nasus, Trematopsis Willughbii, Orthagoriscus Retzii, Ort.
ghini, Ort. Rondeletii, Orth. Blochii, Ort. redi, Ort. aculeatus,
Ranzani, 1. c.

Ostracion hoops, Rich., Ichthyol. Voy. Ereb., Terr., pag. 52,
tab. XXX, figg. 18 — 21; juv. hujus speciei sec. Ltkn, Gthk,
Ann. Mag. Nat. Hist., ser. 4, vol. VIII (1871), p. 320, not.

Orthagoriscus ozodura , Halting, Notices Zool. etc., Verb. Akad.
Wet. Amsterdam, D. XI (1868).

This Sun-fish attains a length of at least 8 feet
(2V2 metres). The largest specimen of which, to the
best of our knowledge, we can speak with positive
certainty, was exhibited from New South Wales at the
Fisheries Exhibition of 1883 in London, and measured
8 feet in length and 12 feet in depth from the tip of
the dorsal fin to that of the analT In young specimens
the form of the body is almost circular. At a length
of 4 V2 — 572 dm. the depth of the body is about 2/3
(69 6 — 64 %) of the length, at a length of about 2 m.,
about 1/2 or 48 % thereof. Thus, the body is elongated
with age to an elliptical form, with the depth fairly
uniform at the middle, while the hind part ends more
abruptly, but the forepart is somewhat prolonged, form-
ing a short muzzle. The thickness is fairly constant,
being greatest above and behind the eyes, where it
measures about 1/i of the greatest depth, and decreasing
without a break, slowly behind and more suddenly in
front. In old specimens, however, on each side above
the eye and back to the perpendicular from the begin-
ning of the insertions of the pectoral fins, we find a
longitudinal, blunt ridge; and a similar ridge may ap-
pear below the gill-opening and the pectoral fin. In
this manner the section of the body at the gill-openings

is rendered more or less distinctly hexagonal, with the
upper and lower angles acute, the latter more so. Both
the dorsal and the ventral edges are sharp and carinated,
the former from the forehead behind the eyes back to
a point a little in front of the beginning of the dorsal
fin, the latter from a point a little behind the lower
jaw back to the vent. The forehead, on the other hand,
is convex; and the tip of the snout is furnished in old
specimens with a rough and sclerous, mobile, round pad,
which projects some distance above and in front of the
mouth. At the ventral margin, a little below and be-
hind the lower jaw, we find in old specimens a similar
osseous growth in the skin, consisting of two, or even
three, fusiform, bony plates, set in a row one after an-
other. In old specimens, too, the ventral profile is
distinctly more arcuate than the dorsal. The base of
the caudal fin passes evenly into the body, so that the
fin projects like an attenuated dermal margin at the
end of the sharply rounded (convex) hind edge of the
body. In young specimens the caudal fin is united
above and below to the lower posterior angle of the
dorsal fin and the upper posterior angle of the anal fin;
but in old specimens it is distinctly separated from
these fins. Its form, though, as we have mentioned
above, it may sometimes present a singular abnormity
— perhaps a malformation or the cicatrice of a
wound — in general corresponds to that of the hind
margin of the body; but at the margin, opposite the
tip of each ray, we find a round incision, which is
more or less completely filled by a compressed, osseous
growth. The dorsal and anal fins rise in the form of
acute-angled triangles — though usually with the apex
rounded — with the anterior margin thick but in profile
concave, the posterior sharp (attenuated) and convex.
These two fins cannot be depressed like normal fins in
their longitudinal direction (the second dorsal and the
anal fins of the Tunnies, for example, are also stiff
when pressed in this direction); but they are highly
flexible laterally, and their movement in this direction0
is assisted by the texture of the skin. This last or-
gan, which is covered with numbers of stiff wrinkles
and ridges, crossing one another in irregular squares,

a Rondelet states, it is true (1. c.), that the Sunfish may attain a length of 6 cubits (23/4 ni.) or more; but this assertion has not
been confirmed, to the best of our knowledge, in modern times, unless we accept Lacepede’s statement that in 1735, on the Irish coast, a
Sunfish 25 feet long was found, “which consequently appeared at night like a shining disk more than 400 square feet in area” (Hist. Nat.
Poiss., tom. I, p. 511).

b Sometimes 72, according to Kr0yer.
c With respect to the musculature see above.

SUN FISH.

627

is extraordinarily thick over the greater part of the
body. In a Sunfish 965 mm. long the skin on the
trunk, according to Cleland, was about 25 mm. thick".
The rigidity of the skin is increased by the spiny and
nodose, small, osseous tubercles4 * which are densely spread
over its surface, and which give the skin both on the
body and the tins a shagreened appearance. But a
rather broad band of thinner and looser skin, with more
granulated or even smooth surface, runs on each side,
along the very margin of the body, from the vent along
the bases of the anal, caudal, and dorsal tins. This
band of flexible and movable skin, which is folded
round the bases of the dorsal and anal tins, admits of
the bending of these two fins at right angles to the
sides of the body. The first, six rays in these tins are
thick and stiff, neither articulated nor branched, and
suddenly increase in length, thus forming the greater
part of the anterior margin of the tin — only the apex
belongs to the seventh ray, which is branched. The
posterior rays, again, suddenly decrease in length be-
hind, are branched, and expand backwards at the tip
like a fan. Thus, the whole of this apparatus forms, to-
gether with the caudal tin, a swimming-blade, elastic
behind, which by alternate movements to right and
left of the dorsal and anal tins drives the fish forward,
just as a boat is propelled by sculling. The compara-
tively short pectoral tins, which, as well as the other
tins, are rough with small osseous tubercles on both
sides, are of rounded form, with the first, and last, rays
simple, the others multifid at the tip, and the middle
ones (the fifth and sixth) longest. They are set, half-
way up the body and just behind the middle point
between the tip of the snout and the beginning of the
dorsal or the anal fin. They are inserted horizontally,
as in the Opah, and move up and down, their true

function thus being probably to maintain the ecjui-
librium of the body. Just in front, of the insertion of
the pectoral tin, with height, about, equal to the length
of this insertion, lies the transversely set, gill-opening
on each side of the body, elliptical in shape, but pointed
at both ends, and with the anterior half covered with
a thin skinc, which is a continuation of the true branch-
iostegal membrane, while the anterior margin is formed
by the skin itself, which covers the operculum. These
small openings are the only external boundaries between
the head and the body. This circumstance has caused
the comparison of the whole fish to a swimming head**.

The head of the Sunfish (from the tip of the snout
to the gill-openings) measures in young specimens
(7a — 1 metre long) between 7s and 3/10 of the length
of the body. In older specimens, which acquire a more
elongated form of body, it becomes comparatively some-
what, smaller, sinking to at least, 29 % of the length of
the body6. In old specimens, which have their short
snout tipped with the hard osseous disk, this disk forms
the extreme end of the snout. In young specimens
the mouth is set, exactly at the tip of the snout, and
shows between the thin lips, which only partially hide
them, its two white dental disks, that of the upper jaw
hooked at the tip, that of the lower jaw even. Behind
(within) these disks we find in young specimens 4 or 5
similar disks or rows of divided, transverse disks, close
behind each other, which are gradually worn away in
course of time, until they disappear. The gape is small, as
in all the Plectognates; in a Sunfish 47 cm. long it can be
opened to a height of only 33 mm. and measures 24 mm.
in breadth, or rather less than the longitudinal diameter
of the eye. The eyes are set very low, in comparison with
their position in the rest of the Plectognates, but much
nearer to the dorsal margin than to the ventraP; their

4 In a large Sunfish dissected by Turner and Goodsir (Nat. Hist. Review 1862, p. 185) the skin varied in thickness on different

parts of the body between 6 and 100— 127 mm. In another specimen, which was unusually large, Goodsir found the thickness of the skin
at certain spots to be 152 mm.

6 The osseous tubercles vary in size, large and small being interspersed with each other; but even the largest ones in large Sunfish
are scarcely of the size of small pins’ heads. At the base they bear radiating striae, and are irregularly incised and dentated at the margin, the
teeth of one tubercle fitting into the incisions in the nest.

e When this membrane lies in folds, or when it bursts — as often happens in stuffed specimens — it may appear as though there
were two gill-openings on each side; and this is perhaps the explanation of the character of two gill-openiDgs on each side that was given
by Rafinesque to his genus Diplanchias.

d “Der schwimmende Kopf”: Bloch, 1. c. From the form of the snout and the small size of the mouth the fish also acquires a
singular resemblance, which has struck many, to the human head.

e According to Campbell’s measurements of a specimen 236 cm. long (Proc. Nat. Hist. Soc. Glasg., vol. V, 1881 — 82, p. 178) this
percentage may sink to 24'7.

f In the specimen which Hasting examined, he found (1. c.) an asymmetry in the position of the eye (the right eye — like the right
pectoral fin — situated perceptibly higher than the left), which he proposed to explain by the habit possessed by these fishes, of lying at the
surface and swimming on one side. Cf. above, on Trachypterus , p. 319.

628

SCANDINAVIAN FISHES.

inferior margin lies in the line between the mouth and
the upper corner of the gill-opening, and the distance
between them and the tip of the snout is somewhat
greater than that between them and the gill-openings.
Externally they are rather small, somewhat oblong (el-
liptical) in form, with a longitudinal diameter of about
16 or 17 % of the length of the head or 1/3 of the
breadth of the interorbital space. The eyeballs them-
selves, however, are remarkable for their great size; in
Hastings specimen, which was nearly 1 1/s m. long, the
transverse diameter of the eyes was 6V2 cm. A most
remarkable point in the eyes of the Sunfish is the inner
eyelid, the mobile, nictitating membrane, pierced in the
middle, with which this fish is furnished according to
Cuvier and Owen". A similar nictitating membrane is
common, it is true, among’ the Sharks and finds its ana-
logue, to a certain extent, in the motionless, adipose
membrane of the Mackerels, the Gray Mullets, and other
Teleosts. But here this membrane is furnished with a
special closing muscle (sphincter) and five radiating
opening muscles, the latter of which originate from
the bottom of the orbit. The nostrils are very small,
the anterior in each pair elliptical and transversely-set,
the posterior round. They are set on about a. level
with the centre of the eyes and rather near each other,
the anterior one at a distance from the tip of the snout
that measures about % — 3/4 of the length of the latter.

The tongue, according to Kroyer, is very large and
fleshy, rather like the human tongue, but with very short
and flat papillae. Both the tongue and the palate are
sharp as a rasp, almost as sharp as the skin of the body.
In front of the tongue we find in the lower jaw (but
not in the upper) a large, transverse fold (velum). The
gill-rakers are only slightly developed arid not dentated.
The fourth branchial arch coalesces with the clavicular
arch* 6. The upper pharyngeals form on each side an
almond-shaped disk, set with three transverse rows of
5 — 8 pointed, scattered and narrow, but fairly large,
curved teeth; the lower pharyngeals are toothless.

The abdominal cavity is lined with a firm, hard, and
white peritoneum (fig. 156, p). The liver ( h ), which is
of a rounded, oval shape, occupies about half, or even
nearly the whole, of the abdominal cavity on the left
side, but has only a small lobe to the right. The gall-
bladder (fig. 157, vf) is large and has a long duct,

which opens into the stomach (v). The latter, which is
scarcely divided externally from the short oesophagus,
extends along the greater part of the dorsal wall of the
abdominal cavity, and under the end of this wall passes,
without external boundary but with an internal con-
traction, into the intestine, which first runs straight for-
ward and then, with several bends within its special
peritoneal (mesenteric) sac (fig. 156, i), coils backwards
and forwards, until it passes straight down into the
rectum (r) and pierces the thick wall of the abdominal
cavity at the vent (fig. 156, a). The entire intestinal
canal is about 372- — 5 times as long as the body, and
is especially remarkable for its thick wall. The spleen
is of a flat, rounded shape, dark reddish blue in colour,
and lies between the liver and the stomach. The urin-
ary bladder (fig. 157, vu) is large and of an elongated

v u au vu

Fig. 157. Intestinal canal of Ortliagoriscus viola, on a reduced scale.

After Cleland.

v, the stomach; u, the ureter; an, the fissure-like mouth of the
urethra in the wall of the urinary bladder; vu, the urinary bladder;
ov, the ovary; r. the rectum; i, the intestine; vf, the gall-bladder.

pear-shape. Its upper wall is pierced by the ureters
(it), subsequent to their union. In front of this bladder
lies the simple c ovary (ov), with the oviduct following
the front side of the urethra down to the common uro-
genital aperture behind the vent. The rudimentary air-
bladder, which was discovered by Costa, occupies an
oblique position, high up at about the middle of the
length of the abdominal cavity.

In the leading features of its coloration the Sunfish
shows itself to be a strictly pelagic fish, von Wright’s
figure tells us better than words the appearance of a
Sunfish about half a metre long; but if we compare
this figure with Costa’s figure of a somewhat older fish,
we find that quite a considerable variation may occur
in the colour. The ground-colour is bluish gray,

“ Cuvier, Leqons d’Anat. Comp. (ed. Dumeril) tome II, p. 434; (ed. Meckel) tome II, p. 437; Owen, Comp., Anat., Physiol., vol. I, p. 336.

6 According to Kr0Yer. According to Cleland it is free.

c According to Cleland and Wahlgren. According to Costa there are two distinct ovaries.

SUNFISH.

629

shading on the back into slate-gray or brown, down the
sides and on the belly into white, and with a silvery
lustre, brightest in old specimens. On the sides of the
head, in front of and below the eyes, as well as on the
chin and throat, we see irregular, clouded spots of red-
dish brown or gray. The dorsal and anal tins are brown,
blackish at the tips. The pectoral tins are also brown,
but, according to Costa, crossed by transverse bands of
grayish white. The pupil is bluish, the iris, according
to most descriptions, silvery with yellow or red inner
margin and with a spot of the same colour above and
below.

Both in the internal organs of the Suntish and on
its gills and skin there live an extraordinary number of
parasites, quite justifying, in the case of old specimens
at least, the appellation of living “ hotels garnis” which
has been bestowed upon them". The skin of large spe-
cimens is coated with a layer of tough slime, two-
tifths of an inch thick, in which there crawl lice of the
family Caligidce , and to which parasitic crustaceans of the
genus Penella — on which Cirrhipeds of the genus Concho-
denna may be found — and Trematods of the families
Tristomidce and Monostomidce attach themselves. The
gills are infested with Caligoids of the genus Lcemargus
and Trematods of the family Bistomidce, the eyes with
Filar ice. In the muscles and, in still greater numbers,
in the liver we find roundworms of the order Acantho-
cephali, and in the intestine, which is generally full of
a fetid, tough, grayish white mucus, roundworms of
the genus Ascaris and flatworms of the genera Bistoma ,
Tetrarhynchus, and Bothriocephalus.

Of the daily life of the Sunfish we have but little
information. All we know with certainty, is that it is
strictly a pelagic fish. It has been most often met with
lying on one side at the surface of the sea — we have
already made the same observation in the case of several
deep-sea fishes — to which the fish has been supposed
to ascend in fine weather in order to bask in the sun* 6.
Its lethargy on most of these occasions, however, may
well be due to its having wandered from its proper
home. Still, it has sometimes appeared nearer land and
swimming forward, with the dorsal fin above the water,
as Gossec relates of a Sunfish, 137 cm. long, that was

° Malard, Le Naturaliste, 2e serie, No. 46 (ler fevr. 1889).

6 Hence, perhaps, the name of Sunfish.
c Zoologist, 1852, p. 3579.
d Proc. Zool. Soc. 1849, p. 6.
e Zoologist 1850, preface, p. XI.

taken in the Bristol Channel oft' Ilfracombe. “It was
slowly moving at the time of its discovery, with a wav-
ing motion from side to side, “like a man sculling a
boat,” to use the comparison of the sailor who helped
to take it; the back-fin appearing above water. The fish
permitted the boat to come close up without exhibiting
alarm, nor was he even disturbed when her side came
into contact with his bulky person. The fellows made
a bowline-knot, and slipped it over his head, tightening
it before his dorsal and anal, so that the knot came in
the middle of his side. Thus they hauled him in, not
without a wetting, for with a flapping action of his
ample fins (again a sort of sculling) he scooped up the
water and threw it over them and into the boat. He
survived his introduction to the public about an hour.”
One of the largest specimens that have been found on
the English coast, 19 dm. in length, was caught off
Chesil Bank (Dorset) in June, 1846'2. This specimen
was more active from the very first, and swam straight
into the middle of the Mackerel-nets. The first net burst;
but in the outer net the progress of the fish was checked;
and with the help of 40 persons the catch was hauled
ashore. “Here it dashed about the pebbles, according
to the fishermen’s account, like a shower of grape. It
expired in about three hours, after uttering “hideous
groans,” like those of a horse dying of the staggers.”
During the summer and autumn of 1850 the Sun-
fish was observed in the English Channel more often
than usual; and from a comparison of these observations
Newman6 came to the conclusion that these fishes had
migrated thither from the west. The first find was re-
corded on the Cornish coast on the 9th of June, the
last off Dover on the 8th of September. During his
expedition with the Hirondelle in 1886 the Prince of
Monaco in the month of September fell in with a num-
ber of Sunfish in company, not far south of Great Sole
Bank, outside the entrance of the Channel. All of them
were of insignificant size. The larger Sunfish he found
in more scattered companies, but nearly always several
not far from each other, in the open Atlantic east and
north of the Azores. The Sunfish thus seems to be to a
certain degree gregarious, at least during youth. Though it
is usually sluggish and helpless, as it lies at the surface,

630

SCANDINAVIAN FISHES.

glittering in the sunshine or, in the darkness of night,
shining with phosphorescent lustre, motionless and drift-
ing with wind and tide — perhaps dazed or senseless
after a too sudden ascent from deep water, as is the
case with the true deep-sea fishes — , still it sometimes
shows considerable activity in its movements. On such
occasions it may be seen leaping high out of the water,
as Day tells us on the authority of Couch’s notes and
of the verbal statement of Dunn. A large Sunfish that
had been harpooned in the open Atlantic by the crew
of the Hirondelle, had such power in its fins that, when
it took to flight on feeling the blow of the harpoon, it
almost succeeded in drawing boat and all with it down
to the depths.

That the Sunfish also visits comparatively shallow
water off sandbanks, islands, or the coast of the main-
land, appears from the fact that its intestine-like stomach
has frequently been found to contain Zostera and alga?,
even those that only grow at a depth of a few fa-
thoms. Still, its proper food is probably animal, though
varied by these vegetable substances. Among the frag-
ments of these plants Kroyer found in its stomach and
intestine the scales of some fish, which he supposed
to have been the Sole, together with Sertularice and
Ophiurce. Wahlgren enumerates, besides the vegetable
substances, fragments both of crustaceans and shellfish
among the contents of the specimen he examined. Koren
and Collett found numerous Medusa; in the stomach
of the Sunfish; and this slimy food may perhaps explain
the composition of the fetid fluid, like thin gruel, of
which the intestine is generally full. On some occasions
the Sunfish has taken a hook baited with ordinary
Mackerel-bait or even with worms.

The Sunfish has long been known in the Medi-
terranean and was found on the coast of England (Corn-
wall) by Willughby and Ray. Bloch includes it among
the fishes of the Cape of Good Hope. Schlegel gives
an excellent figure of the species from Japan; and Ram-
say exhibited in London, as we have mentioned above,
an unusually large specimen of the Sunfish from Port
Jackson" (Australia). Its range is, therefore, extensive,
probably embracing all the tropical and temperate seas.
In Scandinavia it must be regarded as rather rare. In
Sweden and Denmark it has most often been found dead,
floating at the surface or cast ashore, during late au-
tumn or winter. Under these circumstances it has been
met with not only in the Cattegat, but also in the Sound
(a specimen 57 cm. long, in November, 1784, according
to Retzius), the Great Belt (a specimen about 18 dm.
long, secured by Fiedler in November, 1862), and
Flensburg Bay (Mobius and Hkincke, soon after 1860).
The Royal Museum has received from Bohuslan two
specimens about 43 cm. long, the first taken by Pro-
fessor F. Sundevall in October, 1834, the second by
Prof. S. Loven in 1856. Gothenburg Museum, accord-
ing to Malm, possesses 4 specimens from this locality,
between 40 and 55 cm. long and taken from October to
December inclusive. The specimen 11 dm. long described
by Wahlberg, stranded in November, 1866 off the fish-
ing-village of Traslof, just south of Varberg. On the coast
of Norway, according to Collett, the Sunfish is more fre-
quent in its occurrence and has been found alive, even near
the Swedish frontier, off Fredrikshald, in October, 1870.
Several large specimens have been taken off Bergen and
in Trondhjem Fjord. Even off the extreme north of Nor-
way, in Alten Fjord, a Sunfish 11 dm. long has been found''.

° G-t. Intern. Fish. Exbib., London 1883, Cat. Exhib. N. S. Wales, p. 46.

b While this sheet was in the press, I received through Mr. C. A. Hansson a fresh specimen, 40 cm. in length, caught off Lesund, at
the entrance of Dynekil, a fjord in the northernmost part of Bohuslan, on the 6th September, 1892. I here give the principal measurements
of this specimen:

Length of the body from the tip of the snout to the middle of the margin of the caudal fin 492 mm.

„ „ ,, head „ „ „ „ „ „ „ „ „ „ „ front margin of the gill-opening, in % of the length of the body 29.9 „

Greatest depth of the body (at the anal aperture), in % of the length of the body 66.7 „

„ thickness „ „ head (at the temples) „ „ „ „ „ „ „ ,, 18.9 „

Distance from the tip of the snout to the beginning of the dorsal fin, in % of the length of the body 67.1 „

>, ,, ,, ii n ii ii ii ii ii ii ii anal ,, ,, ,, ,, ,, ,, ,, ,, ,, 70.7 ,,

Length of the base of the dorsal fin, in % of the length of the body 19.8 „

„ „ „ „ „ „ anal „ „ „ „ „ „ „ „ „ 19.8 „

„ „ „ longest ray „ „ dorsal „ „ „ „ „ „ „ ,, „ 42.2 „

„ „ „ „ „ » ,, anal „ „ „ „ „ „ „ „ „ - — - - — 40.8 „

„ „ „ middle „ „ ,, caudal „ „ „ „ „ „ „ „ „ 12.6 „

Distance between the anal aperture and the beginning of the anal fin, in % of the length of the body 6.5 „

Length of the pectoral fins, in % of the length of the body - 12.6 „

,, „ „ snout, in % of the length of the head 49.3 „

„ „ „ orbit, „ „ „ „ ,, „ „ „ - — 17.1 „

„ „ „ postorbital part of the head, in % of the length of the head.. 38.4 „

Height of the orbit, in % of the length of the head 14.3 „

Longitudinal diameter of the cornea, in % of the length of the head — 13.2 ,,

Vertical „ „ „ „ „ ,, ,, ,, ,, „ ,, ,, _ 12.9 ,,

Breadth of the interorbital space, in % of the length of the head 55.8 ,,

The iris was blue, with white inner and outer margins of a faint golden lustre. The left eye (but not the right) had a white spot
in the lower part thereof. The pupil was of a little deeper blue than the iris.

SCLERODERMS.

631

The Sunfish is seldom, if ever, used as food. All its
various parasites and the slimy coat of its body render
it repulsive to the ordinary fisherman; and the phos-
phorescent light that radiates from it in the dark, has
caused it to be suspected of injurious properties. Ac-
cording to Risso", however, the liver is eaten in the
north of Italy, though no great value is set upon it;

Fam. B A L

Scleroderms with hard scales or scale-like plates ( not co t
or less fully developed) ivith spinous rays. Caudal fin

This family is composed of fishes of far less sin-
gular appearance than the preceding one. Its members
are the least abnormal of the Plectognates; and their
primeval characters are veiled in a great number of
them by an undeniable beauty of colour — several of the
Balistoids belong to the most handsomely coloured fishes
of the modern period. The dermal covering, however,
varies considerably not only between genus and genus,
but also in several forms during the changes of growth.
Sometimes the scales are extremely small and hardly
visible to the naked eye; and the juvenile forms may
be armed with spines, especially on the sides of the tail,
which remind us of the equipment of the Acanthuri,
but in many forms disappear with age. The dermal
covering is not the only character that distinguishes
these fishes from the other Scleroderms, the family of

and in The Field (4th Feb., 1882) we are told how a
person was cheated into the belief that he was eating
the most delicious turtle soup, which was really made
of the flesh of the Sunfish. The only economical value
of the Sunfish lies in the oil into which the flesh and
liver are boiled down, or the glue extracted from the
bones and gristle.

I S T 1 1) M.

itinuous carapaces) and ivith two dorsal fins, the first ( more

with 10 branched rays, and one simple ray at each margin.

the Coffer-fishes ( Ostraciontidce ); they are also marked
by the presence of an anterior, spinous-rayed dorsal fin
— though this fin may sometimes consist of a single
spinous ray — , by the compressed form of the body,
and by their stronger, rodent-like jaw-teeth.

The family belongs properly to the tropical seas —
Jordan and Gilbert have estimated the number of the
species at about 100 - — and may be divided into three
subfamilies: the Triacanthince, furnished with ventral
fins and with the jaw-bones only loosely united; the
Monacantliince, without ventral fins, or with only one
rudimentary ventral ray, with extremely small scales,
with at most two rays in the first dorsal fin, the mem-
brane of which is also rudimentary, and with only 6
teeth in the lower jaw; and the Balistince.

Subfamily BALISTIN M.

Ventral fins wanting or represented merely by one fixed or mobile spine on the end of the pelvic bones, which are
confluent. First dorsal fin with three spinous rays. Caudal fin rounded, or ivith S-shaped hind margin. Scales
middle-sized or large, more or less plate-like. Upper jaw with 8 teeth in an outer row and 6 in a dense row
within ( behind ) the former. Lower jaw with 8 teeth in a single row. Both the upper and the lower pharyn-
geals furnished with teeth. Branchiostegal rays 6 or 5. Vertebrae 17.

Linnasus called these fishes Filareb (Filers), a Swe-
dish expression for the later name of Scleroderma.
In English they are called File-fishes, from the rough
front surface of the first ray of the anterior dorsal fin.

Even by the external characters this subfamily is
easily distinguisTied from the other Balistoids. The com-
paratively large scales of the body separate it both from
the Triacanthince and the Monacantliince ; in the struc-

ture of the first dorsal fin it ranks between them; in
the absence of ventral fins and in the rounding (at
least at the middle of the margin) of the caudal fin it
has characters in common with the Monacantliince , which
have fewer jaw-teeth, and in which we find either one
single row of teeth on the lower pharyngeals or no teeth
at all on the pharyngeal bones. In a Melichthys (Ba-
buniva from Ascension we find the lower pha-

a Eur. Mer., tom. Ill, p. 174.
b Mas. Ad. Frid., tom. I, p. 57.

80

Scandinavian Fishes.

632

SCANDINAVIAN FISHES.

ryngeals like branchial arches, only shorter and thicker
than the ceratobranchial 1 tones, and furnished with
three rows of teeth, the outermost consisting of cylin-
drical teeth, the middle row of pointed ones, but all
these teeth considerably shorter than those of the inner-
most row, which are larger, compressed, and claw-shaped.
Two of the three upper pharyngcals are thick and united
into one bone, which bears two rows of claw-shaped
teeth, at the base broad but compressed, at the tip
sharply pointed. The skeleton of this species contains
7 abdominal and 10 caudal vertebrae, including the
urostyle.

Within the subfamily of the Balistince, of which
about 30 species have been described, Bleekee esta-
blished 4 genera, differing in the form of the base
of the caudal fin and in the form and colour of the
jaw-teeth. The forms are so closely allied, however,
that a division into genera can be defended only as an
expedient to facilitate a general survey of the subfamily.

Among the peculiarities of the skeleton, besides
those mentioned above, the strong supporting apparatus
of the first dorsal fin, with the articulation of the first
spinous ray, which is the principal defensive weapon of
these fishes, is especially remarkable. The supporting
apparatus, which is evidently formed by the coalescence
and more advanced development of the elements of the
first interspinal bones, consists of three bones", the hind-
most of which is wand-like and lies in a backward and
downward direction, so that its hind extremity rests
against the lower part of the first interspinal bone of
the second dorsal fin — which bone is here supported
on the front of the upper spinous process (neural spine)
of the fifth abdominal vertebra — while the anterior
(upper) extremity articulates firmly with the lower
end of the middle bone in the supporting apparatus.
This bone is triangular, with one angle directed down-
wards, but longitudinally cloven, its two wing-like
halves forming an angle open above and being firmly

united in front and below to the first and largest bone
in the supporting apparatus. This last bone is can ali-
en late, but its long bottom is sharply carinated, thus
forming together with the middle bone a kind of boat,
with the bows pointing backwards and split (open) and
with a large, elliptical hole — crossed, however, by a
narrow, transverse bridge of bone — in the posterior
part of each side, but with the middle of its stern
(turned forward in the fish) touching the middle ridge
of the cranium ( s-pina occipitalis) above the hind part
of the orbits. The hind part of the gunwale (in the
fish the fore part) is furnished with articular cavities,
one on each side, for the lateral articular processes of
the first spinous ray, and from the middle of the bottom,
in the stern, rises a tap-like process with rounded
head, over which glides the base of this ray, with
its concave centre. In front of this process (i. e.
behind it in the longitudinal direction of the fish) rises
from the keel of the boat another tap-like and round-
topped process, over the head of which the cloven base
of the second spinous ray glides to and fro. The back
of the first spinous ray is concave (canaliculate). The
front of the second spinous ray is tumid at the base,
and this swelling drops into the groove of the first spi-
nous ray, when the rays are erected. In this manner
the first ray is locked fast, and any attempt to force it
back merely exerts a pressure on the second ray, which
is thus kept all the more firmly fixed to its tap-like
articular process, and effectually hinders any backward
curvature of the first ray. In order to attain this result
the second spinous ray must be drawn back, an opera-
tion which is performed partly by means of special
muscles, partly by a ligament which unites this ray
within the fin-membrane to the third spinous ray* 6. In
the Monacanthince , which are without the third spinous
ray, the supporting apparatus is also less developed;
and in the Triacanthince the first interspinal bones are
only slightly metamorphosed.

“ Cf. Hollard, Ann. Sc. Natur., troisieme serie, Zoolog., tom. XX, p. 102; Sorensen, Om Lgdorganer hos Fiske, p. 50.

6 Hence, according to Johnston ( De Piscibus, p. 110), the generic name of Batistes (catapult). The Italians, he says, called one of
these fishes pesce balestra , because a slight pull at the third spinous ray was enough to overcome the rigidity of the first ray and to depress
it, just as a light touch of the hammer or trigger in the lock of a cross-bow sufficed to discharge the weapon. This name reappears in the
English name of Trigger- fishes as applied to these forms. Batistes was introduced into zoological nomenclature, without any special explana-
tion, by Artedi, first in Seba’s Thesaurus , tom. 3, p. 63 and again in Gen. Piscium, p. 53.

FILE-FISHES.

633

Genus BALISTES.

Peduncle of the caudal fin laterally compressed . Jaw-teeth white , those in the lower jaw and in the outer row in
the upper jaw obliquely incised or claw-shaped; no prominent canines. Lips fleshy and naked. First dorsal fin
completely separated from the second , its membrane falling short of the first ray of the second dorsal fin.

Rays of the second dorsal fin less than 30.

Such are the characters by which Bleeker" has
defined the true File-fishes, which are distinguished
from all the other members of the family by their
comparatively thick and also naked lips6. The greater
number of these forms are furnished behind the gill-
openings, on each side above the angle of the pectoral
fin, with the plates whose connexion with the air-

bladder has been pointed out, as Ave have mentioned
above, by Mobius. One or perhaps two species are,
hoAvever, knoAvn, Avhich are Avithout these special axillary
plates, and on this account Bleeker proposed to estab-
lish a subgenus Canthidermis, borroAving the name,
but not its application, from Swainsonc. To this sub-
genus Ave should then refer

THE SPOTTED FILE-FISH (SW. SPATTBALISTEN d OR FLACKFILAREN e).

BALISTES MACULATUS.

Fig. 155.

No special axillary plates. Scaly covering of the cheeks even , unbroken by any longitudinal , naked strips. A lon-
gitudinal hollow in front of each eye just below the naked , oblong depression in which the two small , oval nostrils

are situated f. Tail unarmed , but the scales on the sides of the body furnished during youth on the free part of

their surface with rows of pointed spines and at the middle (at the tip of the next scale in front) with one rather
more prominent spine; in older specimens all the scales granulated on the free part of their surface, and the scales
on the sides of the body retaining only blunt stumps of the spines , among which those at the middle of each scale
are larger and somewhat elongated , together seeming to form longitudinal rows of low carince. About 32 scales
in an oblique transverse row from the vent to the first dorsal fin. The quadrangular, elongated scales of the ventral
side groper He in about 14 obliquely longitudinal rows, the upper ones containing about 17 scales between the
base of the pectoral fin and the anal region. Length of the head- in adult specimens 1/i, breadth of the inter orbital
space about 7io> length of the snout about 15 %, distance between the first dorsal fin and the tip of the snout 28
or 29 %, distance between the second dorsal fin and the tip of the snout 49 or 50 %, distance between the anal
fin and the same point 55 or 56 % of the length of the body. Second dorsal and anal fins in front high9 with the
posterior outer (resp. upper or lower) margin concave. Length of the caudal fin at the middle equal to that of the
snout’1. Length of the pectoral fins 7u of that of the body. Pelvic spine short, sometimes firmly united to the pelvis.
Greatest depth of the body in young specimens rather more than x/21 in old rather less than 1/3 of its length.
Least depth of the body equal in adult specimens to the length of the pectoral fins. Greatest thickness equal to the
length of the snout. Coloration brown or blue, with light blue spots, somewhat smaller than the eyes, scattered over
the whole body, the second dorsal fin, the caudal fin, and the anal fin. In young specimens the spots are smaller,

indistinct, and interspersed with darker spots. The fish is sometimes plain brown or black.

“ Atl. I dull. Ind. Or. Neerl., tom. V, p. 98.

b Linnaeus’s Batistes ringens is thus excluded from this genus; in its case the lips are extremely thin and lie within the scaly dermal
fold that extends outside the roots of the teeth in both jaws.
c Nat. Hist. Fish., Amph., Rept., vol. II, p. 194.
d Malm, Gbgs , Boh. Fn.
e Lilljgborg, Sv., Norg. Fisk.

f The anterior larger than the posterior; between them a longitudinal dermal flap, which partly covers the posterior nostril.
g The length of the longest ray (the 6th or 7th) in the former fin about 1 7 1 , 2 / of that of the body, in the latter fin (the 5th or

6th ray) 16 or 17 % thereof, or respectively 73 % of the base of the second dorsal fin and 84 % of the base of the anal fin — in each case

in specimens between 27 and 30 cm. in length. In younger specimens these fins are more rounded.

A In young specimens the caudal fin is sharply rounded; in older ones the outer corners are prolonged into pointed, but short lobes.

634

SCANDINAVIAN FISHES.

R. hr. 6; D. 3|24— 26: A. 21-24; P. 15; ( V.) 1; 0. 1 +
10 + 1 ; L. I at. 46 — 56.

Syn. Guaperua longa , Lister in App. ad Willughb., Hist. Pise.,
p. 21, tab. I, 20.

Balistes maculatus, Bl., Naturg. Ausl. Fiseli., pt. II, p. 25,
tab. CLI; Blkr, Atl. Iclith. Ind. Or. Ne'erl., tom. V, p. 122,
tab. CCXVIII, fig. 4; Gthr, Cat. Brit. Mus., Fish., vol. VIII,
p. 213; Malm, Ofvers. Vet. Akad. Forh. 1875, No. 7, p. 8;
Gbgs., Boh. Fn ., p. 599; Day, Fish. Ind., p. 687, tab.
CLXXV, fig. 3; Fish. Gt. Brit., Irel., vol. II, p. 267, tab.
CXLV ; Lillj., Sv., Norg. Fish., vol. Ill, p. 420.

Balistes hrevissimus + Bal. angulosns (?) + Bal. maculatus +
Bal. longissimus, Hollard, Ann. Sc. Nat., 4 ser., Zool.,
tom. I, pp. 56 — 62, tab. 3, figg. 1 — 4.

The Spotted File-fish attains a length of at least 4
dm., and is distinguished, when adult, among its con-

as a rarity, having wandered to our waters probably
in the same manner as Antennarius histrio , merely a
casual visitor to Scandinavian regions.

In Scandinavia the File-fish, of course, possesses no
economical importance; and it is not used in any manner, to
the best of our knowledge, in its true habitat. The other
European species of the genus, Balistes capriscits, a form
belonging to the warmer regions of the Atlantic and the
Pacific, not very rare in the Mediterranean, and once
or twice met with on the coasts of Great Britain and
Ireland, is also one of the least useful fishes. According
to CanestrinC it is not eaten in Italy. The flesh of the
whole genus, as we have mentioned above, is also sus-

Fig. 155. Balistes maculatus, 1 2 the natural size. Valparaiso; the F.ugenie Expedition.

geners by its elongated form and by the comparatively
great height and almost falciform shape of the second
dorsal and the anal fins. It is one of the first-
discovered species of the genus, and its range ex-
tends over the tropical and subtropical seas of both
hemispheres. In the West Indies it has long been known,
together with another species, under the name of Sa-
lt itco; and from these regions it sometimes strays to
the coasts of North-western Europe. It has once been
found, according to Gunther, off Polperro in Coni wall;
and in Uddevalla Museum is preserved a stuffed speci-
men, 272 mm. long, which is said to have been taken
during the autumn of 1857 off “Saltk&llan, at the head
of Gullmar Fjord, whence it was forwarded fresh to the
said museum”. Of its habits we have no special infor-
mation. In the Scandinavian fauna it is interesting only

pected of possessing poisonous properties. In those
tropical regions where the species are most plentiful,
the natives suppose that these poisonous properties are
due to the food of the fishes and are especially dangerous
when the fishes live on coralline animals. Day describes* 6,
according to Meunier’s observations at Mauritius, the
symptoms caused by the eating of File-fishes. The
poison operates immediately upon the nerves of the
stomach and causes violent spasmodic convulsions in its
walls. Soon afterwards, these convulsions spread to the
muscles of the whole body. The tongue swells, the
eyes stare, the breath grows difficult, and the patient
dies in the most acute agony. Powerful emetics, fol-
lowed by oleaginous substances and emulsions, are the
best antidotes. Other species of the genus, however,
enjoy a good reputation.

a Fauna Italica, pt. Ill, Pesci, p. 147.

6 Fishes of India, p. 686.

HEMIBRANCHS.

635

PHYSOCLYSTI HEMIBRANCHII.

Physoclysts with free jawbones and pectinated gills, but with the branchial arches more or less imperfect and

with large and externally visible inter clavicles.

Here, as in the case of the Plectognates, the sy-
stematic difference from the other Physoclysts is not at
all too great to admit of the arrangement of these fishes
in a series of families corresponding to the series already
adopted among the Physoclysts and ranged beside these
series. This was also the approximate rank assigned to
them by the writer who first treated of the combination
of these fishes into a systematic whole: in 1861, in his
arrangement of the Acanthopterygians", Gunther adopted
a twelfth division, Acanthopterygii g aster ostei formes, in-
cluding the Sticklebacks and Flute-mouths with their
nearest relatives, and a thirteenth division, Acanthopte-
rygii centriscif ormes , containing the family of the Trum-
pet-fishes. After Parker’s* 6 demonstration in 1868 of
the morphological significance of the interclavicles of
the Sticklebacks as traces of the Ganoid type, Cope0
united Gunther’s two divisions into an order Hemi-
branchii, “connecting the Lophobranchii with ordinary
fishes.”

All these fishes have abdominal ventral fins, and
are thus ranged comparatively low in the scale of deve-
lopment of the Physoclysts. Their principal character
also lies, as we have shown above, in the circumstance
that several of the bones which in more typical Phy-
soclysts entirely partake in the structure of the endo-
skeleton, are here wholly or partly dermal, appearing
in the form of growths belonging to the exoskeleton.
This structural feature is fairly common among fishes.
Scales, differing usually in size, shape, or thickness from
the other scales of the body, are in one fish the pre-
cursors of a still further alteration in a kindred form,
a difference which, during the development of the form-
series, leads to the result that the modified scales creep,
so to speak, into the body and join themselves to parts
of the endoskeleton, and finally, when the transforma-
tion has reached its highest point, lie like covering
(membrane) bones outside other bones, or even enter in

the form of independent bones into the endoskeleton.
A great portion of the skeleton of the Vertebrates has
originated in this manner. The most beautiful and most
perfect example of this is given by the human clavicles,
the development of which has been traced by Parker
(1. c.) from their first origin, at which period they are
morphologically analogous to two lateral plates in the
Sturgeons or the Cuirassed Siluroidsk The intercla-
vicles which we have above remarked in certain Plecto-
gnates, are of similar origin. They occur, it is true,
in several other Physoclysts as well, but in an extre-
mely reduced form and as parts of the endoskeleton.

Here, in the Hemibranchii, as well as in the Lopho-
branchii, they still preserve distinctly the character of
dermal plates. Thus, in the Three-spined Stickleback
(fig. 156, C) for example, they have the form of two
parallel, ventral scutes, contiguous in front, separated
behind, on the outside granulated and striated, like the
other dermal plates of the body, on the inside support-
ing, in front, the lower ends of the clavicles, and be-
hind, the loAver margins of the coracoid bones or more
correctly, according to Parker, of their covering bones,
the procoracoids, which are also said to be of dermal
origin.

rfhe same relative position to the rest of the
skeleton is here occupied by the pelvic bones or rather,
according to the same interpretation, by the covering
bones of the pelvic bones proper. These bones lie in
the Sticklebacks immediately behind the interclavicles,
or even project some distance between them; and in the
Three-spined Stickleback (fig. 157, A, vs) they are
marked by the large, ascending process that meets the
lateral plates of the body in the skin. The character
which in Cope has been chosen to give the Hemibranchii
their name, lies in the absence or persistent cartilagi-
nous structure of one or more of the upper parts of
the branchial arches (the epibranchial and upper pha-

“ System. Synops. Fam. Acanthopt. Fish., Cat. Brit. Mus., Fish., p. VIII.

6 Ray. Soc., Shoulder Girdle and Sternum, p. 40.

c Ichthyology of the Lesser Antilles, Trans. Airier. Phil. Soc., Philad., n. ser., vol. XIV, pp. 456 et 457.
d Cf. Smitt, Ur de hogre djurens utvecklingshistoria, p. 218.

636

SCANDINAVIAN FISHES.

ryngeal bones) on each side. Another character of more
general validity — which indicates the low rank of these
fishes among the Teleosts — is the imperfect ramifica-
tion and, in some cases, the defective transverse divi-
sion (articulation) of the soft fin-rays.

The form of the body is very variable in the Hemi-
brancliii. The Sticklebacks, which possess the ordinary,
most typically piscine form, are so like the Horse-
Mackerels that they have been ranged beside the latter
by many systematists. All the others are of more
singular form, elongated like the Garpikes or laterally
compressed, and all with the head prolonged into a long,
tubular or conical snout, at the end of which the small
mouth is situated. This prolongation, however, does
not depend here, as in the Garpikes, on the elongation
of the jaws, but corresponds more exactly to the struc-
ture we have remarked above in certain Plectognates,
here affecting the ethmoid bone, the three pairs of
pterygoid bones, the preoperculum, and the interoper-
culum, which together form the greater part of the
long rostral tube, special suborbital bones, on the other

Fig. 157. Gasterosteus aculeatus, tracliurus , natural size. From
Eckernforde Bay. A: from the side; B: from above; C: ventral and
anal regions; ds, dorsal plates; vs, plate of the ventral fin (pelvic
hone), in A its ascending lateral process; a, vent; at, anal spine;
vt, ventral spine. After Heincke.

i. 1. 2.

Fig. 156. Shoulder-girdle of Gasterosteus aculeatus; magn. 9 or 1 0 diam.; after Parker.

A: Bones of the left side, seen from without, pt, posttemporal bone (Ganoid plate of the first dermal ring); 1.1.2, first lateral plate (Ganoid
plate of the second dermal ring); 1.1.5, fourth lateral plate; cl, clavicles; sc, scapula; sc./, scapular fenestra; b.l , first basal bone of
the pectoral fin; b.4, fourth basal bone; p.cr, ossified part of the coracoid bone (praecoracoideum, according to Parker); i.cl, inter-
clavicles.

B: Section of the lower part of the clavicular arch, to show the cartilaginous part of the coracoid bones (e.cr, epicoracoideum, according to

Parker) between the clavicles.

C: Interclavicles, seen from the ventral side, with the anterior end up.

STICKLEBACKS.

637

hand, being wanting. We find a trace of this elongation
in one of our Sticklebacks, the Fifteen-spined species.
The other families of the Hemibranchii (the Flute-mouths,
Fist ula ri idee, and the Trumpet-fishes, Centriscidae) are
also distinguished from the Sticklebacks by the elonga-
tion and, in some cases, the coalescence of the first
four or even the first six vertebras. In this manner
the forepart of the body receives an increase of strength
which is still further advanced in the Trumpet-fishes
by the extraordinary development of the anterior inter-
spinal bones and by their dorsal expansion into a more
or less perfect shield, which in the Chinese Amphisile

has been compared, not without reason, to the carapace
of the turtles.

The Flemibranchian group is not particularly rich
in forms. It contains only a score of known and
described species. Some of them are so like each other
that their right to specific rank must still be regarded
as rather dubious; others, again, are so different that
a, comparatively great number of genera have been
established. According to Gill the group contains 11
genera, distributed among 6 families. Most of the fa-
milies live in the tropical seas; the Scandinavian fauna
possesses only one.

Fam. gasterosteid m.

The anterior vertebrae of normal structure. Ventral fins with one spinous and one or two soft raps. In front of
the soft-raped ( second ) dorsal fin 3 — 15 free spinous raps ( not united bp a fin-membrane) ; in front, of the anal
fin one spinous rap. Suborbital ring united to the preoperculum, but without rigid, connexion. Teeth on the inter-
maxillarp bones and in the lower jaw; palate and tongue unarmed. No scales proper. Base of the caudal fin
narrow, but, depressed. Branchiostegal raps 3 on each side. Air-bladder simple. Pseudobranchiae more or less

well-developed ; pploric appendages wanting or rudimentarp.

Within this family, which was founded by Gun-
ther/" and originally ranged by him at the head of the
system of the Teleosts, are collected the Hemibranchii
in the exterior of which the ordinary piscine form is
most typically persistent. Still these fishes show ten-
dencies of transition to the carapaced Trumpet-fishes and
also to the elongated Flute-mouths. An expression of
this may be found in the proposed generic division of
the family. One species, our common Three-spined
Stickleback, with its well-developed dorsal plates and
shorter form of body, especially of the snout, has been
employed as the type of a distinct genus, the true
Stickleback genus ( Gasterosteus ), while another species,
the Fifteen-spined Stickleback, with its more elongated
form and, in particular, its longer snout, is the single
representative of a second genus, Spinachia or Gastraea.
The latter genus is approximated to the Flute-mouths
not only by the form of the body, but also by the
structure of the pelvic apparatus, which in the Fifteen-

spined Stickleback is broken up into its two halves,
which merely meet each other by means of foliate
processes in the median line of the belly.

All these fishes are of small size, but have long
been famous both for their beauty and for the high
degree of instinct that guides them in the propagation
of their species. In spite of their insignificant size
they are not destitute of importance. The gregarious
habits of some species collect them in such enormous
masses that they may easily be applied to industrial
purposes. On the other hand, they are destructive
enough, for their fatness is gained to a great extent
at the expense of the fry and eggs of other fishes.

The range of the family embraces the northern parts
of both hemispheres. Some of these fishes occur both
in fresh water and in the sea or, at least, in brackish
water; but the first genus we shall here describe, is
exclusively marine.

Cat. Brit. Mas., Fish., vol. I, p. 1.

638

SCANDINAVIAN FISHES.

Genus GASTR-^EA. \

Body elongated (, greatest depth less than V2 the length of the head), in the abdominal region pentagonal and only
slightly deeper than broad; the forepart (in front of the anal spine) shorter than the hind part , including the
caudal fin. Number of f ree spinous rays in front of the soft-rayed dorsal fin at least, 13. Ventral fins inserted
at the middle of the long pelvic bones, which lie at the lower lateral edges of the belly and are without any firm

osseous connexion with each other. Jaw-teeth compressed.

As we have mentioned above, this genus contains only one species:

THE FIFTEEN-SPINED STICKLEBACK (sw. tangspiggen).

GASTRiEA SPINACHIA.

Plate XXVIII, fig. 5.

Distance between the ventral spines and the anal spine less than 3 * */5 of the length of the head or than 2/5 of the
distance between the former spines and the tip of the snout, which distance is more than 72 % of that between
the anal spine and the tip of the snout. Free dorsal spines of uniform, size, except the hindmost , which is also
the largest. Branchiostegal membranes united underneath into a broad, free dermal fold across the isthmus.

R. br. 3; D. XIV6— XV|6 c— 7 ; A. I[5— 7 ; V. l|2; P. 9 —
10d; C. x+l + 10+l+a?; L. lat. 40; Vert. 40 — 42.

S 'yn. Aculeatus vel Pungitius marinus longus, Schonev., Ichth. Slesv.

IIols., p. 10: Aculeatus marinas major , Id., ibid., tab. IV,
fig. 3 ; Gasterosteus aculeis in dorso quindecim, Art., Ichth.
Gen., p. 52; Syn., p. 81; Centriscus , A, 1, (aliis Spinochia)
Klein, Hist. Pise., Miss. IV, p. 48; StrSm, Sondm. Beskr.,
part. I, p. 316.

Gasterosteus Spinachia, Lin., Syst. Nat., ed. X, tom. I, p. 296;
Fn. Suec., ed. II, p. 119; Bl., Fisch. Deutschl., part. II,
p. 84, tab. LIII, fig. 1; Retz.. Fn. Suec. Lin., p. 339;
Cuv., Val., Hist. Nat. Poiss., vol. IV, p. 509; Ekstr., Vet.
Akad. Handl. 1831, p. 305; Nilss., Prodr. Ichth. Scand.,
p. 87; Ekstr., v. We., Skand. Fisk., ed. I, p. 21, tab. IV,
fig. 3; Sundev., Stockh. L. Hush. Sallsk. Handl., H. VI
(1855), p. 79; Nilss., Skand. Fu., Fisk., p. 112; Lindstr.,
Gotl. L. Hush. Sallsk. Arsber. 1866, p. 15 (sep.); Gthr,
Gat. Brit. Mus., Fish., vol. I, p. 7 ; Sauv. ( Gastrcea ) Nouv.
Archiv. Mus. Par., tom. X (1874), p. 36; Bncke ( Gasterosteus ),
Fisch., Fischer., Fischz. W., 0. Preuss., p. 76; Handb.
Fischz., Fischer. (M. v. d. Borne), p. 99; Day, Fish. Gt.
Brit., Irel., vol. I, p. 246, tab. LXVIII, fig. 5; Mela, Vert.
Fenn., p. 280, tab. IX.

Spinachia vulgaris , Flmng, Brit. Anirn., p. 219; Kr., Damn.
Fiske, vol. I, p. 193; Mgrn, Finl. Fiskf. (disp. Helsingf.
1863), p. 16; Coll., Forh. Vid. Selsk. Christ. 1874, Tillaegsli.,

p. 14; Winth., Naturh. Tidskr. Kbhvn, ser. 3, vol. XII, p.
5; Mok., Hist. Nat. Poiss. Fr., tom. Ill, p. 171; Hansen,
Zool. Dan., Fiske, p. 31, tab. V, fig. 4; Mob., Hcke, Fisch.
Osts., p. 64; Storm, N. Vid. Selsk. Skr. (Trondhj.) 1883,
p. 15; Lillj., Sv., Norg. Fn., Fisk., vol. I, p. 370.

Spinachia Linnei, Malm, Gbgs, Boh. Fn., p. 373.

Obs. The name of Spinachia or, as Klein writes it, Spinochia
is, according to Cuvier and Valenciennes, a Latinized form, dating
from the Middle Ages, of the French epinoche.

The Fifteen-spined Stickleback is distinguished from
its relatives by the elongated form of the body, with
the greatest depth, at the base of the ventral tins, about
1072 — 872 % of the length. From the head to the
vent the body is pentagonal, then quadrangular, but
more and more depressed behind, until at the base of
the caudal tin it suddenly becomes laterally compressed.
The length of the body may rise, at least in the fe-
males, to 1 8 1/2 cm., but is usually between 13 and
15 cm.

The elongated head measures between 1ji and 1/f
of the length of the body, and in old specimens the

a In order to enable us without tautology to retain the Linmean specific name of spinachia, we here adopt the generic name proposed

by Sauvage in 1874, a Latinized form of Cuvier’s Les Gastres, in spite of the fact that, even in the first edition of his Regne Animal (1817),

the latter author adopted Spinachia as the name of the subgenus which Fleming ( British Animals, 1828) raised to a generic rank.

6 Sometimes 13, according to Heincke; sometimes 17, according to Day.
c Sometimes 5, according to Lilljeborg.
d Sometimes 11, according to Khdyer.

c 25'6 — 2 1 ’ 5 % of the length of the body, according to our measurements of specimens between 53 and 172 mm. long.

FIFTEEN-S PIN ED STIC IvL EB A C K .

639

length of the snout is exactly equal to the distance
between the hind margin of the eye and that of
the operculum. This elongation of the snout is, how-
ever, a character of growth, for in fry 21 mm. long
the length of the snout is about 60 % of the post-
orbital length of the head, in young specimens between
50 and 75 mm. long about 90 % thereof. The elonga-
tion is produced, as we have mentioned above, by the
growth in a forward direction of the ethmoidal region
and the palatine parts as well as of the preoperculum
and interoperculum. At the same time the quadrate
bone with the articular knob for the lower jaw moves
forward, and the cleft of the mouth, which up to the
present has been comparatively larger and almost hori-
zontal, becomes relatively smaller and rather more
ascending. With this elongation is also connected the
relative diminution of the eyes; in a specimen 21
mm. long their longitudinal diameter is equal to
the length of the snout, in specimens 25 mm. long
about 3/i, in specimens 50 mm. long about 2/3 , and in
a female 172 mm. long only slightly more than 1/3
of the same length.

The head is quadrangular, tapering in a. forward
direction, with perpendicular sides and only slightly
convex occiput. The snout is depressed and blunt,
with the upper profile slightly concave. The eyes are
round, and their superior margin lies on a level with
the flat or somewhat hollow forehead, or rises a very
little above the latter, the breadth of which, at the
anterior margin of the eyes, is equal to their diameter.
The small, round nostrils, only one on each side, lie
half-way along the snout. The edges of the mouth are
furnished with fleshy lips, lobate on the sides of the
upper jaw; the lower jaw projects beyond the upper,
which may, however, be protruded by means of the
long nasal processes of the intermaxillary bones. The
length of the lower jaw is about 1/g that of the snout.
When the mouth is closed, the small maxillaries, the
length of which is about 1/g that of the intermaxillaries,
drop under the projecting preorbital bones. The jaw-
teeth are of equal size, chisel-shaped, and most of them
notched at the margin; they are set in front in several
(3 or 4) fairly regular rows, behind in a single row.
The pharyngeal teeth are pointed, and above they form
on each side an almond-shaped card, obliquely divided
into an anterior, smaller patch and a posterior, larger
one. The gill-rakers are setiform, 10 or 11 in number
on the first branchial arch. The pseudobranchiae lie

high, on each side within the articulation of the hyo-
mandibular bone; each of them is made up of five,
digitiform lamellae, set in a transverse row. The pala-
tine fold of the upper jaw is fairly large, the corres-
ponding dermal fold in the lower jaw less developed.
The external bones of the head are granulated and
striated. The operculum is triangular, with the hind
inferior side somewhat convex, the upper side soine-
Avhat concave. The suboperculum has a short, anterior
branch in a vertical direction and a longer, posterior
one, broader below, tapering behind, and curved like
a sabre; the latter forms the whole opercular margin
along the lower posterior side of the true operculum.
The preoperculum forms a right angle, with the an-
terior, horizontal arm, which is coasted by the subjacent
interoperculum, considerably elongated, as we have
mentioned above. The two posterior suborbital bones
are attached to the upper (inner) margin of this arm,
the posterior firmly, the anterior very loosely, thus
forming a cuirass for the cheeks, but only under the
eyes. The hind part of the cheek (between the eye
and the vertical arm of the preoperculum) and the space
between the long preorbital (first suborbital) bone and
the anterior part of the horizontal arm of the preoper-
culum (a strip along the side of the snout, behind the
corner of the mouth) are covered by the glossy skin
alone. The gill-openings extend above the operculum
for a distance measuring about V3 of its length. Under-
neath the branch iostegal membranes are united into a
broad, free fold, the hind margin of which coincides
with the anterior end of the interclavicles.

The form of the trunk is determined principally
by the granulated plates that follow the dorsal margin,
the lateral line, and the ventral margins. The dorsal
margin is occupied by the plates that have been deve-
loped in the skin on the tops of the interspinal bones.
Nearest to the occiput there lie two flat plates, one
behind the other, followed by a row of concave plates,
generally 15 in number, each of which is furnished
with a free spinous ray with a small, triangular mem-
brane behind it. Next follow seven plates, which
support the true, soft-rayed dorsal fin. On the dorsal
margin of the tail this row is continued by fastigiated
and carinated plates, growing more and more flattened,
and finally smoothed out and indistinct. The ventral
edges are formed in the forepart of the body by the
interclavicles, which are comparatively small in this
species, and which by their arcuate union in front

Scandinavian Fishes.

81

640

SCANDINAVIAN FISHES.

together form an elongated horse-shoe, equal in length
to the space between 9 of the free spinous rays on the
back. Within each arm of this horse-shoe projects a
small portion of the anterior extremity of one of the
pelvic bones. The two pelvic bones are also elongated
and narrow in this species, their length being about
equal, in adult specimens, to the space between 10 or
even 11 of the free dorsal rays. They are thin and
almost flat, the inner surface alone being slightly hol-
lowed into a broad channel; and they are strengthened
along the middle by two ridges, an outer and strong
one and an inner and weaker, which is interrupted
at the middle. Above, towards the side of the body,
their thin margin is only slightly broadened at the
middle, and they do not send out any special branch
in this direction. Inwards, on the other hand, at the
lower ventral edge, at the middle of their length and
a little further back, they send out a flat and thin
process, somewhat wider towards the tip, which meets
the corresponding process of the pelvic bone from
the other side of the body, though without entering into
any firm osseous connexion with it. The pelvic bones
extend very nearly to a line with the vent; and here
this angular structure of the ventral edges ends, to be
replaced at the base of the anal fin and on the under
surface of the tail by a row of plates analogous to that
we have just seen on the dorsal side. The plates of
the lateral line, about 40 in number, fastigiated and
carinated, with the carinae partly covering each other,
form a straight row from the temporal region above
the gill-cover out along the lateral edges of the com-
pressed tail, though here they grow thin and indistinct.

The first free spinous ray on the back is set above
the hind part of the gill-cover, at a distance from the
tip of the snout measuring between 241/ 2 and 22 %
(generally decreasing with age) of the length of the
body. The last (hindmost) of these spinous rays lies
exactly opposite the vent, and the distance between it
and the tip of the snout varies individually between
45 and 46 % (exceptionally 43 %) of the length of the
body. Thus, during youth, the space occupied by these
spinous rays is, as a rule, somewhat less, in old spe-

cimens somewhat greater, than the distance between the
first ray and the tip of the snout. The length of the
rays is somewhat greater than the distance between
them; and their articulations lie a little to one side
and in a zigzag, so that, when they are depressed, the
tip of each ray falls beside the base of the next ray
behind it, in the groove formed by the above-mentioned
concave plates". Just behind the last spinous ray rises
the true dorsal fin, triangular, with rounded apex, the
first ray being somewhat shorter than the second, but
equal in length to the third, the other rays decreasing
in length gradually, but sharply. Sometimes both the
first and the second rays are simple, but in most cases
all the rays are more or less deeply branched. The
last ray is united throughout its length by the fin-
membrane to the dorsal edge. When erected, the rays
are recurved, the anterior (as well as the posterior)
side of the triangle being thus convex. The length of
the base of the fin is about 1/2 (varying between 56
and 47 %) of that of the head. Just in front of the
anal fin we find a free spinous ray6, equal in size
to the last spinous ray on the dorsal edge, but situated
further back, vertically below the first or even the
second ray of the true dorsal fin. Thus, the anal fin
being almost exactly similar to the dorsal in form,
structure, and size, the termination of the former lies
somewhat further back than that of the latter. The
form of the caudal fin reminds us, to a certain extent,
of a combination of the anal fin with the true dorsal,
in which combination each of these fins would cor-
respond to a half of the caudal fin, which is fan-shaped
when expanded. In this species, as in all the Scandi-
navian Sticklebacks, this fin contains 12 rays, excluding
a few (generally 2 or 3) small supporting rays at each
margin), among which the 10 middle ones are branched
for about half their length, but the outermost simple.
The middle rays of the caudal fin are always some-
what, though sometimes only slightly, shorter than the
longest rays of the dorsal fin, and measure about 8C
or 9 % of the length of the body.

The pectoral fins are of the form and structure
common to all the Sticklebacks. As a rule they consist

" When the number of these rays is less than usual, it is generally the 14th that is wanting, as KR0YER has remarked, the distance
between the last ray and the last but one being thus greater than that between the other pairs. The interspinal plate of the absent ray is
also wanting. In such specimens we have also found the third spinous ray considerably reduced, and its interspinal plate proportionately
smaller than the others. In some cases Heincke found only 13 free spinous rays.
h In our figure this ray has unfortunately been overlooked.
c Sometimes 7.

FIFTEEN -SP I N ED STICKLEBACK .

641

of' 10° simple rays of fairly uniform length, though
the middle ones are somewhat longer than the outer,
the tip of the fins, when they are folded, being thus
evenly, but sharply rounded. When expanded these
fins are fan-shaped. Their base is almost perpendicular,
and by the great expansion of the shoulder-blade and
the coracoid bone is removed farther than usual from
the clavicle, leaving an even patch, covered with glossy
skin, in front of it. The length of the pectoral fins is
generally about equal to the base of the soft-rayed
dorsal or the anal fin, and is usually somewhat greater
in the males than in the females, varying between
about- 10 and 12 % of the length of the body.

The ventral fins lie a little behind the middle of
the pelvic bones and a little behind the end of the first
third* 6 of the body. They are made up of one spinous
ray, which is distinctly longer than (sometimes nearly
twice as long as) any of the other spinous rays, and
behind this ray they have a- triangular fin-membrane,
set obliquely inwards and backwards, and containing
in its posterior (inner) part two small, soft rays, the
last of which is in most cases difficult- to detect. Like
all the other spinous rays, that of the ventral fins is
also granulated throughout the greater part of its length,
with smooth tip; but in this species all the spinous
rays are without lateral spines and denticles.

The only external difference that we have been
able to discover between the sexes of this species, is
that pointed out by Lilljeborg, namely that the pec-
toral fins are generally longer in the males than in the
females; and even this is faint. This uniformity is
highly remarkable, for in our other Sticklebacks the
difference between the sexes is striking enough, even
in the external form.

The peritoneum is white with a silvery lustre, but-
dotted with black, which is often the predominant co-
lour on the dorsal side of the abdominal cavity. The
intestinal canal is simple and short. Just behind the
diaphragm the oesophagus, which is lined with longi-
tudinal folds, passes into the stomach, the inside of
which is reticulate, and which extends scarcely half-way
along the abdominal cavity, to about a line with the
hind extremity of the interclavicles, where it- is bounded
from the intestine by a strong and muscular contrac-
tion. The inside of the intestine is thickly lined with

villi, arranged in oblique, transverse series. In a
straight- direction, and with a breadth almost equal to
that of the stomach, but growing narrower behind, the
intestine extends almost to the end of the abdominal
cavity, where it bends in a sharp curve obliquely up-
wards and forwards, soon t-o return in a curve no less
sharp backwards and downwards to the vent. The only
trace of pyloric appendages that we have been able to
discover, is an extremely short process, directed back
towards the stomach, on the intestine just behind the
pylorus.

The liver lies in the form of an undivided mass
below and along the sides of the oesophagus and sto-
mach, with a blunt point in a downward and forward
direction towards the union of the interclavicles. On
the left side it has a short lateral lobe, which does not
extend further back than the main lobe; but the lobe
on the right side is longer, and follows the intestine
for about half of its anterior, straight- part, or a little
further. At the end of the middle lobe, under the ter-
mination of the stomach, a little to the right, lies the
gall-bladder, and on the left side, above the end of the
stomach, the spleen. The air-bladder is situated in the
posterior part of the abdominal cavity, and its anterior
extremity lies above the end of the stomach.

The ovaries and testes are paired and saccate. The
former at least, with their comparatively large and few
eggs — Kr0yer estimated their maximum number at
300 — occupy the greater part of the abdominal cavity
during the spawning-season.

In coloration the Fifteen-spined Stickleback is one
of our most beautiful fishes, though under the influence
of mental agitation, of terror for example, it- often
suddenly loses its beauty and brightness of colour, and
does not regain them for a long time. The upper part
of the body is olive green, shading into brown, the
lower yellowish with white, silvery belly. The sides
are, as it were, transparent, showing glimpses of the
spinous processes of the vertebrae in the form of dark,
transverse, parallel streaks. Below the lateral line, on
a ground of lustrous silver, lies a row of large, crescent-
shaped and oval, olive green spots, which grow smaller
behind and finally disappear on the tail. A dark brown
band runs along the side of the head from the cor-
ner of the mouth to the base of the pectoral fin.

a Sometimes 9; exceptionally 11.

6 The distance between the ventral spine and the tip of the snout varies in different individuals between about 34 and 37 % of the
length of the body.

642

SCANDINAVIAN FISHES.

The pectoral tins are yellow with a brassy lustre. The
true dorsal and the anal tins are grayish brown with
yellow rays in front; behind the membrane is trans-
parent. The caudal fin is olive-green, with transparent
outer margin and yellowish brown rays. The iris is
gray, above darker with black margin, below whitish
yellow; the pupil is surrounded by a narrow, orange
ring.

The geographical range of the Fifteen-spined Stick-
leback is known to extend from North Cape along the
west coast of Europe to the Bay of Biscay. In Iceland
and America it is unknown. The most westerly parts
of its range are the Faroe Islands and Ireland. It pene-
trates into the Baltic as far as the south coast of Fin-
land. Off Hogland, in the Gulf of Finland, it is com-
mon, according to Mela; in the island-belt, of Stockholm
it is rare. Being distinctly a marine fish, never enter-
ing fresh water, it is commoner on the west coast of
Sweden than on the east. It is also common on the
coast of Norway, off the Orkney and Shetland Islands,
and southward at least to the coast of Brittany; but
in the Bay of Biscay it is rare.

The Fifteen-spined Stickleback is a shore-fish and
lives among the seaweed, ascending into half-a-fathom
of water and even up to high-water mark. “Its favourite
haunts,” says Malm, “are such spots as possess a sandy
bottom, studded with large or small stones covered
with Fucacece and interspersed with groves of Zostera.
Among these it gracefully threads its way by short
stages, now hither and now thither.” It is not very
active nor very timid; but when frightened it darts
forward with the speed of an arrow. In contradistinc-
tion to the other Sticklebacks it leads a more solitary
life, or at least does not assemble in so dense or large
shoals. It is tenacious of life" and a greedy eater. Its
food consists principally of small crustaceans (especially
of the genera My sis and Idothea ) and worms; but it is
also accused of preying upon the roe and fry of other
fishes. Couch* 6 once satv a Fifteen-spined Stickleback
seize and partly devour a young Eel 3 inches long,
though it was at last compelled to disgorge its victim.

During the spawning-season, which occurs in
spring and summer, the Fifteen-spined Stickleback

makes its way to shallow water, within a harbour or
in some little inlet sheltered from the waves, often
between the tide-marks. Here it builds a nest for its
eggs and young. This is constructed either on the
bottom or, perhaps more frequently, floating in the
water, under a pendent tuft of seaweed or some
other object — Couch found one of these dwellings be-
tween the loose twists of the end of a rope °. When
the nest hangs free, it is usually pear-shaped and of
the size of one’s fist. Buckland^ gives the following
description of the nursery of the Fifteen-spined Stickle-
back: “The main body of the nest is formed of very
soft weed — in fact, as soft as sponge — and, strange
to say, a.s though for the sake of ornament, our little
architect has placed at the point where the nest is
thickest a bit of brilliant blood-red weed. Nay, more,
it appears as though the builder of the nest, fearing
the risk of discovery, had worked in great broad por-
tions of brown ribbon Aveed, Avhich should act, firstly,
as a covering, or band, to keep the whole structure
together; and, secondly, to serve the purposes of con-
cealment. So beautifully, indeed, is this nest con-
structed for the concealment of the eggs, that unless
the naturalist to whom the nest Avas sent had been
previously aAvare Avhat the structure meant, he might
easily have been pardoned if he had not seen the eggs
at all. After a careful dissection of the outer coatings
of the nest, the eggs themselves came into vieAv. These
eggs are round little bodies about the size and colour
of mustard seed. They are in bunches like grapes, and
we try Avhether they are to be separated from the bunch
Avith ease. Another marvel; the body of eggs are all
sewed into a compact mass by a very thin, delicate
fibre, Avhich in the sun glistens like a cobAveb or the
very finest floss silk. The filaments Avhich hold the
eggs are Avorked through, over and round them, so
as to form a complete netAvork. ’ Threads of the
same sort also serve to hold the Avails of the nest
together, and are secreted by the fish in the form of
a glutinous substance that hardens in the Avater.

Malm has described (1. c.) his observations of the
spaAvning of the Fifteen-spined Stickleback on the 3rd
of August, 1854, off Kristineberg (Gullmar Fjord).

a KR0YER, however, found it less tenacious of life than the true Sticklebacks.

6 Hist. Fish. Brit. Isl., vol. I, p. 183.
c 1. c., pi. XXXVIII.

d Nat. Hist. Brit. Fish., p. 250. For further information on this head see Hancock, Ann. Mag. Nat. Hist., 2nd ser., vol. X
(1852), p. 246.

FIFTEEN-SPINED STICKLEBACK.

643

“It was a female", greenish in colour, that was busy
near the landing-stage, in about 7 or 8 dm. of water,
building a dwelling, or a kind of nest, for the eggs
soon to be deposited. In a round IioIIoav, about 2 7 2
cm. deep and 15 cm. in diameter, in the sandy bottom,
she placed, several times an hour, small fragments of
dead plants, which she found during her wanderings
in the neighbourhood and carried in her mouth to the
spot. Now and then she worked herself into the heap
thus collected, turning quickly round and round, and
emerged on the other side of the pile. In this manner
she carded the building-materials with the stiff and
sharp rays of her fins, and at the same time oiled them
with the slime secreted by her own body, thus forming
an elastic tube, in which she finally deposited her eggs.
The male in his more bluish dress kept guard, swim-
ming round and round the spot, and iioav and then
rewarded by a friendly greeting from the female, Avhile
the enemies of the roe, the shrimps and shore-crabs
etc., Avere repulsed by the powerful onslaughts of the
brave sentinel, as soon as they ventured to approach
his post. The female was often compelled to come to
the rescue,' but the fish Avere always victorious. But
if another male of the same species came up, he Avas
at once joined by his host in some friendly sport, Avhich
ended in a regular game of tig among the stones and
seaweed several yards from the spot. But as soon as
the host saAv that he had led astray his uninvited guest,
Avhom he had evidently enticed from the spot by this
stratagem, he returned at full speed, to resume at the
next moment the defence of his home and family. They
paid no attention to fishes of other species, such as
Gobies etc.” From this observation Malm concludes
that the Fifteen-spined Stickleback lives in pairs, at
least during the spaAvning-season, and is not polyga-

mous like the other Sticklebacks; while it would also
seem to be peculiar to this species that the female
should build the nest and take part in its defence.
Other observers, hoAvever, are of opinion that here too
the male is the actual builder, but that the nest is
guarded both by the male and the female. The parents
continue their vigilant care of the eggs and the new-
hatched young for at least three or four Aveeks. Ene-
mies are repulsed; if the nest is damaged, they repair
it; if they are driven aAvay, they soon return. If the
nest be built between the tide-marks, they must, of
course, leave it to its fate during the ebb; but the
moisture retained by the Avails of the nest suffices to
keep the eggs or young alive until the tide rises again.

Of the growth of the fry Ave learn from the col-
lections of the Royal Museum that during the expedi-
tion of the gunboat Gunhild, on the 25th of July, 1878,
a young specimen 3l72 mm. long Avas taken at a depth
of four fathoms on a stony and Aveedy bottom, in the
south of the Sound between Malmo and Falsterbo. It
belonged, no doubt, to the fry of the same year. On
the Irish coast Thompson6 found young specimens 19
mm. long in June and others, 25 mm. long, in July.
Through Professor Lindstrom the Royal Museum has
acquired some young specimens between 57 and 71
mm. in length and taken on the coast of Gothland in
August, 1852. These last specimens, hoAvever, can
hardly have been less than a year old.

The solitary life of the Fifteen-spined Stickleback,
which prevents it from ever being caught in any quan-
tity, deprives it of all special importance to man in an
economical respect. It is taken only by accident among
other fishes in small-meshed nets or the seine; and it is
utilised only in the manufacture of oil and as manure.

(Ekstrom, Smitt.)

“ As the nest was afterwards destroyed by accident and the fish were frightened away, Malm was prevented from ascertaining by
closer examination whether this specimen was really a female, and her companion a male.
b Nat. Hist. Irel ., vol. IV, p. 90.

644

SCANDINAVIAN FISHES.

, Genus GASTEROSTEUS a.

Body mackerel-like , compressed ( greatest depth more than half the length of the head), without other carince than
the lateral edges of the kelly and the elevation of the lateral line on the hind part of the tail; forepart of the
body {in front of the anal spine) longer than the hind part, including the caudal fin. Number of free spinous
rays in front of the soft-rayed dorsal fin , less than 13. Ventral fins inserted at the outer , front corner of the
pelvic hones , which coalesce with each other in the median line of the kelly into a triangular ventral plate ,

tapering behind. Jaw-teeth conical.

The genus of the Sticklebacks contains the most
widely-spread of all the Scandinavian fishes. There is
hardly a single brook in which we fail to see these
small and lively creatures swimming merrily about,
often within narrow limits which the heat of summer
reduces more and more each minute. For economical
purposes they generally possess far too little importance
to interest the practical fisherman, who usually knows
them merely by name. The zoologist cannot content
himself with so slight an acquaintance. To him a
knowledge of the Stickleback is quite as important as
of its far larger and far more useful fellow, the Salmon.
Both are equally small fractions in the great series
of natural forms which it is his task to investigate,
and both are equally interesting, equally advantageous
subjects of study. When we add to this the wonder-
ful instinct of the Sticklebacks, which guides them to
an architectural skill exceptional among fishes, we have
sufficient grounds for bestowing on them more than
usual attention.

To the system atist. they are also of absorbing in-
terest, for the course of development may be traced
with especial distinctness in their differences of form
and changes of growth. Their relation to the preceding
genus has been expressed in a masterly way by Dr.
Heincke6, in an elucidation of the significance of the
variations affecting the dorsal spines in the Three-
spined Stickleback. With this object he had examined
about 10,000 specimens of the species in question.

The Three-spined Stickleback is generally furnished
— and this is one of its most important characters —
with 3 free spinous rays at the dorsal edge in front
of the soft-rayed dorsal fin. It had long been known,
however, that the number of these spinous rays might

be increased to 4C; and a variety of this kind, with
the four spinous rays comparatively short, had been
described by Cuvier and Valenciennes'* as a distinct
Italian species, Gasterosteus tetracanthus. Heincke found
about 1 percent of the Three-spined Sticklebacks that
he examined, furnished with four spines; and of this
number he examined 61 specimens in order to locate
the supernumerary ray. Each of the spinous rays is
attached to and articulated, in a singular manner which
we shall describe more fully below, with one of the
interspinal plates on the back. These plates are as a
rule (fig. 157 and fig. 158, B) 6 in number and arranged

3

Fig. 158. Gasterosteus aculeatus, gymnurus. Natural size. From
Italy. After Heincke.

in a row: 2 small ones just behind the head, 2 larger
ones, each with a spinous ray, the poster.^, usually
the larger, 1 smaller plate with no spinous ray, and
lastly 1 supporting the hindmost spinous ray. The
largest two of these plates, however, are evidently due
to the coalescence of at least 2 plates respectively. In
fig. 159 we see the anterior one broken up into two
parts, and in fig. 1 60 the posterior of them shows the
same division. The anterior also rests on the upper

° The generic name is derived from Artedi, Genera Piscium , p. 52.
1 Ofvers. Vet. Akad. Fork. 1889, p. 395.
c Cuv..' Val., Hist. Nat. Poiss., vol. IV, p. 491.
d 1. c., p. 499.

STICKLEBACKS.

645

spinous processes of two vertebras, the posterior on
those of four. Thus we have as a rule to deal with
8 interspinal plates at the dorsal edge in front of the
soft-rayed dorsal tin. In one single case, however,
Heincke could trace 9 of these interspinal plates, the
last two each with a distinct, well-developed spinous
ray, and the third, counting from behind, with a ru-

Fig. 159. Gcisterosteus aculeatus, trachurus. Natural size. From
the Frisches Halt, off Kouigsberg. — - ® the posterior, detached part
of the third dorsal plate (ds. 3 ). After Heincke.

dimentary spinous ray “in the form of a rigid spine.”
The Three-spined Stickleback has thus shown at least
traces of as many as 5 free spinous rays at the dor-
sal edge.

If we now observe the arrangement of these 5
spinous rays, we shall find that each one of the 9

Fig. 160. ' Gasterosteus aculeatus , trachurus , tetr acanthus, with the
supernumerary spinous ray set on the penultimate dorsal plate. Natural
size. From Kiel Bay. — 0 the posterior, detached part of the fourth
dorsal plate (ds. 4). After Heincke.

dorsal plates, with the exception of the first, may be
furnished with a spinous ray, though not simultaneously.
In order to express the differences that occur in this
respect, we shall denote each of the interspinal plates
by an Arabic figure (1 — 9), the plates which in each
case must be regarded as double, by brackets ( — — ),
the three ordinary spinous rays by the Roman num-

erals I — III, the most usual supernumerary ray by IV,
and the rudimentary and supernumerary ray that has
only once been observed, by V. The cases (varieties)
observed by Heincke are then as follows:

A: in Three-spined Sticklebacks with 3 free spinous rays at
the dorsal edge.

a: with the regular number (6) of interspinal plates, the
forms most typical in this respect (figs. 157 and 158).

0 0 1 _n_ 0 0 III

1 — 2 — 3 — 4 — 5 — 6— 0 — 8 — 9

b: with 7 interspinal plates (the 3rd and 4th detached
from each other, fig. 159).

0 0 1 0 II 0 0 III

1 — 2 — 3 — 4 — 5 — 6 — 0 — 8 — 9

Fig. 161. Gasterosteus aculeatus, trachurus, tetracanthus, with the
supernumerary spinous ray set on the penultimate dorsal plate.
Natural size. From Eckernforde Bay. After Heincke.

B: in Three-spined Sticklebacks with 4 free spinous rays at
the dorsal edge.

c: with a supernumerary spinous ray on the penultimate
interspinal plate, otherwise as in variety a. 36 speci-
mens out of 61 showed this arrangement (fig. 161).

0 0 1 II 0 IV III '

1 — 2 — 3 — 4 — 5 — 6 — 0 — 8 — 9

d: similar to c, but with the 5th and 6th plates detached
from each other (fig. 160). 16 specimens out of 61.

0 0 I II 0 0 IV III

1 — 2 — 3 — 4 — 5 — 6 — 0 — 8 — 9

e: with a supernumerary spinous ray on the 6th plate and
with this plate detached from the 5th. 4 specimens
out of 61.

0 0 1 II IV 0 0 III

1 — 2 — 3^4 — 5 — 6 — 0 — 8 — 9

f: with only 5 interspinal plates, otherwise similar to c.
2 specimens out of 61.

0 0 I II 0 IV III

0 — 2 — 3^4 — 5^~6 — 0 - 8 — 9

g: with only 6 interspinal plates, otherwise as in d. 1
specimen out of 61.

0 0 1 II 0 0 IV 111

0 — 2 — 3 — 4 — 5 — 6 — 0-8 — 9
h: with a supernumerary spinous ray on the 4th inter-
spinal plate, which is detached from the 3rd, the latter,

646

SCANDINAVIAN FISHES.

on the other hand, being confluent with the 2nd (fig.
162). 1 specimen in 61.

0 I IV II 0 0 111

1 — 2 — 3 — 4 — 5 — 6 — 0 — 8 — 9

i: with a supernumerary spinous ray on the 2nd inter-
spinal plate, otherwise similar to a. 1 specimen in 61.

0 IV I II 0 0 III

1 — 2 — 3 — 4 — 5 — 6 — 0 — 8 — 9

C: in one Three-spined Stickleback with a rudiment of a fifth
spinous ray and with eight interspinal plates.

0 0 I II 0 V IV III

1 — 2 — 3 — 4 — 5 — 6 — 7 — 8 — 9

Now, as at least 7 of the interspinal plates —
though only 3 — 5 at once — may be furnished in the
Three-spined Stickleback with true spinous rays, and
as 7 — 12 free spinous rays occur at the dorsal margin

Fig. 162. Gasterosteus aculeatus , trachurus , tetracanthus ( spmulosus )
with the supernumerary spinous ray (*) set on the posterior, de-
tached part of the third dorsal plate. Natural size. From the Frisches
Haff, off Konigsberg. After Heincke.

of the Ten-spined Stickleback, while in the Fifteen-
spined Stickleback the limits of the variation in this
respect are 13 and 16, the series, with decreasing num-
bers, is complete from the last species to that first
mentioned. Heincke assumes that the development has
taken this direction, with constantly sinking numbers.
Thus, where more than 3 free spinous rays occur at
the dorsal edge of the Three-spined Stickleback, we
have a retrogression to an older stage of development,
through which the species has already passed. The
Fifteen-spined Stickleback, according to this theory, re-
presents the earliest stage of the development of the
whole family, whose descent should thus be traced
from fishes of the Fistularioid family.

The alterations of development, however, lead us
to a different conclusion, namely that the Ten-spined
Stickleback stands nearest the probable origin of the

family. We have already seen that the elongated form
which is the chief characteristic of the genus Gastrcea

— though we meet with it in the youngest forms of
all the Sticklebacks — especially in the character that
suggests a transition to the Flute-mouths, namely the
elongation of the snout, is a character of age, a cha-
racter which develops during growth from a juvenile
stage with the typical form of the species perfect in
other respects, but with the length of the snout only
slightly greater than in the true Sticklebacks. Even
during the later (older) stages of growth this general
rule holds good, for the length of the snout shows
increase, e. g. in proportion to the postorbital length
of the head (see the following table), both in the Fifteen-
spined Stickleback and the true Sticklebacks, but is least
in the Ten-spined species. Another relation, in which
the changes of development also follow the same direc-
tion in all the three species now under consideration,
gives the same result. If we endeavour to formulate
an expression for the position of the soft-rayed dorsal
and the anal fins with respect to each other, we find
that, as a general rule, the anal spine lies somewhat behind
the perpendicular from the beginning of the former fin

— in the Ten-spined Stickleback alone it may excep-
tionally lie somewhat in front of this line — and that
the percentages for the distance between the former fin
and the tip of the snout compared with that between the
anal spine and the same point, run in all three species
in inverse proportion from the earlier stages of growth
to the later. The highest percentages, which are thus
expressions of the lowest stages of development, occur
in the Ten-spined Stickleback (see the following table).
But this is not all. In the relations in which the Fifteen-
spined Stickleback shows a deviating direction of deve-
lopment,- it stands nearest to the Ten-spined Stickleback.
For example, if we compare the length of the soft-rayed
dorsal fin with that of the head, we find that this fin
is considerably longer in the true Sticklebacks than in
Gastrcea , and longest in the Three-spined Stickleback.
The percentage for this relation also increases, as a ge-
neral rule, during the growth of the Sticklebacks, but
decreases with age in Gastrcea (see the following table).

Average in:

Gastrcea

spinacliia.

• Gasterosteus
pungiiius.

Gasterosteus

aculeatus.

Length of the body expressed in millimetres..

64.9

132.2

41.2

56.3

55.8

69.7

„ ,, „ snout in % of the postorbital length of the head

96,i

96.3

51.2

56. s

73.6

74.8

Distance between the soft-rayed dorsal fin and the tip of the snout in % of the distance between the anal
spine and the same point...

97.2

95.4

99.9

97.8

90.7

86.i

Length of the base of the soft-rayed dorsal fin in % of the length of the head

52.6

49.i

84.i

88.i

86.2

95.7

STICKLEBACKS.

647

Thus both the Fifteen-spined and the Three-spined
Sticklebacks stand nearer to the Ten-spined Stickleback
during youth than they do in later life. During growth
they diverge from this species, each in a special direc-
tion. The first direction of development tends towards
the Flute-mouths, the second, as we have mentioned
above, towards the Trumpet-fishes.

This conclusion differs essentially, it is true, from
that arrived at by Heincke, but agrees with the latter in
the most important point, namely that the Three-spined
Stickleback is descended from a species that was extre-
mely closely allied to the Ten-spined Stickleback, unless
indeed its ancestor be this very species. As evidence
in favour of this Heincke cites Gasterosteus Bussei , the
form described from Amur in 1887 by Herzen stein and
Warpachowski, with 9 free spinous rays at the dorsal
margin (like the Ten-spined Stickleback), but with a
complete row of scutes along the sides of the body (like

the Three-spined Stickleback). A similar form had al-
ready been described in 1869, from David’s Chinese
collections, by Guichenot" under the name of Gastero-
steus sinensis , and in 1880, from Dybowski’s collections
in the bays of Northern Japan, by Steindachner/' un-
der the name of Gasterosteus japonicus. We have an-
other intermediate form in Gasterosteus ( Eucalia ) in-
constans from North America arid Greenland, with the
branchiostegal membranes inferiorly free, at least to
some extent, from the isthmus (as in the Ten-spined
Stickleback) and without lateral plates on the body, but
with only five free spinous rays at the dorsal edge”.

The range of the genus Gasterosteus embraces both
fresh and salt water in the Arctic and Temperate Zones
of the Northern Hemisphere, both in the Old World and
the New; but the number of the species is probably no
more than five, two of which belong to the Scandi-
navian fauna.

THE THREE-SPINED STICKLEBACK (sw. storspiggen).
GASTEROSTEUS ACULEATUS.

Plato XXVIII, figs. 1 (9) and 2 ((f).

Three (exceptionally four) free spinous rays at the dorsal margin. Branchiostegal membranes inferiorly united

to the isthmus throughout its length.

R. br. 3; D. Ill— IV|l 0—12 A. I|8— 10; P. 1 0 e — 1 1 ;
V. 1 1 1 ; C. x+ 1 + 10 + 1 +(c; L. lat. 3—34 ;Vert. 31—32 A

Syn. Gasterosteus aculeis in dorso tribus, Art., Gen. Pise., p. 52;

Syn. Pise., p. 80; Spec. Pise., p. 96; Lin., Fn. Suec., ed.
I, p. 103; Westbeck, Vet.-Akad. Handl. 1753, p. 261;
Str6m, Sondm. Pester., vol. I, p. 315; Odmann, Vet.-Akad.
Handl. 1782, p. 167.

Gasterosteus aculeatus, Lin., Syst. Fat., ed. X, tom. I, p. 295;
Fabr., Fn. Groenl., p. 169; Retz., Fn., Suec. Lin., p. 338;
Pall. (G aster acanthus) Zooyr. Ross. Asiat ., tom. Ill, p. 229;
Cuv. ( Gasterosteus ), Reyn. Anim., ed. I, tom. II, p. 320;
Ekstr., Vet.-Akad. Handl. 1831, p. 296; Nilss., Prodr.
Ichth. Scand., p. 85; Jenyns, Man. Brit. Vert. Anim., p.
348 ( + G. spinulosus, p. 350); Ekstr., v. We., Skand. Fisk.,
ed. I, p. 17, tab. 4, fig. 1, a et b ; Kr., Damn. Fiske , vol.
1, pp. 169 et 590; Lillj., Vet.-Akad. Handl. 1850, p. 309;
Hancock, Ann. Mag. Nat. Hist., ser. 2, vol. X, p. 241;

Sund., Stoekh. L. Hush. Sallsk. Hand]., H. 6 (1855) pp. 79,
85, 176; Nilss., Skand. Fn., Fisk., p. 103; Thomps., Nat.
Hist. Irel., vol. IV, p. 82; Heck., Kn., Siisswassei1/. Osterr.
Mon., p. 38 ( + G. bracliycentrus, p. 41); Sieb, Susswasserf.
Mitteleur., p. 66; Mgrn, Finl. Fisk.-Fn. (Disp. Helsingf.)
p. 14; Lindstr., Gotl. L. Hush. Sallsk. Arsber. 1866, p. 14
(sep.); Canestrini, Fn. D'ltal., pt. Ill, Pesci, p. 25; Coll.,
Forb. Vid. Selsk. Christ. 1874, Tillaegsh., p. 11; ibid. 1879,
p. 2; N. Mag. Naturv. Christ., Bd. 29 (1884) p. 48; Malm,
Gigs, Boh. Fn., p. 371; Seidl., Fn. Balt., p. 128; Winth.,
Naturh. Tidskr. Kblivn, ser. 3, vol. XII. p. 4; Fedders.,
ibid., p. 73; Bncke, Fisch., Fischer., Fiscliz. O., W. Preuss.,
p. 73; Id., Handb. Fischer., Fiscliz. (M. v. d. Borne), p.
98; Mor., Hist. Nat. Poiss. Fr., tom. Ill, p. 163; Mela,
Vert. Fenn ., p. 277, tab. IX; Day, Fish. Gt. Brit., Irel.,
vol. I, p. 238, tab. LX VIII, figg. 1—3; Ah. Aqass., Proc.
Amer. Acad. Arts, Scienc., vol. XVII (1882), p. 288, tab.
IX; Hansen, Zool. Dan., Fiske, p. 27, tab. V, figg. 1 et 2;

“ Nouv. Arch. Mus. cFHist. Nat. Paris, tome V, p. 204, pi. 12, fig. 4.
h Stzber. Akad. Wiss. Wien. Math. Naturw. Cl., Bd. LXXXII, i, p. 264, taf. Ill, fig. 2.
r Cf. Jordan and Gilbert, Bull. U. S. Nat. Mus., No. 16, p. 394.
d Sometimes 8, according to Kroyer.

,, 14, ,, ,, Lilljeborg.

e „ 9, ,, ,, Kroyer.

f Exceptionally 30, sometimes 33, according to Fatio.

Scandinavian Fishes.

82

648

SCANDINAVIAN FISHES.

Storm, N. Viol. Selsk. Skr., Trondhj. 1883, p. 15; Mob.,
Hoke, Fiscli. Osts., p. 66; Lillj., 6'v., Norg. Fisk., vol. I,
p. 343; Mob., Nature, vol. XXXIX, No. 9I>8 (Dec. 13, 1888)
Hcke, dfvers. Vet.-Akad. Forli. 1889, p. 395.

Gcisterosteus biaculeatus, Shaw (ex. Penn.) Gen. Zool., vol.
IV, p. 608.

Gasterosteus trachurus + G. gymnurus ( leiurus ) + G. argyropo-
mus + G brachycentrus + G. tetracanthus + G. noveboracensis
+ G. niger , Cuv., Regn. Anim., ed. 2, tom. II, p. 170; 4- G.
semiarmatus + G. semiloricatus , Cov., Vat.., Hist. Nat. Poiss.,
vol. IV, pp. 493 et 494; G. loricatus + G. dimidiatus, Reinh.,
D. Vid. Selsk. Math. Naturv. Afh., VII (1838), pp. 114 et
119; G. aculeatus + neustrianus 4- semiloricatus 4- semiarmatus
4- leiurus + Bailloni 4- argentatissimus + elegans , Blanch., Poiss.

■ d. eaux douces Fr., pp. 213 — 236; G. aculeatus 4- ponticus
+ noveboracensis + suppositus + obolarius (?) 4- loricatus + niger
+ serratus 4- neustrianus 4- semiarmatus + semiloricatus + texa-
nus 4- leiurus + Bailloni 4- algeriensis 4- plebeius 4- inopinatus 4-
argentatissimus 4- elegans 4- islandicus + biaculeatus + microce-
phalies + argyropomus 4- brachycentrus 4- tetracanthus 4- spinu-
losus , Sauv., Nouv. Arch. Mus., tom. X, pp. 9 — 25; G.
microcephalus 4- biaculeatus 4- atkinsii (ex Bean) 4- aculeatus ,
Jord., Gilb., Bull. U. S. Nat. Mas., No. 16, p. 395.

Obs. The great number of nominal species that have been estab-
lished, according to the above list of synonyms, in the endeavour
to elucidate the inconstancy of form among Three-spined Stickle-
backs, is merely a significant token of the lengths to which this in-
constancy may run in a species that sometimes occurs in enormous mul-
titudes, is found in far distant localities both in salt water and in fresh,
and both in form and colour shows marked sexual distinctions.

The Thi •ee-spined Stickleback attains a length of
at least about 9 cm. Our largest specimen is a female
from Greenland which measures rather more than 91
mm. from the tip of the snout to the end of the middle
rays of the caudal tin, or 93 mm. from the tip of the
lower jaw to the end of the outer caudal rays". The
form of the body reminds us strongly of the Horse-
Mackerel, being fusiform with strong lateral compression.
The body is deepest across the pelvis at the articulation
of the ventral spines, where the depth is about 1/5 of
the length of the body, varying individually between
18V2 and 2272 % thereof. The thickness at the same
spot is about half the depth, varying individually be-
tween 44 and 57 % thereof. Except during the spawn-
ing-season, when the organs of generation become tumid
and render the body more terete, the thickness is almost
uniform from the eyes back to the strongly compressed
base of the caudal fin, the sides being flat throughout
the forepart of the body, but with sharp Carinas on

their posterior parts, except in the variety gymnurus,
which is without caudal plates, and in which the sides
of the tail are also flat. The dorsal profile between the
head and the soft-rayed dorsal fin and the ventral pro-
file between the head and the anal fin are usually com-
paratively straight or only slightly convex, while the
upper and lower contours of the head in front and of
the tail behind are more sharply convergent. Some-
times, however, especially in the females, which gene-
rally show a deeper form of body, the entire upper and
lower profiles of the body run in unbroken curves. The
external form is also affected to a considerable extent
by the more or less advanced development of the co-
vering of plates, this being the origin of most of the
numerous different names by which the species has
been known.

The whole covering of plate-armour shows fairly
great individual variations both in extent and in strength.
In its strongest form it consists of comparatively thick
osseous growths in the skin, externally resembling Ga-
noid scales, granulated and striated by grooves and
ridges, the latter rough with small granules. But, as
Heincke has remarked, both the thickness and the rough-
ness may be reduced, to the greatest extent, as a rule,
in the Three-spined Sticklebacks that inhabit brackish
or fresh b water, or belong to more southern regions.
Still forms with extremely reduced plate-armour also
occur in the Arctic regions, as for instance in Green-
land, where the Sofia Expedition of 1883 took a number
of small specimens of the gymnurus form in a lake near
Ritenbenk (North Greenland). These specimens, like the
Italian argyropomus , have only three plates on each side
of the body, namely the three that adjoin the ascending
lateral disk of the pelvic bones.

The plate-armour of the back we have already exa-
mined for the most part, and we need only add that
here, as in the Fifteen-spined Stickleback, the soft-rayed
dorsal fin rests on a row of interneural plates (here
about 13, i. e. — excluding the plate of the last spinous
ray — one for each ray and one more behind the fin).
This statement also applies to the anal fin, which gene-
rally has 11 interhsemal plates at the base, the first
supporting the spinous ray and the last two usually

a Thus almost exactly equal to the largest specimen in Copenhagen Museum, according to Hansen (Zool. Dan., 1. c.). According to
Pallas it attains a greater size. His G aster acanthus cataphractus from Kamchatka seems to be simply the typical Three-spined Stickleback,
but is stated to attain a length of 122 mm. (longitudine pollicum sesquiquinque). Cf. also Tilesius, Mem. Acad. Petersb., tom. Ill (1809

— 10), p. 226.

b Already remarked by Kr0Yer (1. c., p. 180).

TH 1IEE-SP1NED ST ICIvLEBACK .

without rays. The ventral margin in front of the anal
tin forms a more or less distinct break for the vent;
and in front of the vent, at a. greater or less distance
therefrom, lie the pelvic bones so characteristic of the
Sticklebacks, or rather their covering bones. Here we
see the so-called pelvic bones ( propelvis ) distinctly si-
tuated as osseous growths in the skin, in an homologous
series with the interhsemal plates mentioned above;
while in the other Teleosts the pelvic bones, in their
position and often in their form as well, have proved
quite as distinctly homologous with the interhgemal
bones (spines) themselves. This difference is accompa-
nied by another: in the Sticklebacks the ventral fins
are attached to the middle (in Gastrcea) or to the an-
terior part (in Gasterosteus “) of the so-called pelvic
bones, while in the other Teleosts they articulate with
the hind extremity of these bones.

In the Three-spined Stickleback, as in the following
species, the pelvic bones, with their inner margins unit-
ed by a suture in the median line of the belly, form
a compact, triangular plate, pointed behind, which at
the anterior part of each of its sides sends out a foliate
disk up the sides of the belly. In the Three-spined
Stickleback this disk generally meets three of the lateral
plates of the body on each side, thus forming a com-
plete pelvic girdle, belonging to the dermoskeleton. The
anterior margin of the pelvic plate is straight or only
slightly concave* * * * 6. The two anterior corners touch the
interclavicles already mentioned, which in the Three-
spined Stickleback are comparatively broad — their
breadth at the middle about equal to the distance be-
tween them0 — and so long that the pelvic bones lie
entirely behind the insertions of the pectoral fins.

In the typical Three-spined Stickleback the sides
of the body are almost entirely covered by a corslet
formed by a row of plates belonging to the lateral line,
the only naked parts being the belly itself, up to a level
with the bottom of the insertions of the pectoral fins,
a narrow strip along the soft-rayed dorsal and the anal
fins, and the patch in front of each of the pectoral fins.

641)

Among these lateral plates the middle ones, below or
just in front of the last spinous ray on the back, are
the largest (deepest). Forward and backward the plates
diminish in size; but on the last eight or nine (on the
peduncle of the tail) we find a raised, median carina,
following the longitudinal direction of the body, and
perceptible, though not very distinct, to the twelfth
plate, counting from behind. This median carina ren-
ders the breadth (thickness) of the body no less along
the greater part of the tail than it is in front; but just
in front of the caudal fin the breadth decreases more or
less rapidly. The lateral line, which pierces these plates,
runs from the temporal region parallel to and near the
back. The sixth plate, counting from in front, is the
most constant in position, and unites the 4th dorsal
plate to the top of the ascending disk of the pelvic
bones, with which disk the 5th and 7th lateral plates
are also contiguous in most cases. These three lateral
plates (5 — 7) are also the most persistent, while the
others may be reduced, attenuated, and obliterated, as
we have described above. The first to disappear are
the plates that lie on the posterior part of the ventral
sides and the anterior part of the sides of the tail,
between the seventh plate (from in front) and the ca-
rinated caudal plates: — a form of this description from
Norway has been named liemigymnus by Collett. Later
the fish loses both the foremost plates (in front of the
fifth) and the hindmost, carinated caudal plates, and
the peduncle of the tail undergoes more and more la-
teral compression ( gymnurus ). At the same time the
silvery lustre of the body generally disappears, and the
back develops more or less distinct, dark spots or trans-
verse bands, suggesting a retrogression to the juvenile
characters.

The head is laterally compressed; the cheeks are
parallel; but the snout is attenuated in front even on
the sides. The upper and lower profiles of the head
meet at fairly equal angles (the upper sloping as much
as the lower rises), with the exception of a break formed
by the articulation of the lower jaw. This break is

“ In the North American Apeltes quadracus the pelvic bones are separated from each other behind, as in the Fifteen-spined Stickle-

back; the anterior part of these bones (in front of the ventral spines) is short, broad, and continuous, as in the Three-spined Stickleback,

but without any ascending disk at the lateral margins. The interclavicles are so small that the anterior part of the pelvic bones lies below

the insertion of the pectoral fins, with the ventral spines just behind it.

6 In the variety islandicus (according to Sauvage, 1. c., p. 21, pi. I, fig. 8, ci), as in the above-mentioned gymnurus form from Green-
land, the pelvic plate is both narrow and deeply incised at the anterior margin, just as in the Tcn-spined Stickleback.

c In some specimens of the above-mentioned gymnurus form from Greenland, however, the interclavicles are as narrow and as widely
separated at the middle of their length as in the Ten-spined Stickleback.

650

SCANDINAVIAN FISHES.

most distinct in lean specimens, hardly perceptible in
fat and fleshy ones. Thus in the former the lower jaw
rises somewhat more rapidly in a forward direction,
and projects beyond the true tip of the snout. Some-
times too, especially in lean specimens, the said break
renders the inferior profile of the head behind it almost
horizontal. The length of the head is greater in the
males than in the females, and varies individually be-
tween 23 V2 an<l 28 % of that of the body®. The eyes
are round, and their superior margin lies almost exactly
in the same plane as the interorbital space, which is
flat or very slightly convex and in front narrows regu-
larly, but only slightly; the line from the tip of the snout
to the upper angle of the pectoral fin touches the lower
margin of the pupil. The diameter of the eyes is equal
to the breadth of the interorbital space* and in the fry,
as usual, considerably greater than (sometimes 1 V2
times) the length of the snout, in old specimens equal to
this length or at least 4/5 thereof. The simple, round
nostrils, only one on each side, lie at the upper edges
of the convex snout, about half as far from the eyes as
from the tip of the snout. The length of the snout is
in young specimens about 1/2 (or less), in old specimens
(more than 1/2 dm. long) as much as ?‘j sometimes 2/3 ,
of the postorbital length of the head, which length
measures in young specimens about 3/5 (62 or 63 %), in
old about 2/5 (40 — -48 %) of the total length of the head,
or in the former (though not during the earliest stages)
as much as 1/s (50 — 46 %) in the latter only about 1/3
(37 — 32 %) of the distance between the first spinous
dorsal ray and the tip of the snout. The cleft of the
mouth ascends obliquely and is rather small, the length
of the upper jaw being equal to the diameter of the eyes
or even less than it. Lips are not wanting; but they
are not so tumid as in the Fifteen-spined Stickleback.
Here too, the upper jaw is capable of protrusion by
means of the fairly long nasal processes of the inter-
maxillary bones, which processes extend, when drawn
back, up to the groove in the upper surface of the snout,
a little behind the line of union between the two nostrils.
The length of the lower jaw is about equal to that of
the snout, in young specimens considerably greater than
the latter. The maxillary bones are narrow, only
slightly widened at the hind extremity, and curved
downwards, but about equal in length to the inter-

maxillaries. When the mouth is closed, they are scarcely
covered by the suborbital bones. Among the latter three
bones the middle one is the smallest, and the hindmost,
which is somewhat larger than the foremost (the pre-
orbital bone), more or less completely fills the angle of
the preoperculum, with its lower margin touching the
horizontal branch of the last-mentioned bone, and some-
times with the inferior part of its hind margin also
touching the front of the vertical branch of the same,
while a triangular process, sharply narrowing upwards,
unites this hindmost suborbital bone to the posterior
frontal bone. Thus, the preorbital bone, which forms
the outer margin of the nasal cavity, being also united
to the anterior frontal bone, the orbit is completely
closed; and the naked patches on the cheeks, covered
only by a glossy skin, have the same position as in the
preceding species, though the anterior patch, behind the
corner of the mouth, is much shorter. The jaw-teeth
are of uniform size, set in front in several rows, behind
in one or two, of uniform thickness, cylindrical, more
or less blunt at the tip, and straight or only slightly
curved. The pharyngeal teeth are of similar form, but
somewhat larger and in general more pointed. The gill-
rakers are pectinated, largest in the outer row on the
front of the first branchial arch, where they are about 16
in number, in the other rows small and scattered. The
palatal folds within the jaws are well-developed both
above and below. The tongue is more developed than
in the preceding species, moderately fleshy, and broad
(blunt at the tip). The structure of the opercular appa-
ratus is the same as in the Fifteen-spined Stickleback,
only that the anterior, horizontal branch of the preoper-
culum and the interoperculum are shorter, and that the
ascending process of the suboperculum between the oper-
culum and the vertical branch of the preoperculum is
larger. In form the operculum resembles a semicircle
set on end and superiorly truncated at the end of the
first third of its length or even a little earlier. The
gill-openings are smaller than those of either the pre-
ceding species or the following one, the branchiostegal
membranes coalescing underneath with the isthmus
throughout its length, back to the anterior extremity of
the interclavicles, and thus being separated from each
other. In the typical Three-spined Stickleback (the
trachurus form) all the external bones of the head, as

“ A comparison between 4 males and 8 females gives the result that in the males the length of the head is more,
less, than 27 % of the length of the body.

h In young specimens and in the gymnurus form the breadth of the forehead is comparatively less.

in the females

THREE-SPI N ED STICKLEBACK .

651

well as the other plates and bones that appear at the
surface of the body, are rough with grooves alternating
with granulated ridges, corresponding to the radiating
grooves and striae in the structure of an ordinary scale.
This marking is most handsome and most regular on
the surface of the operculum, where it radiates from the
articulation (the upper anterior corner) of the bone, with
the grooves, in their even and handsome curves, and the
almost smooth ridges terminating at the hind inferior
margin thereof. In the gymnurus form, where the ossified
dermal growths are generally more or less reduced, the
external bones of the head may also be covered to a great
extent by the skin, thus losing their Ganoid appearance.

Of the structure of the fins we know already how
the spinous dorsal rays vary in number and position.
The ordinary place of the first spinous ray lies at about
the end of the first third of the length of the body, of
the second spinous ray at the end of its second fifth,
and of the last spinous ray a little behind the middle
of the body. The most usual supernumerary ray lies
a little in front of the middle of the body. On this
head we must remark, however, that in later life the
first spinous ray is generally removed distinctly in a
forward direction, its average distance from the tip of
the snout thus sinking from 32‘8 to 30‘8 % of the length
of the body while this length increases, on an average,
from 55'8 to 69'7 mm. The spinous rays are of a
more or less broad, triangular form, with concave or
nearly straight sides, and with the front convex and
the back concave. The front is rough and lacunose, like
the above named parts of the dermoskeleton; the lateral
margins are spiniferous for a greater or less distance
from the base out towards the subulate tip. The rough-
ness and the spines partially disappear, however, under
the same conditions as the plate-armour of the body,
in the gymnurus forms. These spinous rays also vary
considerably in length. As a rule the second is the
longest, though the first may sometimes equal it in
length; the maximum length we have found in the latter
case, is 42l/2 % of the distance between the first spinous
ray and the tip of the snout, while the minimum lengths
(in percent of the same distance) have proved to be
16 in the case of the first ray, 17 in that of the second.
The length of the last spinous ray varies between 7 */2
and 17 % of the said distance; and this ray
forms a sharp, almost geniculate curve.

these spinous rays stand straight up; when depressed,
they lie in a straight line with each other, the tip of
each ray sometimes (when the length of the rays is so
great) resting on the base of the ray behind it. The
interspinal plates on which the rays are set, are fur-
nished with a groove in which the latter are partly re-
ceived when depressed. These spinous rays, as well as
the others (those of the anal and ventral fins), are ar-
ticulated in such a manner that they can be locked fast
by the fish at any angle whatever — though generally
and at most in an upright position — and cannot be
depressed by force, unless they are simultaneously lifted
a little way. This result is attained by the presence
at the middle of the base of each spinous ray, in ad-
dition to its true articulation — a trochlea articulation
between each lateral half of the base of the ray and a
transverse hollow in the interspinal plate — of a deep
fissure (articular cavity), into which is fitted a disk-
like, circular process of the interspinal plate. The po-
sition of the former articulation is a little eccentric
relatively to this disk, and still more so relatively to
the articular pivots of the spinous ray, which bound
the said fissure, and which are more elliptical in out-
line. Thus, when the spinous ray is erected, these
pivots with their elliptical friction-surfaces fall into the
circular articular cavity on each side of the central
disk of the interspinal plate. The lower lever of each
pivot attains a greater size than the radius of the disk
and of the said articular cavities. Any attempt to bend
the spinous ray results merely in a pressure partly of
the pivots against the walls of the articular cavities,
partly of the bottom of the middle fissure of the spi-
nous ray against the top of the disk. As soon as the
spinous ray is lifted, however, and the said level* thus
shortened, the pressure ceases, and the ray falls. Each
of the spinous rays has a small, triangular fin-mem-
brane behind it. By means of this membrane the last
spinous ray is united to the base of the first ray of the
soft-rayed dorsal fin, which in this species begins dis-
tinctly in front of the perpendicular from the vent.
This fin is of an elongated triangular form, sloping even-
ly backwards from the first rays, and finally passing
almost imperceptibly into the dorsal margin of the pe-
duncle of the tail. Its length (base) is about equal to
the length of the head in old specimens, in young ones
less than the same"; and its height (longest ray) is

generally
When erected,

The specimens we have received from Archangel, are marked in this respect, as in several others, by the persistency of the juvenile

characters.

652

SCANDINAVIAN FISHES.

about 2/5 of its length. All its rays are branched at
the tip. The peduncle of the tail (the space between
the dorsal and caudal tins) is twice or three times as
long as deep. The anal tin begins perceptibly behind
the perpendicular from the beginning of the soft-rayed
dorsal tin, but ends almost exactly opposite the ter-
mination of the latter. The distance between it and
the tip of the snout is generally greater in the females
than in the males, the length of the head in all the
females we have examined, being less than 43 % of
this distance, in the males more than 44 % thereof.
The soft-rayed anal tin is preceded by a spinous ray,
which is usually exactly similar to the last spinous ray
on the back and joined in the same way to the tin.
The length (base) of the soft-rayed anal tin varies be-
tween 14 and 17 % of the length of the body, and
measures as a rule about 2/3 , exceptionally 3/4 ", of the
length of the soft-rayed dorsal tin. It is otherwise
similar to the latter in structure and form. The ventral
tins contain two rays, the spinous ray and one shorter,
soft ray at the hind margin of the tin-membrane. The
spinous ray is always larger and stronger than any of
the spinous rays on the back, but otherwise similar to
the latter and varying under the same conditions as they.
Thus, its length may vary between more than half (54
or 55 %) and about one-third (sometimes 29 %) of the
distance between the first spinous dorsal ray and the
tip of the snout. The position of the spinous ventral
ray is subject to the following rules: in the males the
distance between it and the beginning of the anal fin
is usually perceptibly less than half of that between it
and the tip of the snout (exceptionally equal to half
of the latter), in the females, on the other hand, usually
considerably greater (60 — 68 % of the distance between
the ray and the tip of the snout), though the young
females come nearer the males in this respect, or are
even similar to them. The same distinction of age and
sex may also be observed in the relation to the length
of the head: in old females the distance between the
spinous ventral ray and the anal fin may lie even
greater than the length of the head, in younger females
(though more than 5 cm. long) we have found it to be
only Ys o;f this length, and in the males we have never
found it to be more than 3/4 thereof. The structure
and position of the pectoral fins are the same as in the
preceding species, with the same thin-skinned, glossy i

patch in front of their vertical insertion; but their tips
are more sharply truncate. Their length varies between
about 13 and 17 % of that of the body, measuring in
the males, according to our measurements, more than
72 % (73 — 90 %) of the distance between the spinous
ventral ray and the beginning of the anal fin, in the
females less than 72 % (7 1 — 47 %) of the same distance.
The caudal fin is straight or slightly forked at the hind
margin. In all our specimens the 106 middle rays are
branched, the others simple. The length of the fin at
the middle is about equal to the postorbital length ot
the head.

The internal organs are similar to those of the
Fifteen-spined Stickleback. The peritoneum is silvery,
but punctated with brown. The air-bladder is com-
paratively large, coasts the whole dorsal side of the
abdominal cavity forward to a line with the first spi-
nous ray on the back, and sends out, according to
Kroyer, “from its under surface a canal to the middle
of the upper surface of the stomach. This canal (pneu-
matic duct) finally shrinks, however, into a ligament.

“The most remarkable characteristic of the Three-
spined Stickleback,” writes Ekstrom, “lies in its varia-
tions of colour. In winter the body is of a pure sil-
very white, with the upper part of the head and the
back blue. In summer the top of the head and the
whole of the back down to the lateral line are blackish
gray. When the spawning-season sets in, the fish ex-
hibits a surprising play of colours. The back first be-
comes brownish, developing darker, transverse bands,
and the silvery white sides show a dash of brassy
yellow. This applies in particular to the females. The
males at the same time acquire a red spot, which be-
gins under the head at the meeting of the branchio-
stegal membranes, and which expands rapidly, the red-
dish tinge usually extending throughout the lower part
of the fish, from the tip of the lower jaw to the vent.
The extent of this red colour varies in different indi-
viduals. In some cases it rises up the sides a little
above the lateral line; in others the whole body is red,
except the highest part of the back, which is then
reddish brown, and the forehead, which is always of a
deep bluish-green colour. The iris is of a handsome
green. We have represented a male in this attire in
Plate XXVIII, fig. 2, and fig. 1 in the same plate is a
reproduction of a female in her winter dress.”

“ In exceptional cases also only 62 /.
b Kroyer found only the 8 middle rays branched.

THREE-SPINED STICKLEBACK.

653

The geographical range of the Three-spined Stickle-
back embraces the whole of Europe, with the exception
of the Danube °, and extends south to the fresh water
of Algiers; but the species has never been found in
the Mediterranean itself, though it occurs in the Black
Sea. In the north of Asia (Siberia) it is common. Both
this species and the next one were found on Behring
Island by the Vega Expedition. In North America the '
Three-spined Stickleback lives at least as far south as
Southern California and New York. In Greenland and
Iceland it has long been known. In Spitsbergen, on
the other hand, it has not been met with. The Three-
spined Stickleback is thus a Boreal-arctic species; and
as we have remarked above, it occurs in its most ty- j
pical form, with most developed armour, chiefly in the
sea, while the strength of the plate-armour and of
the spines is generally reduced in brackish and still
more in fresh water.

Throughout Scandinavia the Three-spined Stickle-
back is found not only in all the lakes, large and
small, streams, and brooks, but also in both island-
belts, the Eastern and the Western. It is often met
with in collections of water so small and so isolated
that it appears difficult, if not impossible, to explain
the manner in which it has been conveyed thither,
or the sources from which it derives its support. It
is fondest of calm water, and in summer frequents
shallow spots close in shore, especially where the sun-
shine has free play. Here it leads a merry life, and
one may often see it leap several centimetres out of
the water; while at other times it keeps still at the
same spot, as though there were nothing in the world
to disturb it. But in a moment it is all life and spirit,
and darts off a few paces with the speed of an arrow.
In stormy weather it is tossed on the waves, and has
thus been cast even into a boat. Towards autumn it
retires to deeper spots on the shores of channels, estu-
aries, or larger inlets. In late autumn and at the be-
ginning of winter, however, it roams about in large
companies, which in many parts of our eastern archi-
pelago yield rich harvest to the fisherman. When the
cold is most severe, it probably lies packed in large
shoals at the bottom, at least in the island-belt; for
when, as sometimes happens, it is taken in the seine
at this time of year, it is caught in great numbers.
Sociability is also a characteristic trait of the 'Three-
spined Stickleback; solitary specimens are seldom met

with, and it generally lives collected in large or small
companies.

The Three-spined Stickleback devours with avidity
worms, insects, larva), small mollusks, and small fry.
Though of insignificant size it is a greedy and bold
fish-of-prey, and furiously attacks fishes which are even
larger than itself. It sometimes does damage by no
means to be despised; and in their internecine battles
it happens often enough that a Stickleback rips up
the belly of his antagonist with one of his ventral spines.

The spawning-season of the Three-spined Stickle-
back, which period is attended by several remarkable
phenomena, occurs about the end of June or even earlier.
Even in May both the male and the female begin to
change colour in the manner above described. This is
the prelude of the spawning, which begins as soon as
.the fish have acquired the proper colour. The shoals
which now ascend to shallow, weedy shores in order
to spawn, distribute themselves in such a manner that
the males and females are separated. Each of the males,
which seem to be much fewer than the females, chooses
a certain spot, at some distance from his neighbours,
where he exercises undisputed authority. Here he builds
the nest to which he entices one female after another
to lay her eggs. During this period the males are vio-
lently jealous of each other, and a slight action on the
part of one of them is often all that is required to
goad his fellow to madness and provoke a duel.
These combats are waged as follows: the two rivals
rush with the speed of arrows against each other, deal
a powerful side-stroke with their sharp ventral spines,
and hasten with undiminished speed each back to his
own domain. After a few onsets the superiority of the
stronger combatant is demonstrated, his territory is ex-
tended, and he signalises his triumph by a splendour
of colours, while the vanquished lays aside his brilliant
dress as though overcome with shame. While the males
disport themselves in these chivalrous tournaments, or
rather fight for their nests, the females swim about in
long troops of greater or less strength outside the
battle-ground, and now and then a male selects his
temporary mate from the company. The female that
heads the troop, swims forward with rapid darts, fol-
lowed by the others, suddenly stops, and assumes a
vertical position, with head turned towards the bottom.
The others assemble round her and range themselves
in the same manner, as densely packed as possible.

a “Der Stickling fehlt dem Donaubeeken im engeren Sinne.” Hckl., Kn., Siisswasserf. Oesterr. p. 41.

654

SCANDINAVIAN FISHES.

When she has thus collected the troop around her, she
suddenly deals a blow that scatters the whole crowd
in an instant. This sport is often repeated, but the
rapidity with which they disperse, renders it impossible
to observe whether it is always the same female that
takes the lead, or whether they change places. These
operations are continued, as long as the sun is high

Fig. 163. Male of Gasterosteus ciculeatus in front of its nest.
Natural size.

in the heavens, for 4 — 6 days, according to the weather.
It seems more than probable that during these evolu-
tions the females lose some roe, which adheres to water-
plants, and that this is fertilized by the males that,
perhaps only for the time being, have not built any
nest for the eggs. Benecke has also ascertained that
under certain circumstances, as for example when he
finds a suitable crevice or secluded nook among the

water-plants, the male does not build any nest properly
so called. Thus we have to deal with two methods of
spawning, the free spawning, in which the eggs are
developed where they fall, among the water-plants, and
the more connubial method, in which the eggs are de-
veloped in a nest built by the male. But in any case
the nest-building is one of the most interesting parts
of the life of the Three-spined Stickleback, and one
which many have been in a position to observe (fig. 163).

The earliest account we possess of the nest-building
of the Sticklebacks is nearly two hundred years old.
In 1721 the English writer Richard Bradley tells
us" that one Sergeant Hall had seen a Stickleback
build a nest, and had secured the latter in order to
present it to Bradley, who gives a figure of it and
says that he “supposes it was rather composed to lav
the spawn in, than for a lodgment for the fish itself."
Since that date several naturalists have watched the
building operations of the male Three-spined Stickle-
back in its native haunts; but the most accurate ob-
servations of its habits have been made in aquaria,
where the Three-spined Stickleback constructs its dwell-
ing in the same way as when at liberty. Brehm has
described its method of building in his work on The
Life of Animals b as follows:

“Warrington®, CostfT, and Evers have observed
their captive specimens at work. During the spawning-
season, as we have mentioned, the male wears a dress
of the most gorgeous colours, and lie displays his en-
hanced activity and vivacity in other ways as well. As
soon as he has decided on a certain spot at the bottom,
he first drags together some roots and other parts of
several aquatic plants. These pieces may be larger than
himself, and are often fetched from considerable distances.
With great pains he tears off comparatively large bits
even from living plants, tests their weight by letting
them fall to the bottom, and employs in his work only
those that sink quickly, rejecting all that prove too light.
The building-materials are thus carefully selected, piled,
and still further adjusted, until the little architect finds
them arranged according to his wishes. To attach the
nest to the bottom he employs sand or small pebbles®.

“ A Philosophical Account of the Works of Nature , p. 61, pi. VIII, fig. II. It may be questioned, however, whether this nest be-
longed to the Three spined Stickleback or to the following species.
b Thierleben. 2:te Aufl., 3:te Abth., 2:te Bd., p. 88.
c Ann., Mag. Nat. Hist. 2nd ser., vol. X, p. 27 6.
d Mem. Sav. Etrang. Inst. Fr., tome X (1848), p. 575.

e The Three-spined Stickleback may also build its nest above the bottom and among the branches of the water-plants, though
this method of building seems to be more characteristic of the next species.

T II B EE- S T I N ED STICKLEBACK.

rPlie tubular chamber is constructed, and the form and
durability of the whole nest attained, l>yr the Stickle-
back swimming slowly over the parts already in po-
sition, and at the same time gluing them together
and cementing them. Evers distinctly observed how
the builder, when he had added new layers, shook his
tins, raised his head, bent his body upwards, and slid
his belly over the structure, emitting at the same time
a drop of a viscid substance, which could be clearly
distinguished in the water, and the effects of which
might immediately be perceived in the now cemented
building-materials". At times he shook the building
and then pressed it together again; at times he kept
swimming over it. With his fins, which he kept in
continual and rapid motion, he produced a current,
and thus washed away from the nest the pieces that
were too light and the loose stalks, which he then
took up again and tried to fit in more durably. It
took about- four hours to procure the various building-
materials; but at the expiration of this time the out-
lines of the nest were ready. Its completion, the re-
moval of the parts that are too light, the arrangement
of the separate stalks, the plaiting of their ends, and the
addition of the sand to weigh them down, require several
days. While the Stickleback is building his nest, he
thinks only of his work and endeavours merely to pro-
vide against any interruption in its progress or hind-
rance to its completion. He labours indefatigably and
watches with suspicion every creature that approaches
the nest with or without evil intentions, whether it be
another Stickleback, a newt, a water-beetle, or a larva.
A water-scorpion ( Nepa ) in one of Evers’ aquaria was
seized by the cautious builder thirty times or more, and
carried in his mouth over to the opposite side of the
aquarium.

The size of the nest varies pretty considerably, de-
pending both on its situation and on the materials of
which it is composed. As a rule it seems to be of the
size of one’s fist. It is generally ellipsoidal and entirely
closed above, but furnished at the ends with an en-
trance and an exit. At first only the entrance is
visible, but subsequently, exactly opposite it, we dis-
cover the exit. When the Stickleback has finished his

655

building-operations, he endeavours to attract a female
to the nest. Warrington says that a completed nest
excites the attention of the female; but Coste asserts
that the male sallies forth to guide her thither, and
that he ushers her into the nest with a shower of ca-
resses. The last statement, however, is also accepted by
Warrington. The male distinctly shows his delight at
the arrival of the female, swims round her in all di-
rections, enters the nest, sweeps it out, returns in a
moment, and tries to drive the female in by thrusting
at her from behind with his snout. If she will not
obey of her own accord, he also employs his spines,
or at least his caudal fin, to overcome her reluc-
tance; but in case of need another female is fetched.
If tlie male succeeds in persuading a female to enter
the nest, she lays a few eggs within it, according to
Coste only two or three, and then bores a hole through
the wall of the nest on the side opposite to the en-
trance, and departs. The second opening in the nest
does not exist until it is formed in this manner. The
current that now flows through the openings, is of
benefit to the eggs. On the following day the male
sets out again in quest of a new female, whom he com-
pels by kindness or force to lay eggs, and repeats this
process until a sufficient number of ova is procured.
During or immediately after the laying of the spawn
he enters the nest, rubs his side against that of the fe-
male, and then glides over the eggs in order to ferti-
lize them.

From this hour his zeal and watchfulness are re-
doubled. He has to guard and defend the eggs against
every aggressor. Every Stickleback that approaches is
furiously attacked and put to flight, whether it be a
male or a female, for both are equally dangerous ene-
mies of the eggs, and the latter is perhaps even more
greedy of the ova or the new-hatched fry than the
former. Until the young have emerged from the eggs,
the male shows his care in other ways as well. With
his mouth he repairs every damage to the nest, and
often stations himself in front of or within the cham-
ber, keeping his pectoral fins in vibration and thus
renewing the water in the nest, as though he knew
that- the eggs require fresh oxygen.

" M6bius ( Nature , vol. XXXIX, No. 998, Dec. 13, 1888, p. 168) saw the male .Stickleback keep spinning round the nest new threads,
which originated from the urinary bladder. The chemical reactions showed that these threads consist of mucin, which is secreted, however,
not by the urinary bladder, but by the kidneys. The section of a kidney, treated with osmic acid and coloured with hematoxylin, showed
that only a few of the cells lining the tubnli urinifevi partake in the development of the mucin, while most of them do not undergo any
change of this description. After the end of the spawning-season, no mucinous cells can be' found in the kidneys, which are now less tumid.

Scandinavian Fishes.

656

SCANDINAVIAN FISHES.

At last the time for the ripening of the eggs ap-
proaches, and new troubles come. He has now to pro-
tect and defend the helpless young. In Warrington’s
aquarium a female laid her eggs at night on the 8th
of May, and on the very next day the mother was
driven away by the male. The latter continued his
guardianship until the 18th of the same month, and
then began suddenly to destroy the nest, leaving only
a few stalks at the foundation. All the mud and
sand that covered the eggs was removed in his mouth, |
and a space about 8 centimetres in diameter carefully I
cleared. Astonished at such conduct on the part of the
nest’s protector, Warrington took a magnifying glass to
ascertain the cause, and discovered the new-hatched fry.
Henceforth the male Stickleback never ceased swimming
in every direction over the cleared space, redoubling his
vigilance, and driving away every other fish that came
too near. As the young gained in size and strength,
they seemed anxious to leave the nest, but the father
intercepted them time after time, carefully taking the
truants in his mouth, and returning them to their
quarters. Not until they were strong enough to swim
with ease, did the father’s watchful care gradually begin
to slacken, but at last, when they were quite capable
of procuring their own food, he left them entirely to
shift for themselves.”

When the spawning is over, the handsome colours
of the fish gradually disappear, and its normal tem-
perament returns.

From the great numbers in which this species oc-
curs, especially during certain years, we might suppose
it to be very prolific: but this is not the case. The
female has no more than 110 — 150 eggs in both ova-
ries; and as the restricted number of the males leads us
to assume that not all the eggs are fertilized, the fe-
cundity of the species cannot be set very high. In
spite of this, as we have already mentioned, in certain
years enormous shoals of Three-spined Sticklebacks are
met with. Several reasons have been suggested to ex-
plain this fact. The most probable explanation of these
cases in general is Cuvier’s, that the years in which
these large shoals appear have been more than usually
favourable to the process of reproduction; but Sundevall
explains a portion of these variations by the fact that,
during the years in which the Herring-fry are plentiful
in the outer part of the island-belt, the Stickleback re-

tires thither and is comparatively scarce nearer to the
mainland; while the largest catches of Sticklebacks on
the coast of the mainland seem to be made during
periods when the opposite is the case.

The Three-spined Stickleback, which is surpassed in
voracity by few predatory fishes, becomes very fat, and
is probably rapid of growth, soon attaining its full size.
But the statement of certain authors, that it does not
live more than 3 years, seems to require corroboration.

This fish is hardly used at all as human food —
though in England, together with Herring-fry, it of-
ten tempts consumers under the name of Whitebait.
Steller and Tilesius state that the Stickleback which
occurs on the coast of Kamchatka, is delicious when
boiled and an excellent ingredient in soup. Still, even
the Kamchatkans themselves reject it for their own use,
but keep it to feed their dogs in winter. The single
specimens which in Scandinavia are caught at almost
every haul of the seine for other fish, are generally
thrown among the offal and used as food for swine. Pre-
datory fishes and seafowl, however, eagerly devour the
Stickleback, and it is the principal food of the Garpike.

When the Three-spined Stickleback is taken in
quantities, which happens only in certain years, it is
boiled down into oil, a manufacture to which so early
a writer as Westbeck (1753) directed the attention of
his countrymen. A barrel (4 V2 bushels) of fresh Stickle-
backs yields more than one gallon of clear oil. The foot
or sediment that settles at the bottom of the cauldron
is an excellent manure, two barrels (9 bushels) of this
sediment, mixed with a sufficient quantity of water,
being reckoned equal to 10 lasts0 of dung.

The only method of fishing employed exclusively
for this fish is simple enough. The fishery commences
at the beginning of November — some fishermen state
that the Stickleback does not come in before the ground
has been covered with snow — and continues until the
inlets and channels have begun to freeze. Towards
evening, when it has been observed that the Stickle-
backs are collected in a shoal, the presence of which is
betrayed at sunset in calm weather by the surface of
the water being rippled as if by fine rain or a shower
of sand, two fishermen repair to the spot in a boat. In
the bow of the boat is erected a kind of fire-pan (fig.
164), on which dry wood is laid and kindled. When
the boat has reached that side of the shoal towards

Sw. parlass = a load drawn by two horses.

THREE-SPINED STICKLEBACK.

657

which the heads of the fish are turned, one of the two
fishermen keeps the boat still by means of a stake,
which is thrust into the bottom, while his companion
scoops the fish out of the water into the boat with a
tine hand-net (tig. 165). In this way several boat-loads
may be taken in a night. In December, 1887 a cor-
respondent writes from the island-belt of Ostergothland :
“'fhe violent storms of this autumn have driven into 1
our creeks and bays a multitude of “prigg" or spigg
(Sticklebacks), for which a fishery has now been carried
on for some time. The Sticklebacks must have fed extra-
ordinarily well this year, for they are so fat that 2l/3
gallons of oil may easily be extracted from a barrel of ;
fish. The owners of the stretches of shore where the
Sticklebacks swim in shoals, esteem themselves especial-
ly fortunate, for the Stickleback-fishery takes the palm
of all the fisheries among our islands. On the strip of

approaches them, they generally remain quite still at
first for a few moments, apparently unconcerned; but
suddenly a fish starts up, casts itself to one side, and
joins some comrade disturbed in the same manner, or
takes its place in an army already formed and advanc-
ing in the immediate neighbourhood, an army which
like a long, moving Avail roves along the bottom, at
first, as a rule, in a circle round the light. Gradually
the advancing Avail increases in height, length, and
breadth, Avhile it circles hither and thither, as if to col-
lect more and more stragglers, in curves of greater or
less extent, until at last, Avhen the army appears to be
sufficiently strong, it suddenly dashes up and assembles
beneath the light. Here the crush that noAV folloAvs is
tremendous, and the movements of the fish culminate
in a strange confusion, exactly as though they intended
Avith their immense numbers to overpoAver and vanquish

shore belonging to a small crofter, for example, the value
of the catch has proved to be 600 croAvns (Tod), and
on the coast-line of a farm 2,000 croAvns ( £ 1 10). The
shoals press on Avithout a pause to their destination, so
that the fisherman may keep seining the Avhole day at
the same place, and the catch in spite of this be quite
as great in the last draught as it Avas in the first".”
In his annual report for 1869 Baron G. C. Ceder-
strom, the pisciculturist, describes the singular behavi-
our of the Sticklebacks during this fishery as folloAvs:
“In autumn the Sticklebacks generally repair toAvards
evening to the shalloAv Avater near the shores, Avhere they
keep still, resting on the bottom as though the spinous
rays of the ventral fins hoav served as feet. In the
morning they again return to someAvhat deeper water.
It is highly interesting to Avatch the effect of the torch-
light on them, Avhile they are at rest. When the light

° See the Dagens Nyheter for the 15th
" Qvod nomen “paullo urbanius”
c So called because it thrives in

the fire. In spite of the hand-nets hoav plied, they
still rush on, undaunted as before. When their num-
bers are so feAv that the fisherman does not think it
worth Avhile to use the net there any longer, he moves
to another spot between 50 and 200 yards off, or some-
times even nearer his former station; and here the same
occurrences are repeated.'

The Stickleback bears different names in different
parts of SAveden. On account of its spinous fin-rays it
is called pigg , spigg , and stagg , in Scania hornstagg, in
the eastern archipelago skot spigg \ horntagg, and liornfisk.
On account of its bright and tin-like colour it is knoAvn
in Avinter as tennfisk , tennfisk med spjut (Speared Tinfish)
etc. In Gothland, according to Lindstrom, it is coupled
with the following species under the name of baingyl
or bainkyl0.

(Ekstrom, Smitt.)

of December, 1887.
quam vulgare Skitspigg : Retzius.

small collections of water ( bain = bone, gyl — pool), Tr.

658

SCANDINAVIAN FISHES.

THE TEN-SPINED STICKLEBACK OR TINKER (sw. smasfiggen).

GASTEROSTEUS PUNGITIUS.

Plate XXVIII, figs. 3 and 4.

Free spinous rays on the hack usually 9 (7a — 12). Branchiostegal membranes united underneath into a fold

posteriorly free from the isthmus.

R. hr. 3; D. 7“— 121 0 — 12; A. 18 — 10 ; P. 9—10; V. '/\ ;
C. x -|- 1 4" 1 0 4- 1 4" x ; Vert. 32 — 33.

Syn. Gasterosteus aculeis in dorso decern, Art., Ichth., Gen., p. 52;

Syn. p. 80; Spec., p. 97; Lin., Fn. Suec., ed. I, p. 104.
Gasterosteus Pungitias , Lin., Syst. Nat., ed. X, tom. I, p. 296;
Fn. Suec., ed. II, p. 119; Retz., Fn. Suec. Lin., p. 338;
P all. (G aster acanthus), Zooyr. Ross. Asiat., tom. Ill, p. 228;
Cuv., Val. ( Gasterosteus ), Hist. Nat. Poiss., vol. IV, p. 506;
Ekstr., Vet. Akad. Handl. 1831, p. 302; Nilss., Prodr.
Ichth. Scand., p. 86; Ekstr., Skand Fisk., ed. I, p. 20,
v. Wright, tab. IV, fig. 2; Kr., Damn. Fiske, vol. I, pp. 188
et 593; Sundev., Stockh. L. Hush. Sallsk. Handl., H. 6 (1855),
p. 79; Nilss., Skand. Fn., Fisk. p. 110; Thomps., Nat.
Hist. Irel., vol. IV, p. 89; Gthr, Cat. Brit. Mus., Fish.,
vol. I, p. 6; Sieb., Siissiv. Mitteleur., p. 72; Mgrn, Finl.
Fisk-Fn. (disp. Helsingf.), p. 15; Lindstr., Gotl. Lans Hush.
Sallsk. Arsber. 1866, p. 15 (sep.); Coll., Forh. Vid. Selsk.
Christ. 1874, Tillregsh., p. 13; ibid. 1879, No. 1, p. 4;
N. Mag. Naturv. Christ., Bd. 29 (1884), p. 49; Day, Journ.
Linn. Soc. Lond., vol. XIII (1876), Zook, p. 110; Malm,
Gbgs, Boh. Fn., p. 373; Seidl., Fn. Balt., p. 128; Winth.,
Nat. Tidskr. Kbhvn, ser. 3, vol. XII, p. 4; Feeders., ibid.,
p. 74; Mor., Hist. Nat. Poiss. Fr., tom. Ill, p. 169; Day,
Fish. Gt. Brit., Irel., vol. I, p. 244, tab. LXVIII, fig. 4;
Mela, Vert. Fenn., p. 279, tab. IX; Fatio, Fn. Vert. Suisse,
vol. IV, part. I, p. 98; Hansen, Zool. Dan., Fiske, p. 30,
tab. V, fig. 3; Storm, Norsk. Vid. Selsk. Skr., Trondhj.,
1883, p. 15; Mob., Hcke, Fisch. Osts., p. 65; JoriD., Gilb.,
Bull. U. S. Nat. Mus., No. 16, p. 393; Lillj., So., Norg.
Fisk., vol. I, p. 363.

Gasterosteus occidental's, Cuv., Val., 1. o., p. 509. Gast. dekayi
+ G. nebulosus, Ag., Lake Superior , pp. 310 et 311. Gast.
concinnus, Rich., Fn. Bor. Amer ., p. 57. Gast. mainensis ,
Storer, Bost. Journ. Nat. Hist., vol. I, p. 464. Gasterosteus
pungitius+G. bur gundi anus -f G. Iceuis (ex Cuvier, Regn.
Anim., ed. 2, tom. II, p. 170) + G. lotharingus + G. breviceps,
Blanch., Poiss. d. eaux douces Fr., pp. 238, cett. ; Gasterostea
pungitia + G. burguncliana + G. occidentalis + G. dekayi + G.
blanchardi + G. mainensis + G. Icevis + G. lotharinga + G.
breviceps +G. concinna+G. globiceps, Sauvage, Nouv. Arch.
Mus. d’Hist. Nat. Par., tom X, pp. 29, cett.

Obs. As appears from this list of synonyms, it has also been
proposed to base a number of independent species on the variations of
the Ten-spined Stickleback. The lack of constancy is, however, just
as great here as in the preceding species; and to judge by our present
knowledge of the forms, only the East Asiatic variety ( Gasterosteus
sinensis, Guiciienot, see above), with its fully-plated lateral line and
its large, comparatively few spinous rays on the back, which are equal

in length to the diameter of the eyes or still longer, deserves to be
especially distinguished as an intermediate form between this species
and the Three-spined Stickleback.

The Ten-spined Stickleback is the smallest fresh-
water fish of Europe. Its ordinary length when full-
grown is about 5 or 6 cm. Lilljeborg, however, speaks
of specimens 81 mm. long from the moats of Upsala
Castle. In form it is usually shallower (more elongated)
and more terete than the preceding species, the depth
of the body at the insertions of the ventral tins being
on an average* 6 about 18 % of its length (as compared
with 20 % in the preceding species), and the breadth
straight across the upward lateral processes of the pelvic
bones on an average 58 — 62 % of the said depth (as
compared with 52 — 51 % in the preceding species). But
in this respect no constant difference can be traced, and
the largest specimens of the Ten-spined Stickleback as
a rule come nearer and nearer the Three-spined species.

The Ten-spined Stickleback is generally without
any external trace of plate-armour on the sides of the-
body; but in most cases the lateral line is raised on
the sides of the tail in the form of a carina, rendering
the breadth of the peduncle of the tail considerably
greater than its depth, and now and then showing plate-
like ossifications in the skin. But quite as frequently
this carina is really membranous, and sometimes hardly
a trace of it is visible, the base of the caudal fin thus
acquiring a terete appearance. Especially in the south-
ern parts of the geographical range of the Ten-spined
Stickleback this last form seems to be the commonest
one, and it has been described by Cuvier under the
name of Gasterosteus Icevis. The plates at the dorsal
margin and along the base of the anal fin are hidden
in the skin; but the former are more concave on the
upper surface than in the preceding species, thus form-
ing a groove both deeper and broader, into which the
spinous dorsal rays may be depressed. In number these
spinous rays vary within the limits given above —

“ Abnormal specimens with only 2 or 4 free spinous rays on the back are mentioned by Day (1. c.).

6 In our specimens, which are between 38 and 60 mm. in leDgth.

TEN-SPINED STICKLEBACK.

659

sometimes even the greater number of them may lie
wanting. They do not stand at right angles to the
back, but lean alternately towards the sides. They are
also set, not in a straight line, but in a zigzag (v.w).
They are subulate in form, somewhat curved but only
slightly widened at the base, with sharp points and
smooth edges. In size they are fairly equal, though the
hindmost ray is generally somewhat larger than the
others" and equal in length to the spinous ray in front
of the anal tin. All of these spinous rays are usually
shorter than the diameter of the eyes; but the anal
spine may sometimes equal this diameter in length.
Each of them has a small, triangular fin-membrane be-
hind it. In this species too, the ventral spines are the
largest, their length varying between 2/'5 and l/i of the
distance between the first dorsal spine and the tip of
the snout. At the outer margins, at least in full-grown
specimens, they are usually granulated or even spiny
with osseous tubercles arranged in several rows. Their
articulations are constructed in the same manner as in
the preceding species.

The pelvic bones are narrower than in the Three-
spined Stickleback, and the sinus in their anterior mar-
gin is deeper but narrower. Their upright lateral pro-
cesses are somewhat broader and more obliquely cut at
the top. The interclavicles are also narrower than in
the preceding species, but of about the same length,
the pelvic bones lying entirely behind the perpendicular
from the insertion of the pectoral fins.

The head as well as, in most cases, the whole body
is more elongated (shallower) than in the Three-spined
Stickleback; but its relative length is about the same,
being on an average about 1/4 of that of the body. Its
structure is also the same as in the Three-spined Stickle-
back, with the exception of the character that lies in

the above-mentioned arrangement of the branchiostegal
membranes.

The soft-rayed dorsal and the anal fins are also of
a low, triangular form. The first ray in each of them
is simple but distinctly articulated. They are fairly
analogous in position, the beginning of the dorsal fin
never lying in front of the perpendicular from the hind
part of the vent. The base of the soft-rayed dorsal fin
is always perceptibly shorter than the head, but varies
between 20 and 24V2 % of the length of the body.
The length of the base of the anal fin varies between
about 16 and 1 9 1 /2 % of that of the body.

The hind margin of the caudal fin is straight, some-
times slightly forked or even convex, an indication of
the transition to Gastrcea. Its length at the middle is
always perceptibly less than the postorbital length of
the head. The pectoral fins are similar to those of
the Three-spined Stickleback both in form and rela-
ive size.

The internal organs resemble those of the Three-
spined Stickleback; but the air-bladder is somewhat
narrower. K rover found some instances to corroborate
Cuvier’s observation to the effect that the intestine runs
from the stomach straight to the vent, instead of pre-
viously forming a circular coil, as it generally does
both in this species and in the two preceding ones.

The relation between this species and the Three-
spined Stickleback is of especial interest when we con-
sider the external differences of. growth and sex, which
show that the Ten-spined Stickleback represents partly
the earlier (lower) stages of development and partly
the male characters. The following table of averages
contains the changes of growth most important in this
respect, together with the sexual differences especially
conspicuous in the Ten-spined Stickleback.

Average in

10 speci-
mens of
Gcister-
osteus
pungitus.

14 speci-
mens of
Gaster-
osteus
aculeatus.

8 speci-
mens of
Gastrcea
spinachia.

Obs.

Length of the body expressed, in millimetres... -

48.7

62.7

107. 0

—

Longitudinal diameter of the eyes in % of the length of the snout

102.3

91.8

42.7

juv. sen.

Postorhital length of the head in % of its total length

49.8

43.2

43.8

d

O

A

>

Length of the base of the anal fin in % of the distance between the anal spine and the tip of the snout..

33 9

24.7

23.3

juv. sen.

» » s head in % of the distance between the first dorsal spine and the tip of the snout

93. o

79.o

100.2

juv. sen.

n n n n ?? n 11 5? ii ii ii anal ii ii ii ii ii ii ii -

45.9

-ll.o

49.8

d*>9

Postorbital length of the head in % of the distance between the first dorsal spine and the tip of the snout

4(3.4

34.3

43.9

tf>9

Distance between the ventral spines and the anal spine in % of the length of the head. ...

79.3

89.3

44.9

cf <9

„ „ „ „ ,, „ „ „ „ „ „ „ ,, distance between the ventral spines and the tip of the snout

55.2

58.fi

30. o

<? <9

ii ii ii ii ii ii ii ii ii i ' ii ii ii ii ii ii spine ,, ,, ,, ,, y,

35.9

36.9

22.9

c? < 9

EkstrSm (Vet. Akad. Handl., 1. c.), however, found the second ray longest and the hindmost shortest

660

SCANDINAVIAN FISHES.

In the first three relations the development has ad-
vanced in the same direction from pungitius both to
aculeatus and to spinachia — in each case, except the
second, furthest in the last species — , but in the fourth
spinacliia has retained the most primitive character of
the Sticklebacks. In the last five relations spinacliia
is the nearer to pungitius, and stands in each case,
except the sixth, on the opposite side to aculeatus.

The direction of development is fixed in aculeatus by
the female, in spinachia by the male characters.

The coloration of this species is quite as varied as
that of the preceding one, and is affected by the differ-
ent seasons of the year, the different moods of the fish,
and the different bottoms and lights of the water in
which the fish lives. Here too, we find darker (Plate
XXVIII, fig. 4) and lighter forms. In winter the upper
part of the head and the back are of a bluish drab, and
the sides silvery white, with extremely fine, dark dots.

In summer, on the other hand, the head and the upper
part of the whole body are of a marked olive-green.

The lower part of the body is now light green, with a
dash of brassy yellow, which shades more into red
under the head and at the bases of the pectoral fins.

The whole body is thickly strewn with distinct, black
dots. It is this dress which is depicted in Plate XXVIII,
fig. 3, painted from a large specimen, a female taken
in the neighbourhood of Haparanda in June, 1832.
During the spawning-season the males are black on the
sides of the belly.

The Ten-spined Stickleback has the same geogra-
phical range as the Three-spined species, but in Europe
does not penetrate so far south. Moreau sets the south-
ern limit for the range of the Ten-spined Stickleback in
France at hit. 45° N. According to Fatio this species
is wanting in Switzerland. Heckel and Kner do not
include it among the fishes of the Austrian Empire. It
is also unknown in Italy (Canesthini, Giglioli); but it j
occurs in the Black Sea, according to Pallas, and Nord-
mann“ says that it is found in several streams in the
south of Russia. On the other hand, its occurrence in
Greenland and Iceland is uncertain. Neither Reinhardt
nor Lutken includes it in their lists of the fishes of Green-
land, though Hansen says that it lives there; and Faber
had never seen it in Iceland. Richardson’s concinnus
belonged, however, to the extreme north of North Ame-

“ Demid off’s Travels, tom. Ill, p. 382.

* Cat. Collect. Fish. U. X’. Nat. Mus., Gt. Intern. Fisher. Exhib.

c See Ransom, Ann., Mag. Nat. Hist., ser. 3, vol. XVI, p. 449.

rica (Saskatchewan and Great Bear Lake) and, Pallas
knew the species from Siberia and the Sea of Okhotsk.
The Vega. Expedition found it together with small Three-
spined Sticklebacks, as we have mentioned, on Behring
Island. Bean6 assigns it to Alaska ; and it was already
known to Cuvier from Newfoundland ( occidentolis ).

In Scandinavia the Ten-spined Stickleback is com-
mon from the extreme north to the southernmost pro-
vinces, both in most of the Swedish lakes and streams
and on the Baltic coast. In Norway, according to Col-
lett, its occurrence is only sporadic and confined to
fresh and brackish water.

The habits of this species are essentially the same
as those of the preceding one. It prefers clear, running
wafer. It is sociable, and therefore lives collected in
large shoals during the greater part of the year. It also
joins company with the Three-spined Stickleback, when
the latter sets out on its autumn wanderings. In tem-
perament it seems to be very sluggish, but can move
with great rapidity. Its food consists of worms, insects,
fish-roe, and bits of grass, which it devours with avi-
dity. We often find this little fish, like its congener,
very fat.

The spawning-season generally occurs in June and
July — according to Benecke even in April — when the
fish makes its way to small brooks and rills, where
during the spawning we find it packed in multitudes.
Males and females are promiscuously assembled, and try
to crowd as near to the grassy bank as possible. The
roe is orange and, in comparison with the size of the
fish, coarse, the eggs being about 1 mm. in diameter.
It is deposited either on the grass or in a nest, for here
as in the preceding species the male builds a shelter
for the eggs until the exclusion of the fry0. As long
as the spawning lasts, and as long as the male lias to
defend the eggs, the fish is not at all timid, and suffers
itself to be taken in the hand.

The Ten-spined Stickleback, which in different lo-
calities bears different names, e. g. benunge , signalling
etc., is utilised in the same way as the Three-spined
species. It is said to yield a finer oil. It is taken
chiefly in November, when it collects in shoals together
with the Three-spined Stickleback.

(Ekstrom, Smitt.)

London 1883, p. 19.

LOPIIOBRANCHS.

661

PHYSOCLYSTI LOPHOBRANCHIT*.

Physoclysts with tufted branchial laminae.

This single character is enough to divide from the
rest of the Physoclysts a series extremely remarkable
in many respects, which contains two families, and of
which our Pipefishes and the Sea-horses of warmer re-
gions are well-known types.

The gills of the Teleosts generally consist, as is
well known, of a great number of branchial I a mi me —
on the first branchial arch of a Cod 3 dm. Iona:, for
example, we can count about 100 pairs — which are
arranged in a double row on the outside (the convex
side) of each branchial arch, like the teeth of a comb
but in pairs, each lamina alternating with (obliquely
opposite) the other lamina of the pair, though they are
united to each other for a greater or less distance from
the base. Each of these laminae (fig. 1 66) is furnished
on each side (above and below) with numerous, trans-
versely-set lamellae — in the Cod we can count as many
as 1,000, but in other Teleosts they are much fewer, in
the Gudgeon ( Gobio fluviatilis) for example, only 50 or
60 — which are honeycombed by the capillary network of
the respiratory vessels with a view to the oxygenation of
the blood and its transmission from the branchial arteries
to the branchial veins. In the Lophobranchs the num-
ber of the branchial land me is considerably reduced, in a
Pipefish, for example, being only 5 — 8 pairs on each
arch, the minimum number on the foremost and hindmost
of the four branchial arches, the maximum on the two
middle arches. But to compensate for this the com-
paratively few branchial lamella: are considerably en-
larged, being visible even to the naked eye, or at least
with the help of a weak magnifying-glass. They are
largest at the middle of each lamina, their size decreas-
ing regularly towards the base and the tip thereof; and
the lamina itself is curved, not unlike a peacock’s feather.
“By means of this arrangement,” says Retzius, “the
branchial laminae (fig. 167) acquire an appearance some-
thing like that of bunches of peacocks’ feathers, set
alternately and touching each other with the convex
part of their curves. When we see the two rows of
these bunches from without (fig. 168), the figure also
reminds us somewhat of the foliation of Salvinia .”

Fig. 166. A pair of branchial lamina? attached to a section of a
branchial arch in the Cod. b, the lamella?. Magnified and schema-
'* tized. After Tit. Williams.

Fig. 167. A pair of branchial laminae attached to a section of a
branchial arch in Syngnathus. Magnified. After A. Retzius.

Fig. 168. A branchial arch with its alternating lamina? and their
lamellae in Syngnathus. Magnified. After A. Retzius.

L’ordre des Lophohranches, Cuv., Regne Anim., ed. I, tom. II, p. 155.

662

SCANDINAVIAN FISHES.

This character is accompanied, however, by several
other peculiarities, most of which call to mind the Hemi-
branehs. The body is scaleless, but in general completely
covered with Ganoid-like plates, though these may
sometimes be thin and coated with skin, so that they
become indistinct. They are arranged in longitudinal
and transverse rows. The number of the transverse
rows is usually equal to that of the vertebrae, to the
processes of which they are more or less closely united;
and in consequence they generally form distinct seg-
mental rings round the bod}'. In most of the Lopho-
branchs and especially in the Pipefishes the body is
highly elongated, like that of the Eels or the Flute-
mouths, and the snout is produced as in the latter
forms. The branchial arches are incomplete, consisting
in each ease of a single cartilaginous bone; the epi-
branchial and upper pharyngeal bones, as well as the
hypobranchial bones, are wanting. The interclavicles
are present, but are not externally visible.

The organs of motion are feeble and the muscula-
ture as a whole is not much developed; but in the motory
muscles of the fins Rollett" has been enabled to observe
a fact that seems calculated to possess the greatest im-
portance for the explanation of the structure and func-
tion of the striated muscles. Each of these muscles
consists, as is well known, of a number of small fibres,
corresponding to large cells, each with its envelope, a
thin membrane, and its contents. Within the thin mem-
brane, the sarcolemma , that confines the muscular fibre,
we find two essentially distinct substances, the first,
which has been named by Kuhne rhabdia, the staff -
substance, divided into a number of cubical, parallelo-
pipoid, virgulate, or lamellar parts, arranged beside
each other in longitudinal or transverse rows, but in
general at very minute intervals, which are filled by
the second substance, the sarcoplasma, a kind of pro-
toplasm with nuclei distributed among it. The great
muscles of the body in the tiny Sea-horse are also of
this structure; but Rollett has pointed out a remark-
able difference in flic relative proportions of these two
substances in the muscles that set the fin-rays in vibra-
tion. Here, where the movements are too rapid for the
eye to follow them, the sarcoplasm preponderates greatly
over the rhabdia. Up to the present time science has
not fully succeeded in elucidating the significance and

a TJeber die Flossenmuskeln des Seepferdchens ( Hippocampus
XXXII (1888), p. 233.

co-operation of the two substances in the contraction
of the muscles. But, as only the sarcoplasm together
with the sarcdlemma comes in immediate contact with
the end apparatus of the fibrillse of the motory nerves,
we may at least conjecture both that the physiological
significance of the two substances is different, and that
the operation of the nervous power varies according to
the different quantity and distribution of the two sub-
stances in a muscular fibre. Roli.ett’s observation of
the Sea-horse — together with several other observations,
e. g. of the wing-muscles in insects — entitle us at
least to the assumption that extremely rapid muscular
motion may be promoted by the more advanced deve-
lopment of the sarcoplasm.

The Lophobranchs are characterized, it is true, by
a considerable reduction of the true organs of motion;
but one genus, Solenostomus from the Indian Ocean and
Australia, the type of a distinct family, is fully equip-
ped in this respect, in particular with a high anterior
dorsal fin, a large caudal fin, and both long and broad
ventral tins. All the other genera, which together form
the family Syngnathidce, are destitute both of the first
dorsal fin and of the ventral fins; the anal, caudal, and
finally even the pectoral fins are reduced in most of
them, and entirely disappear in some forms. When the
caudal fin has disappeared, the tail becomes a prehensile
organ, more or less developed, which fishes of this
description, e. g. the well-known Sea-horses, curl round
the branches of seaweeds or round other suitable objects,
in order to hold themselves fast.

All these fishes are feeble creatures, which depend
for their protection chiefly on their resemblance to the
surrounding objects. Our small Pipefishes closely re-
semble the thread-algae ( ulvce ) to which they often
attach themselves, and several Sea-horses develop foliafe
growths sometimes of the most fantastic shapes, by
means of which they acquire a deceptive likeness to
the seaweed. They must seek their food among the
weakest minute animals, for their toothless mouth is
most like a sucking tube (cf. above, pp. 263 and 264,
on the internal folds of the mouth in the Gobies).

In the method of their reproduction all the Lopho-
branchs' of which we have any information on this
head, show the peculiarity that they carry the impreg-
nated eggs and, until they can help themselves, the

antiquorum') und tiler MuskelstrucUir im Allgemeinen, Arch. Mikr. Anat.

LOPHOBRANCHS.

663

excluded embryos on their ventral side. In the great
majority of cases it is the males that have to perform
this duty, just as the males of the Pycnogonoid fa-
mily carry the lumps of eggs on their feet. But in
Solenostomus the ventral fins of the female coalesce

longitudinally with the ventral side so as to form a
sac, in which the fertilized eggs are stored and deve-
loped.

Of the two families of this series there occurs in
the Scandinavian fauna only the

Fam. S YNGJN ATHIDfE.

The greater part of the margin of the branchiostegal membrane united to the body , leaving only at the top , above
and just behind the end of the operculum , a small , fissure-like gill-opening on each side.

This family, as defined" by Kaup/j, includes the
great majority of the Lophobranchs, with about 150
species, distributed among 15 genera. All the Syn-
gnathoids that occur in the Scandinavian Fauna, belong
to Gunther’s0 subfamily Syngnathince, which is dis-
tinguished from the Sea-horses (Hippo campince) — with
their descending (prone) head, elevated occiput, and
more or less tumid body — by an evenly elongated
form of body, with the head prolonged in the same
direction as the trunk and with the latter only slightly
thicker than the caudal part. In the Syngnathince , too,
the tail is never so developed into a prehensile organ
as in the Hippo campince.

The above-mentioned rings formed by the plate-
armour of the body are composed (fig. 169) on the

Fig. 169. Schematic transverse section of a Syngnatlms. Magnified.
After Moeeatj. A, transverse section of the trunk; 1 — 1 ', 2 — 2',

and 3 — 3' the paired plates; 4, the ventral plate. B , transverse

section of the tail.

trunk (in front of the vent) of 7, 8, or 9, and on
the tail (behind the vent) of 6 or 4 plates, so ar-
ranged that the former make up 3 or, at the end of
the trunk, 4 pairs and an unpaired ventral plate, which
disappears behind the vent, while either the uppermost
or the lowest pair disappears either behind the end of
the dorsal fin or even behind the vent. As the paired

plates are bent at greater or less angles, longitudinal
carinae are formed on the sides of the body, three on
each side of the trunk or even to the termination of
the dorsal fin, and two on each side of the tail, where
its section is quadrangular. But in many cases the said
angularity is slight or absent, while the skin that covers
the plates grows comparatively thick, rendering the
body smoother, the plates and carinae more indistinct
or even imperceptible.

As in the rest of the Lophobranchs and a great
proportion of the Hemibranchs, the snout is elongated
and bears at the tip the gape, which is almost vertical
when closed, with the ascending uncler-jaw forming the
extreme margin of the head in front. In the Deep-
nosed Pipefish (Syngnatlms typhle) the elongation of
the snout is produced in the following manner. The
ethmovomerine part is elongated like a staff, and coasted
below by the long and narrow parasphenoid bone, while
the frontal bones extend forward above in the form of
long and narrow covering-bones over about half of the
said elongation. The hyomandibular bone is an oblong,
quadrangular but irregular, vertically set disk, which
is united at a right angle below with the abnormally
developed os symplecticum, which is directed forward,
extends below the eyes, and sends out a branch obli-
quely upward towards the lateral ethmoid (prefrontal)
bone, while a second, still longer, horizontal branch
meets a process in a backward direction from the
quadrate bone. This horizontal branch of the symplec-
ticum is partly naked (without covering bones) exter-
nally, but is covered behind and below, throughout the
greater portion of its extent, by the preoperculum. The
vertical (posterior) branch of the preoperculum lies

“ The family Syngnathidce as established by Bonaparte (Cat. Metod. Peso. Eur., 1846, p. 89) was differently defined, so as to
exclude the Sea-horses.

h Cat. Lophobr. Fish. Brit. Mus. (1856), p. 5.
c Cat. Brit. Mus., Fish., vol. VIII, p. 153.

Scandinavian Fishes.

84

G64

SCANDINAVIAN FISHES.

outside the hyomandibular bone and is united above
to the inferior margin of the suborbital bones. The
obliquely ascending branch of the symplecticum, on the
other hand, is separated by a space, occupied by the
masticatory muscles, from the two posterior suborbital
bones, which bound the orbit below, but is united to
the hind superior corner of the anterior suborbital (the
preorbital) bone, where the latter meets the lateral
ethmoid bone. The foremost suborbital bone forms the
greater part of the side of the snout, being united in
front as a covering bone to the ento-(meso-)ptery-
goideum and the quadrate bone, but leaving behind the
latter an opening in the middle for about a third of
the depth of the snout; the middle suborbital bone is
united below to the symplecticum; the hindmost sub-
orbital bone both to the symplecticum and, behind, to
the preoperculum. We refer to the opercular apparatus
a narrow, lancet-shaped, thin bone which lies along the
inside of the horizontal, forward branches of the pre-
operculum and the symplecticum and the horizontal,
backward branch of the quadrate bone. This lancet-
shaped bone is united by ligaments behind to the upper
part of the ceratohyoid bone and in front to the an-
gular part of the lower jaw. The latter union clearly
shows that the bone must be an interoperculum0, cor-
responding most nearly in form and position to the
interoperculum in Tetrodon for example, though here
it lias passed within the symplecticum and become united
behind to the upper part of the outside of the cerato-
hyoideum, instead of retaining its union to the sub-
operculunT', which is here reduced to an extremely
thin disk within the lower margin of the operculum.

The lower jaw is remarkable for its extraordinary
depth behind and its sharply curved and toothless dental
part. The maxillary bones are comparatively well-
developed and their hind (lower) extremity is expanded;
but the toothless intermaxillaries are small, styliform,
and without nasal processes. The palatine bones are
shorter than the maxillaries and of fairly uniform
breadth. At the anterior angle of the union of each
palatine bone to the top of the ascending branch of the

“ J. Playfair McMurrich {On the Osteology and Development
(Oct. 1883), p. 623), assumes that this bone is a metapterygoid.

b See, however, below, on the ligamentous connexions between
in Silurus.

c See for example our figure of Syngnathus acus. The ent
tuberance on the throat just behind the perpendicular from the eye
body, B ) the urohyoid bone extends back from the ceratohyoid bones

d Anatomisk undersokning dfuer ndgra delar af Syngnathus A

quadrate bone we find the narrow pterygoid bone, which
is bent at an obtuse angle, and behind the said union,
close to the ridge of the snout, the entopterygoid bone,
which is united in the same manner to the quadrate
bone.

The tongue is wanting, and the glossohyoid bone
can hardly be discerned; but the urohyoid bone is com-
paratively long. When the last bone is drawn back,
the anterior (lower) end of the ceratohyoid bone with
the tip of the whole branchial apparatus is forced down-
wards", the cavity of the mouth being thus widened
and filled with a fresh supply of water. In this manner
the Lophobranchs pump in their food together with the
water needed for their respiration. The method in
which the transverse dermal folds {vela) just behind the
jaws assist to this end, has already been touched upon.
The gill-rakers are small, denticulate, and pectinately
arranged along both margins of the inside of the branchial
arches. Their function, the prevention of food and fo-
reign substances from penetrating into the gill-cavities
when the lish swallows, has already been pointed out
by Retzius''.

The intestinal canal (tig. 170) is simple in all the
Lophobranchs, but shortest in the Syngnathince. In the
Hippocampince (the Sea-horses) the intestine forms at
least a few curves, but in the Syngnathince it is straight
or nearly so, thus rendering the whole digestive canal,
from the pharynx to the vent, equal in length to or
only slightly longer than the abdominal cavity. The
lines of demarcation between the oesophagus, the sto-
mach, and the intestine are only internal, and consist
merely in the difference of the mucous membrane,
while the beginning of the last division is marked as
usual by the mouth of the gall-duct. But the small
intestine is divided from the rectum by an annular
valve. No pyloric appendages can be traced. The me-
senterium is wanting or only partly developed, in front
and behind, to bind the stomach and intestine to the
dorsal wall of the abdominal cavity. Both the ovaries
and the testes are paired, simple, and elongated. Their
discharging ducts are short and distinct down to their

of Syngnathus Peckianus — Quart. Journ. Micr. Sc., u. ser., No. XCII

os epihynle , the interoperculum, and the angular part of the lower jaw

re figure (.4) shows the ceratohyoid bones depressed to a pointed pro-
; and in the lower figure (the anterior part of the ventral side of the
and between the opercula.

cus och Opludion, Vet. Akad. Handl. 1833, p. 146.

LOPHOBRANCI-IS.

665

A: Male, witli the abdominal wall opened and folded outwards and the
marsupium, in which appear the cavities left by the eggs, distended.

h, liver; va, air-bladder, with the anterior, thin-walled part pointed
and obliquely divided from the posterior; t, testes; pa, anal fin; vu,
urinary bladder; i, intestine, opened for a short distance at the vent;
ug, urogenital aperture, behind (below) the vent, which is opened.

B: Female, with the abdominal wall opened and folded outwards.

/), liver; vf, gall-bladder; pa, anal fin; 7,, intestine, at the mouth of
the gall-duct coming from the right side (in the figure the left); ?2,
posterior part of the intestine, opened in order to show the valve that
divides the intestine from the rectum, which is also opened as well
as the vent; ov, ovaries; vu, urinary bladder; tig, urogenital aperture.

opening just behind the vent. The urinary bladder is
long and narrow, lies on the left side, and coasts the
left sexual organ. The air-bladder is large and coasts
the upper wall of the abdominal cavity for at least
one-half or two-thirds of the length of the latter.

The ribs are wanting. Among the other peculiari-
ties of the skeleton the structure of the shoulder-girdle
is especially remarkable. The posttemporal bone (fig.
171, pt) coalesces with the mastoid bone ( epoticum ) of
the skull and is covered in a Ganoid fashion by a plate-
like layer, pointed behind. The supraclavicular bones are
wanting; but the clavicle (cl) is all the larger. This
bone is T-shaped, with the upper (horizontal) bar united
to the transverse processes of the first two vertebra?.
The first transverse process is directed backwards, and
so long that all the anterior upper branch of the cla-
vicle may be contiguous with it: while the second of
these processes, which is comparatively short, and only
slightly expanded at the top, adjoins the angle between
the horizontal and vertical branches of the clavicle. The
outside of the clavicle is covered by the three plates on
each side of the body which form the first (foremost)
ring; and in Syngnatlms acus the inner prong of the
vertical shaft, which is forked at the bottom, in the T-
form of the clavicle, is supported on the first abdominal
plate belonging to the same ring. This plate is wanting
in Syngnatlms typhle , where the support is therefore
given by the lower lateral plate. Only the anterior
margin, however, of the middle lateral plate in this ring
is united to the middle of the shaft in the T-form of
the clavicle, the rest of this plate lying as a. covering over
the muscular mass of the pectoral fin, in the middle of
which mass we find the extremely thin scapulo-coraeoid
disk (sc) together with the four basal bones (fig. 171,
has) of the pectoral fin. This disk is not divided here
into its ordinary component parts (the shoulder-blade
and the 'coracoid bone), and is extended between the
hind part of the upper (horizontal) branch of the clavicle
and the inner prong of the vertical shaft of the clavicle,
without touching this bone at any other point. At this
lower end of the clavicle, however, within the lower end
of the scapulo-coraeoid disk and above the inner side of
the first two abdominal plates in Syngnatlms acus , lies
one interclavicle (id), which meets both the clavicle and
the said disk (fig. 171, sc). Neither in Syngnatlms acus
nor in S. typhle have we succeeded in finding more
than one such interclavicle, though according to Par-
ker the former species possesses two.

666

SCANDINAVIAN FISHES.

The skeleton is otherwise extremely simple and only
slightly ossified, with the vertebrie elongated and their
processes straight and of uniform breadth, or slightly
broader at the top; but in the vertebrae above which
the dorsal fin is attached, the upper spinous processes
are divided sagitally (in the longitudinal direction of the
body) into three or four divergent branches. Distinct
haemal arches appear only at the beginning of the
caudal region, where a few may be found.

By far the greater number of the Syngnatliince are
marine fishes; they are met with everywhere, where
seaweeds cover the bottom. Out at sea they accom-
pany the tufts of sargasso-weed, or drift about at the
surface with the current and the waves. Some of
them, however, also live in brackish water, and a
few species occur in fresh water alone. The species'
that occur in Scandinavia may be distinguished as
follows :

1

Fig. 171. Right scapular arch of Syngnathus acus , seen from within, 7 times the natural side. Principally after Parker.
pt, posttemporal bone; cl, clavicle; slix, lower lateral plate in the first ring ( scutum laterale inferius annuli primi ); sli.,, lower lateral plate
in the second ring; svl, ventral plate in the first ring; sv„, ventral plate in the second ring; icl, interclavicle; slmx, middle lateral plate in

the first ring; has , basal bones of the pectoral fin; sc, scapular disk.

A: The adult, fishes without pectoral
fins. The males carry the impreg-
nated eggs in a layer of mucus on
the ventral side of the trunk.
a: Beginning of the dorsal fin far
in front of the perpendicular
from the vent, its distance from
the tip of the snout being at
most about 4/5 (less than 85 %)
of the distance between the

vent and the same point Nerophis cequoreus.

b: Distance between the dorsal fin
and the tip of the snout at least
9/io °f drat between the vent
and the latter.

aa: Length of the trunk and
head (distance between the
vent and the tip of the snout)
more than 45% of the length
of the body Nerophis ophidion.

bb: Distance between the vent
and the tip of the snout
less than 35 %> of the length

of the body Nerophis lurnbriciformis.

B: The adult fishes furnished with pec-
toral fins. The males" carry the
impregnated eggs in a canaliculate
sac on the ventral side of the tail.

a: Length of the head more than 15

% of that of the body. Syngnathus typlile.

b: Length of the head less than 14
% of that of the body.
aa: Length of the head greater
than that of the base of

the dorsal fin. Syngnathus acus.

bb: Length of the head less
than that of the base of

tire dorsal fin Syngnathus rostellatus.

In our Scandinavian species.

PIPEFISHES.

667

Ever since Fries" pointed out the remarkable fact
that the larva? of Nerophis lumbriciformis are furnished
with fully developed and actively employed pectoral fins,
it has been evident that the former of the two above
groups, into which we have after Fries divided the
Scandinavian Syngnathince, has been developed by a
retrogressive metamorphosis from forms agreeing more
closely with the better equipped types. The Needle-
fishes ( Syngnatlii Ophidii in Fries) are also distinguished
by a considerably smaller head, in adult fishes less than
710 of the length of the body, and a smoother body,
without or at least with only indistinct longitudinal
Carina?'. It is also a rule among the Lophobranehs that
the larvae and the young fishes have sharper carinae or
even spines on the plates of the body, where the older

fishes show fainter traces thereof. Thus the Needle-
fishes, in this respect too, represent older (more advan-
ced) stages of development. This cannot prevent them,
however, from being regarded as less fully equipped
even in respect to the covering of the body. An exotic
genus within this group, Protocampus , which otherwise
comes fairly near Nerophis, also retains the larval ver-
tical fin even in the case of full-grown specimens, at
the dorsal edge both in front of and behind the true
dorsal fin, and at the ventral edge of the trunk. Again,
the manner in which the male Needle-fishes carry their
eggs, must surely be regarded as more primitive than
the corresponding arrangement in the rest of the Syn-
gnathi. We therefore begin our description with the forms
which Fries called Tdngsndllor ( Syngnatlii Marsupiales).

Genus SYNGNATHUS'.

The adult fishes furnished with pectoral, caudal , dorsal, and anal fins. The uppermost row of plates on each,
side of the trunk terminates posteriorly at the end of the dorsal fin or just in front of this point. Vent situated

impregnated eggs in a canaliculate sac underneath the tail.

in the anterior half of the body. The males carry the

Some fifty species, known and defined with greater
or less certainty, are contained within this genus, which
is spread over all the seas of the Tropical and Tem-
perate Zones, and is represented even in some of the
rivers and lakes of the Tropics.

The upper marginal carina on each side of the tail
runs forward in this genus below the posterior part of
the upper lateral carina of the body, on each side of the
dorsal fin, either to meet the middle lateral carina of the
trunk or to disappear above its termination. The egg-sac
of the males is formed in this genus in the following
manner. The anterior part of the lower marginal carina
of the tail grows obliquely downwards on each side for
a greater or less part of its length, and along the un-
der surface of the free margin develops a dermal fold,
a continuation of the anal mucous membrane. These
dermal folds on each side of the body converge towards
each other, but without coalescing, their free margins
being closely approximated to each other, but leaving a
slit, which the fish can widen or firmly close at pleasure.
“When the spawning-season approaches,” says Ekstrom
of Syngnathus typhle, “the foliate lids (dermal folds)

that close the marsupium of the male, become swollen,
and the marsupium is gradually filled with a white,
clear, and thick mucus, which serves as a bed for the
eggs, and decreases in quantity as the young grow,
until, by the time they are large enough to swim and
make independent progress though the water, little or
none of it remains. One day in the month of July 1
was present at the hauling of a seine in the island-
belt, and secured a male of this species that had fully
developed young in the marsupium. I at once con-
structed a dam with stones from the beach, and the
fish was set in the pool soon after it had been taken
from the seine. After it had swum to and fro for a
while, it opened the marsupium by a downward move-
ment of the tail, whereupon the young crept out one
after another and swam under and on both sides of the
male, but always kept close to its body at a. little di-
stance from the marsupium. As soon as I tried to capture
the male, it made a sudden movement, at the same
time bending the body in an arch upwards, and the
young at once crept into the marsupium, the lids of
which Avere then shut. The same experiment Avas re-

“ Metamorphos, cinmarkt lios Lilia ILafsnalen , Vet. Akad. Handl. 1837, p. 59.

b Artedi, Iclithyol., Gen. Pise., p. 1. In his works, however, this genus represented the whole of the modern series Lophobrancliii.
In Aristotle the genus was called (SeXovrj, a name which Athen^eus subsequently transferred to the modern Rhampliistoma. Gaza translated
/SeA.ot'tj by acus, which was the most general name of the genus during the Middle Ages; but Belon called it Typhle or Typliline.

668

SCANDINAVIAN FISHES.

peated several times, but both the male and the fry
invariably behaved in the same manner.

As the genus is defined in the above diagnosis, it
contains six of Kaup’s" and DumekilV genera, two
of which, Siplionostoma and Syngnathus, are retained
by Gunther®. The distinction between these two genera
is stated to consist in the character that in Siplionostoma '
the lower covering plates of the shoulder-girdle (the
lowest pair of lateral plates in the first ring) are only

loosely joined to each other at the middle of the belly;
while in Syngnathus they are firmly coalescent, and
their juncture is usually covered by the first ventral
plate, which is always wanting in Siplionostoma , this
genus thus having an oblong, rhomboid a t patch, co-
vered only with skin and without plate, at the middle
of the belly just behind the shoulder-girdle. The dif-
ference is of little importance, at least in the Scandi-
navian fauna, which contains only three of these species.

THE GREAT PIPEFISH (sw. STORA TANGSNALLAN cl).

SYNGNATHUS ACUS.

Fig. 172.

Snout comparatively narrow and shallow. Length of the head less than 14 % of that of the body and at most
24 % of that of the tail , which is more than twice the length of the trunk. Distance between the dorsal fin and
the tip of the snout at most 39 % of the length of the body, and the length of its base less than 1/3 of this
distance and also less than the length of the head. Length of the marsupium of the male more than twice that

of the head. Hind margin of the caudal fin rounded.

B.

Fig. 172. Syngnathus acus from the Great Fishing-bank, S. W. of Bergen, where the depth was 100 — 150 fathoms, taken by Fisherman
Andersson of Bobus! fin in 1874. A, of the natural size; B, the head seen from below, natural size.

R. hr. 2; I). 38-42; A. 3—4; P. (10) 11—12 (13); C. (9)
10; Vert. 66; Ann. 62 -66 = (20 — 22) + .-r.

Syn. Syngnathus corpore medio heptagono, cauda pinnata, Art.,
Ichthyol., Spec. Pise., p. 3, No. 3; Lin., Fn. S'uec., ed. I,
p. 126, No. 336.

Syngnathus Acus (p. p.), Lin., Syst. Nat., ed. X, tom. I, p.
337; (“Laminae corporis trunci 20, cauda? 43”); B h., Fiscli.
Deutschl., pt. Ill, p. 113, tab. XCI, fig. 2; Flmng, Brit.
Anim., p. 175; Jen., Man. Brit. Vert., p. 484; Yarr.,
Hist. Brit. Fish., ed. 1, vol. II, p. 325; Fr., Vet. Akad.
Handl. 1837, p. 27; Kr. ( Siphostoma ), Damn. Fiske, vol.
Ill, i, p. 692, Sund. ( Syngnathus ), Ofvers. Vet. Akad.
Forh. 1852, p. 85; Nilss., Skand. Fn., Fisk., p. 684; Kp.,
Cat. Lophobr. Fish., Brit. Mus., p. 41 ; Gthr (p. p.), Cat.

Brit. Mus., Fish., vol. VIII, p. 157; Dum., Hist. Nat. Poiss.
(Nouv. Suit, a Buff.), tom. II, p. 552; Coll. (p. p.), Vid.
Selsk. Forh. Christ. 1874, Tillsegsh., p. 200; ibid. 1879,
No. 1, p. 101; Winth., Naturh. Tidskr. Kbhvn, ser. 3, vol.
XII, p. 53; Hcke (p. p.), Arch. f. Naturg. 46 (1880), I,
p. 332; Mor., Hist. Nat. Poiss. Fr., tom. II, p. 42; Day
(p. p.), Fish. Gt. Bril., Irel, vol. II, p. 259, tab. CXLIV,
fig. 1; Storm, Vid. Selsk. Skr. Trondhj. 1883, p. 42; Lillj.,
So., Norg. Fisk., vol. Ill, p. 454.

Syngnathus Typhle, Bl., Fiscli. Deutschl., pt. Ill, p. 112,
tab. XCI, fig. 1; Malm (p. p.), Ofvers. Vet. Akad. Forh.
1852, p. 84.

Ohs. Artedi’s description ( Descr . Spec. P/sc., 1. c.) leaves no
room for doubt that he really distinguished between the two species

a Cat. Lophobr. Fish., Brit. Mus. 1856.

b Hist. Nat. Poiss., tome II (Nouv. Suites a, Buffon, Paris 1870).
c Cat. Brit. Mus., Fish., vol. VIII, pp. 154 and 155.
d Fries.

GREAT PIPEFISH.

G69

that according to his opinion had already been represented by Ron-
delet ( De Pise., lib. VIII, cap. II 1 1, p. 229) in distinct figures, but
combined in one chapter (as one species). The first of these figures
evidently represents the species called Typhle in Belon (Nat., Divers.
Poiss., p. 446); the second (lower) figure should thus correspond to
the species called by Linnveus Syngnathus acits, assuming that this
species occurs in the Mediterranean, from which locality Rondelet
had procured his fish". After Artedi’s observation: “Numerus in-
cisurarum seu crenarum transversarum, in hoc genere, omnino atten-
detidus est’ , when we employ the Linnaaan name, we must first be
guided by the character which Linnaeus gives, and which is drawn
from the number of the rings on the body. Both Fries (1. c.) and
Sundevall (1. c.) have adequately shown that Linnaeus otherwise con-
founded these two species, and cannot be regarded as an authority on
this question. As Sundevall also points out, Donovan, in his descrip-
tion of Syngn. typhle, was the first after Artedi clearly to define the
distinction between the two species, and the first to give to the name
of acus the employment which has subsequently and rightly been
observed by the English faunists.

The Great Pipefish attains a length of about half
a metre (Storm). From Dynekil (Bohuslan) the Royal
Museum has received through Mr. C. A. Hansson spe-
cimens up to 442 mm. long. The elongated, wliip-like
body shows a distinct break, especially in adult speci-
mens, at the boundary between the trunk and the tail;
and the trunk is thickest, as usual, in the females. The
greatest depth of the body, at the middle of the trunk,
may sometimes rise to nearly 5 % of its length, but is
usually only 3 or 4 % thereof, or in the females about
13 Vu — 15 %, in the males about 11 — 12 % of the length
of the trunk* * * * 6. The greatest breadth is as a rule about
4/5 of the greatest depth. At the beginning of the tail
the depth of the body in old specimens is at most
about 3 %, sometimes only 272 %c of its length, and
from this point it decreases regularly to the shallow
base of the caudal fin. The length of the trunk behind
the gill-cover is abc ut 28 — 26 % of that of the body;
and the length of the tail, including the caudal fin,
varies between about 59 and 61 % of that of the body.

The plate-armour is distributed in such a manner
that 21 or 22, sometimes, in young specimens, 20 rings
(including the ring of the shoulder-girdle) belong to
the trunk, and 43 — 45 to the tail. Here, as in the
other Syngnathi, these plates are striated with trans-
verse ridges and grooves between them, starting from
the middle carina, which is finely dentated. The upper-

most plates, which meet at the middle of the back, are
almost square and bent at a slightly obtuse angle, ren-
dering the dorsal side flat or slightly concave. In the
middle lateral row on the trunk the plates are more
distinctly hexagonal or even octagonal, and bent at a.
very obtuse angle. In the lowest lateral row on the
trunk the angle of curvature varies according to the
greater or less distension of the belly, but the form of
the plates is more like that of those in the middle
lateral row. The middle ventral row also consists of
hexagonal or octagonal plates. The two hindmost rings
on the trunk are without ventral plate, its place being
occupied by the anal region. On account of the form
of the plates two rows of diamond-shaped patches, co-
vered only' with skin, are left on each side of the trunk
and also on its ventral side; but on the back there is
only one row of these rhombs, which generally extend
across the whole dorsal plane, but sometimes, especially
in front, are indistinct, the contiguous plates being per-
fectly rectangular. The middle lateral row of the trunk
ends in the hindmost ring of this region, as a rule
obliquely below the beginning of the row that forms
behind the dorsal fin the upper lateral margin of the
tail, and below the posterior part of the dorsal fin (be-
hind the vent) a middle lateral row belonging to the
tail. But sometimes the last-mentioned row wedges
itself into the hindmost ring on the trunk, which ring
may thus contain four plates on each side. The dif-
ference in this respect is, however, so irregular that
we sometimes find the former to be the case on one
side of the body, the latter on the other side. The
four sides of the tail behind the dorsal fin are con-
structed in the same manner as the dorsal plane of
the trunk; but the ventral side is broader than the
dorsal, which is parallel to it, the difference being
greatest in old specimens. The marsupium of the males
extends under 24 — 26 rings.

The length of the head measures 12 — 1 3 x/2 % of
that of the body. At the occiput it is only slightly
shallower than the beginning of the trunk and not
very sharply marked off therefrom. Here as in the
Syngnatln in general, the trunk is furnished at the

° Whether the Atlantic Syngnathus acus also occurs in the Mediterranean, is not yet decided with certainty. Moreau had no know-

ledge of it from this locality, and Canestrini (Fauna DPtalia, Pesci ) does not include it among the fishes of Italy. On the other hand,

Moreau gives Syngnathus ettion, and Canestrini Sygn. ta’nionotus, which seem to stand in the same relation to the Mediterranean Syngn.

rubescens, as Syngn. rostellatus to Syngn. acus.

6 In young specimens about I1/., din. long the greatest depth of the body may be only about 21/., o/o of its length and 10 % of the
length of the trunk.

c In young specimens still less, sinking at least to 21/- "l of the length of the body.

SCANDINAVIAN FISHES.

670

extreme front of its dorsal plane with two unpaired
plates, which have been called occipital, because their
middle carina forms a continuation of the occipital
carina. These plates also lie so far forward that the
anterior one partly covers the crest of the occiput, and
the small gill-openings lie beside the front part of the
posterior plate. Thus the hind part of the head with
the gill-openings lies to a great extent within the limits
of the trunk. The anterior occipital plate is shorter
than the posterior, sometimes only half as long. The
true occipital carina is hardly as high as these so-called
occipital plates; but still the occiput rises considerably
from the forehead, which in its turn slopes forward
towards the snout. The occiput is also convex, and the
sides of the hind part of the head are considerably
tumid, on account of the alveated form of the oval
opercula. These bones are striated with smooth ridges,
radiating upwards and downwards and leaving between
them rows of small cavities in the surface; and an hori-
zontal ridge, granulated by a row of small round tu-
bercles and scarcely half as long as the operculum, runs
back from the articulation of this bone. The occiput,
the top of the head, and the temples are cavernous
in the same manner, but with coarser and more irre-
gular depressions. The snout, on the other hand, is
smooth, with the exception of the bars (carina?) that
follow its dorsal side. One of these bars, the dorsal
carina of the snout, runs in a line with the occipital
carina, which has terminated and been obliterated on
the concave forehead. This bar is coasted on each side
by another, an immediate continuation in a forward
direction of the strongly elevated upper orbital margin,
which is continued backwards in the same manner by
a ridge that runs obliquely up towards the beginning
of the anterior occipital plate. The orbital margin itself
is also continued downwards and sharply defined in old
specimens, in which the praefrontal bone also forms a
prominent knob at the middle of the anterior orbital
margin. All these ridges are granulated at the free
margin, more or less distinctly and most so in old spe-
cimens, by a row of small round tubercles. In front
of the said praefrontal protuberance, which is not very
distinct in young specimens, lies a triangular depression,
within which an obliquely longitudinal elevation of the
skin bears a small round nostril at each end. When
the mouth is closed, the snout is somewhat compressed
from the sides, with the breadth gradually decreasing
in front; but when the mouth is open, it is more terete

and, when seen from above, of more uniform breadth.
The tip of the snout is turned upwards, and rises per-
ceptibly above the ascending tip of the lower jaw.

The length of the snout varies considerably in
different individuals, but especially according to age.
In young specimens, up to a length of 2 dm., it may
be only slightly greater or even somewhat less than half
the length of the head; but in older specimens it mea-
sures about 56 — 59 % thereof. The variation is as
great, but reversed, in the case of its least depth, which
we have found in young specimens to measure 20 — 18
% of its length, while the corresponding proportion in
old specimens varies between about 15 and 13 %. As
a rule the least depth of the snout is 2/s of the length
of the lower jaw, which is about the same as the dia-
meter of the eyes. The postorbital length of the head,
occupied almost entirely by the gill-cover, measures about
V8 (31 — 34 %, exceptionally 36 %) of its entire length.

As we have mentioned above, the branchiostegal
membranes are entirely coalescent, below and on the
sides, with the front of the shoulder-girdle. The gill-
openings, with their crescent- shaped lids, are small hori-
zontal crevices, about half as long as the diameter of
the eyes, and situated in a line with the latero-dorsal
carina? of the trunk, which begin just behind them.

The only paired fins, the pectorals, occupy an obli-
quely vertical position about half-way up the body, and
are of almost uniform breadth, with sharply rounded
tip. Their rays are simple and unarticulated, somewhat
widened at the tip, and usually 12, exceptionally 10 —
13, in number. The length of the longest (middle)
rays is always less than the depth of the body at the
beginning' of the tail.

The dorsal fin begins on the last or the penulti-
mate ring of the trunk, at a distance from the tip of
the snout which we have never found to exceed 39 %
(37 — 381/2 %) of the length of the body, and which is
greatest as a, rule in the females. It is of almost uni-
form height, though it rises slightly in a backward direc-
tion for about three-fourths of its length and then sinks
again. Its rays (usually 35 — 42, exceptionally 43 — 45),
like those of the other fins, are simple and unarticulated;
they are somewhat broader at the tip and compressed
(flattened in the longitudinal direction of the body).
It more or less entirely occupies 10 or 9 rings of
plates, and the length of its base, which is always
shorter than the head, measures 11 or 12 % of that
of the body.

GREAT PIPEFISH.

671

The anal fin lies just behind the vent, at a
distance from the tip of the snout not exceeding
41 % (39 — 40 V2 %) of the length of the body. It is
extremely small, resembling a thin and narrow dermal
flap, a little expanded towards the tip, and consists of
three (according to Moreau four) rays, which do not
attain a length of even half that of the longest rays of
the dorsal fin. It is often so entirely enveloped by the
marsupium of the males as to be invisible externally.

When expanded the caudal fin is fan-shaped, with
rounded hind margin. As a rule it contains 10 rays,
more seldom 9. Its length at the middle is about 3 %
(2'8 — 3’2 %) of that of the body, or at most about 1/i
(26 — 22 %, least in old specimens) of that of the head.

The coloration is reddish or yellowish brown, with
about 12 or 13 broad, dark brown, transverse spots
across the hind part of the head, the back, and the
tail. These spots are intersected, however, by longi-
tudinal, oblong, smaller spots of the ground-colour.
The sides of the snout are also marked in a similar
manner. The ventral side is yellow or red. The lower
part of the gill-cover gleams with a silvery lustre. The
caudal fin is dark brown. The anal and pectoral fins
are transparent, but the edges of the rays are dark
brown. The dorsal fin resembles the last-mentioned
fins, but has several transverse spots arranged in rows
on the rays.

The geographical range of the Great Pipefish ex-
tends along the whole west coast of Europe south of
the neighbourhood of Trondhjem, and according to
Gunther the British Museum has also received speci-
mens through Sir A. Smith from the Cape of Good
Hope. On the English coast the species is very com-
mon. It seems hardly probable that it occurs in the
Mediterranean, as . Kroyer has already remarked. It
rather appears to be represented there by one or per-
haps two very nearly allied species, Syngnathus rubes-
cens and S. tenuirostris. In the west of the Atlantic
it is unknown. Kroyer states that it occurs in the
Cattegat; and according to Wintiier he received two
young specimens from the northern entrance of the
Sound, oft' Ilornbaek. In Christiania Fjord it is com-
paratively plentiful, according to Collett; and the
Royal Museum has received through Mr. C. A. IIansson
several specimens, between 270 and 442 mm. in length,

from Dynekil and Stromstad Fjord. Thus it cannot
be considered rare on the north coast of Bohuslan; but
further south it has never been met with on the Swe-
dish coast.

The Great Pipefish generally lives, like other Lopho-
branchs, among the seaweed in comparatively shallow
water, even between the tide-marks. But now and
then we meet with the Syngnathi , and with this species
among others, at the surface, even far out at sea, where
they show a high degree of activity in their move-
ments, especially at night. During the Atlantic ex-
pedition of the corvette Josephine in 1869 I stood many
a night in the bows, and watched the wriggling motion,
in lines of phosphorescent light through the dark waves,
of Syngnathus pelagicus, a species which comes very
near the Great Pipefish, but is distinguished from it
in several respects, for example by a comparatively
larger head and shorter tail and by the much shorter
marsupium of the males. Thompson also tells us of
the Great Pipefish a: “A friend who has frequently
watched the movements of pipefishes in Belfast Bay
describes them as skimming along the surface of the
water, in the summer evenings especially, like a
slate thrown horizontally. — He lias seen them skip-
ping for 20 or 30 yards at a time, and also spring-
ing a foot high into the air.” The specimen which is
represented in our figure, was taken by a fisherman
from Bohuslan, Aniiersson by name, on “the Great
Fishing-bank S.W. of Bergen in 100 — 150 fathoms of
water;” but we have no information whether it was
taken at the bottom, or perhaps found at the surface.
In its daily life, according to Kroyer, the Great Pipe-
fish “swims slowly, with singular, stiff, angular, and as
it were contorted movements.” At these times, here
as in the Lophobranchs in general, it is the vibrating
dorsal fin that is the real instrument of locomotion, the
pectoral fins, which move in the same manner, seeming
rather to steer the course of the fish. The fish lies in
every possible position, with the head turned downwards
or upwards or forwards at will, as it glides on in quest
of its food*, which seems to consist principally of cru-
staceans0, usually of minute size, though “not unfre-
quently,” says Couch, “shrimps of comparatively no small
size are swallowed; and there have been found in the
stomach some so large as to raise our wonder how they

“ Nat. Hist. Irel., vol. IV, p. 239.

b Cf. Couch, Hist. Fish. Brit. Isl., vol. IV, p. 353.

c Cf. Olsson, Iakt. fislc. foda , Lunds Univ. Arsskr., VIII (1871), p. 10.

85

Scandinavian Fishes.

672

SCANDINAVIAN FISHES.

could have been made to pass between the jaws and
through the gullet,” this depending, of course, on the
distensive powers of the snout.

The spawning-season occurs in spring and summer.
Couch found a male with eggs in the marsupium in
April. On the 7th of May, 1869, at the west entrance
of the Channel (Lat. 47° 14' N., Long. 9° 9' W.), the
Josephine Expedition took several young specimens of
this species, swimming freely about at the surface.
These specimens were between 10 and 21 mm. long,
of light colour and transparent, but already marked
with transverse spots. In the smallest of them the
whole of the embryonic vertical fin was still persistent.
The largest ones had fully developed fins, except the
anal, of which not a trace was yet visible; and their
plates, as is usual in the fry, were tipped with spines.
We found the number of rings in front of the dorsal

fin to be 19. Malm saw in the possession of Mr.
G. Kolthoff a male 440 mm. long, with almost
fully developed embryos, that had been taken in
the neighbourhood of Skafto (Gullmar Fjord) on the
1st of August, 1877. In two males which were taken
in Stromstad Fjord by Mr. C. A. Hansson in Sep-
tember and October, 1888, the marsupium is quite
empty.

It was in this species that John Walcott first
discovered, either in 1784 or 1785 according to Yarrell,
that it is really the male who carries the impregnated
eggs during their further development, and not the fe-
male, as had previously been supposed ever since the
time of Aristotle. Walcott’s observation was first
published, however, by Yarrell in 1836, or five years
after Ekstrom had published his account of the Deep-
nosed Pipefish ( Syngnathus typhle).

THE LESSER PIPEFISH (si v. lilla tangsnallan")-
SYNGN ATH US ROSTELL ATUS.

Plate XXVIII, figs. 6—8.

Snout terete and shallow. Length of the head less than 13 % of that of the body and at most 20 % of that of
the tail , which is more than twice that of the trunk. Distance between the dorsal fin and the tip of the snout at
most 37 % of the length of the body, and the length of the base of this fin more than 1/3 of this distance and
also greater than the length of the head. Length of the marsupium of the males more than twice the length of

the head. Hind margin of the caudal fin rounded.

D. 33—39; A. 36; P. 9—11; C. (9) 10; Ann. 52—56 =
(15—17) + x.

Syn. Syngnathus pelagicus, Donovan, Brit. Fish., tab. LV1II (nec
Osbeck).

Syngnathus typhle, Malm (p. p.), Ofvers. Vet. Akad. Fork.
1852, p. 84.

Syngnathus rostellatus, Nilss., Skand. Fn., Fisk., p. 687 ;
Schleg., Pier. Nederl., Vissch., p. 179, tab. 17. fig. 2;
Ltkn, Vid. Meddel. Naturh. For. Kbhvn 1865, p. 222; Malm,
Gbgs, Boh. Fn., p. 595; Winth., Naturh. Tidskr. Kbhvn,
ser. 3, vol. XII, p. 53; Petersen, Vid. Meddel. Naturh. For.
Kbhvn 1884, p. 159; Lillj., Sv., Norg. Fisk., vol. HI,
p. 459.

Syngnathus acus (p. p.), Gthr, Cat. Brit. Mas., Fish., vol.
VIII, p. 157; Coll., Forh. Vid. Selsk. Christ. 1874, Tillasgsh.,
p. 200; ibid. 1879, No. 1, p. 101; Day, Fish. Gt. Brit.,
Irel., vol. II, p. 259, tab. CXLIV, fig. 2.

Syngnathus Dumerilii , Dum., Hist. Nat. Poiss. (Nouv. Suit, a
Buff.), torn. II, p. 556; Mor., Hist. Nat. Poiss. Fr., tom.
II, p. 49.

Ohs. As appears from the title of Plate XXVIII we originally
held the same opinion as Gunther and Day, that this little Pipefish
was merely a juvenile form, singular in certain respects, of the pre-
ceding species. The collections made by Hedenborg in the Bospho-
rus for the Royal Museum show, however, that Syngnathus rostellatus
is certainly distinct from the Mediterranean species with which it has
been identified by Gunther ( Syngn . hucculentus, Rathke; S. brevi-
rosti'is, Hempr. and Ehkenb.). The latter species has a shorter snout
and longer postorbital region etc., the length of the snout in all
Hedenborg’s specimens being less than 47 % of that of the head,
the postorbital length of the head equal to or at least more than 4/5
of the length of the snout, and the least depth of the snout ]/4 — 1/3
of its length, while the depth of the body at the beginning of the
tail is more than 1/5 (22 — 26 %) of the length of the base of the
dorsal fin. These two species cannot, therefore, be ranged as mutually
complementary forms of transition to Syngnathus acus. Furthermore,
in 1842 the Royal Museum received through Mr. Pontin, Member of
the Swedish Medical Council, a thing previously unknown, to the best
of our knowledge, namely a young specimen 155 mm. long of the
Atlantic Syngn. acus, taken “by a Dutch captain, probably in the
North Sea.” This specimen has 64 (20 + 44) rings on the body;

a Malm, 1. c.

b Sometimes 4, according to Lilljeborg and Nilsson.

LESSER PIPEFISH.

673

the length of the head is somewhat more than 13 % (13'2 %) of that
of the body, the length of the base of the dorsal fin less than l/3
(31 '6 %) of the distance between this fin and the tip of the snout or
only 88 % of the length of the head, and the temporal carinse — which
in Syngnathus rostellatus are never more than rudimentary ■ — are dis-
tinct, starting backwards from the supraorbital margins. Thus, as we
know the true juvenile form of the Great Pipefish, we are compelled to
retain the Lesser Pipefish as a distinct species, however nearly these
two species are allied.

The Lesser Pipefish attains a length of 16 cm."
The form of its body so closely resembles that of young
specimens of the preceding species that in this respect it
is difficult to state any constant distinction between them;
but the plate-armour confirms the rule already formu-
lated by Artedi, that within this genus we must look
for characteristic differences, first of all, in the number of
rings on the body. The Lesser Pipefish has only 15 — 17
rings on the trunk and 37 — 40 on the tail, or at most
57 in all, thus at least 6 less than the minimum num-
ber in the preceding species. The middle lateral carina
of the trunk is usually coherent in this species with the
upper lateral edge of the tail. It is stated, however,
that this character may also be found in the preceding
species, at least in young specimens thereof; and on the
other hand, the two carinae may be separated in this
species in the same manner as in the preceding one —
see the largest of the specimens figured in Plate XXVIII
(fig. 8, a; twice the natural size in fig. 8, b). The mar-
supium of the males occupies the first 23 — 26 rings
of the tail.

The length of the head measures about 10V2 — 1 2 1/2
% of that of the body, and its structure is otherwise
the same as in the preceding species; but behind the
eyes, on the top and on the temples in front of the
small occipital carina, it is evenly convex, without the
carinas which in the Great Pipefish run back on each
side from the upper orbital margin, or with only a rudi-
ment of a similar carina immediately behind each orbit.

The pectoral fins are comparatively larger than in
the Great Pipefish, and are inserted lower down the
sides. The length of their longest rays is greater than
the depth of the body at the beginning of the tail.

The dorsal fin is supported on 10 or 11 plates,
but sometimes partly occupies as many as 13. It be-
gins at a distance from the tip of the snout that mea-
sures 34 — 36* 6 % of the length of the body; and the
length of its base is about 15 — 1 2 1/3 % of the same

length, though, so far as we have been able to ascertain,
the variations are so restricted that it is always greater
than the length of the head. Its upper margin is more
regularly curved than in the Great Pipefish, with the
longest rays nearer the middle.

The anal fin is of insignificant size; but the caudal
fin is comparatively larger than in the preceding spe-
cies, its length being at least 3 7 2 % of that of the
body and 38 — 31 % of that of the head.

The three figures which are given in Plate XXVIII,
represent three individuals, one female (fig. 8), a young
male with undeveloped marsupium (fig. 7), and a spe-
cimen of the fry (fig. 6). These specimens were taken
at the beginning of July, 1887 in Koster Fjord (North-
ern Bohuslan) at the surface, where the depth of water
was about 100 fathoms. They were forwarded alive
by Mr. C. A. IIansson to the Royal Museum, where
they lived a few days longer. Their coloration was
above greenish, shading into brown; but the dorsal side
itself was lighter, shading into gray. The hover part
of the sides and the belly were silvery, the former with
a golden lustre in the male. The darkest colour thus
appeared like a longitudinal band from the sides of the
snout, across the eyes, on the postorbital part of the
head, and on the sides of the body just below the upper-
most lateral carina. But behind the dorsal fin, at the
transition to the tail, it was broken off short, and die
liberal band of the tail overlapped it below. The caudal
fin was brownish violet. The other fins were trans-
parent. The iris was golden. No traces of transverse
bands appeared on the body; but that this is not a,
constant character of the species, seems more than pro-
bable, and is easily explained, with the knowledge we
possess of the generic power of changing colour. In
some of the specimens belonging to the Royal Museum,
which have lain in spirits for more than forty years,
transverse bands of a darker colour appear between the
rings on the trunk, and in the males the strips be-
tween the marsupial plates are in some cases lighter,
in others darker. According to Nilsson too, “the co-
loration is above dark gray or brown with darker trans-
verse bands, below yellowish white; the caudal fin with
transverse streaks of brown.”

The Lesser Pipefish is fairly common in Bohuslan,
according to Malm the commonest species of the genus

a According to Collett. The largest specimen in the Royal Museum is a male 155 mm. long.

6 Moreau’s measurements of a female Syngnathus Dumerilii, which is probably identical in species with *S'. rostellatus, show, how-
ever, that this percentage may rise to 37.

674

SCANDINAVIAN FISHES.

except the Deep-nosed form. It was forwarded by
v. Duben to the Royal Museum in 1845 from Bergen.
Baron Gyllenstigrna found it in Sk elder Bay off Kul-
len, and handed over his specimens to Nilsson, who was
the first to describe this species. In the Sound it has
been met with by several Danish zoologists, among
others by Lutken off Hveen. Hansen has taken it in the
Cattegat off La? so. It goes southward in the Atlantic
to the Bay of Biscay, according to Moreau, but is rare
on the French coast.

The habits of the Lesser Pipefish probably resemble
in the most essential respects those of its larger con-
gener;- but its comparatively larger fins are suggestive
of a more active life, and its insignificant size must

compel it to seek its food among the most minute
marine animals. Its spawning-season occurs in spring
and summer; from May to the middle of June Malm
found eggs and young in the marsupium of the males.

In order finally to express, with the greatest bre-
vity, the difference of form between this species and
its nearest two relatives in the Atlantic region, we give
below the most important proportions in this respect
of three specimens of equal size, a male of S. rostel-
latus from Bohuslan, a female of S. acus from the North
Sea, and a male of S. pelagicus from the Sargasso Sea
(Lat. 32° N., Long. 43° W.), all three 155 mm. long
from the tip of the snout to the hind margin of the
caudal fin.

Syngnatlius

rnstellatus.

acus.

pelagicus.

Number of rings on the trunk

16

20

17

,, ,, ,, ,, ,, tail

40

44

32

Length of the head in % of that of the body

10.0

13.2

13.9

Distance between the dorsal fin and the tip of the snout in % of the length of the bodv

34. o

36.8

38.7

„ anal „ „ „ „ „ „ „ ., -

35.5

39.o

43.2

Length of the trunk behind the gill-covers in % of the length of the tail. _

40. o

42.3

51.7

., base of the dorsal fin in % of the distance between the dorsal fin and the tip of the snout..

35.4

31.6

33.3

,, ., .. ,, ., ., .. „ ., ,, ,, ,, ., ,, vent ,, ., ,, ,, ,, „

34.5

29.7

29.8

,, ,, ,, ,, ,, .. ,, length of the head

121.8

87.8

93.o

., „ „ middle rays of the caudal fin in % of the length of the head

31.4

24.4

27.9

THE DEEP-NOSED PIPEFISH (sw. KANTNALEN OR TANGSNALLAN).

STAGNATE US TYPHLE.

Plate XXIX, fig. 1.

Snout laterally compressed and comparatively deep. Length of the head more than 15 % of that of the body and
at least 1 4 of that of the tail , which in the males is generally more , in the females less, than twice the length of
the trunk behind the gill-covers. Distance between the dorsal fin and the tip of the snout at least 39f2 % of the
length of the body , and the length of the base of this fin less than 1/3 but more than V4 of this distance , and
also less than the length of the head. Hind margin of the caudal fin more or less obtusely pointed.

R. hr. 2; D. 35 — 38«; A. 3; P. 13 — 15; C. (8) 10; Vert.

55; Ann. 52—57 = (17 — 19) + x.

Syn. Syngnatlius corpore medio lieptagono, cauda piimata, Art.,
Ichthyol. , Spec. Pise., p. 2; Lin., Fn. Suec.,e d. I. p. 126,
No. 335.

Syngnatlius Typlile , Lin. (p. p.), Syst. nat., ed. X, tom. I,
p. 336 (»Lamina3 corporis trunci 18, cauda 36»); Donovan,
Brit. Fish., tab. LVI; Flmng, Brit. Anim., p. 175; Jen.,
Man. Brit. Vert., p. 485; Yarr., Hist. Brit. Fish., ed. 1,
vol. II, p. 332; Fr., Vet.-Akad. Handl. 1837, p. 28, tab. Ill,

fig. 2; Bonap. ( Siphostoma , ex Raf.), Cat. Met. Peso. Europ.,
p. 89; Kr., Damn. Fiske, vol. 3, p. 673; Nilss. ( Syngna -
thus), Skancl. Fn., Fisk., p. 689; Sundev., Stkblms L. Hush.
Sallsk. Handl., H. VI, pp. 83 et 165; Mgrn ( Siphostoma ),
Fail. Fiskfn. (disp. Helsingf.), p. 69; Lindstr. ( Syngnatlius ),
Gotl. L. Hush. Sallsk. Arsb. 1866, p. 24 (sep.); Gthr
( Siphonostoma ), Cat. Brit. Mus., Fish., vol. VIII, p. 154;
Coll., Forh, Vid. Selsk. Christ. 1874, Tillaegsh., p. 199,
ibid. 1879, No. 1, p. 100; Winth., Naturh. Tidskr. Kbhvn,
ser. 3, vol. XII, p. 52; Bncke ( Syngnatlius ), Fisch., Fischer.,
Fiscliz. 0. W. Preuss., p. 189; ILcke (Siphonostoma), Arch.

a 34—40, according to Moreau.
b 38—46, „ „ Day.

DEEP-NOSED PIPEFISH.

675

f. Natnrg., Jahrg. 46, Bd. I (1880), p. 321; Mgr., Hist.
Nat. Poiss. Fr., tom. II, p. 55; Mela, Vert. Fenn., p. 360,
tab. X; Day, Fish. Gt. Brit ., Irel., vol. II, p. 257, tab.
CXLIV, fig. 3; Mob., Hoke ( Siphonostcnnum ), Fisch. Osts.,
p. 102; Coll. ( Syngnathus ), N. Mag. Naturv. Christ., Bd. 29
(1884), p. 113; Lillj. ( Biphonostoma ), Sverg., Norg. Fisk .,
vol. Ill, p. 439.

Byngnatlius acas, Ekstr., Vet. Akad Handl. 1831, p. 271, tab.
II, figg. 1 et 2 ; Schagerstr., Physiogr. Sallsk. Tidskr., p.
314; Malm ( Siphonostoma ), Gbgs , Boh. Fn., p. 592.

The Deep-nosed Pipefish attains a length of 33
cm., according to Yarreel. The largest Scandinavian
specimen in the possession of the Royal Museum is a
female 27 cm. long (from Norway, through Marklin);
and according to Ekstrom the size of the species never
exceeds 22x/4 cm. in the island-belt of Sodermanland.
In form of body it is much the same as the Great
Pipefish, the only difference being that it generally has
a longer trunk and shorter tail. It thus represents in
general the sexual characters that distinguish the fe-
males. As a rule too, the depth of the body is some-
what less; at the beginning of the tail it seldom (and
only in the males and young specimens) exceeds 2x/2
% of the length of the body".

The plates of the body essentially correspond in
form and arrangement to those of the two preceding
species. As we have mentioned above, however, the
foremost inferior plates, the lowest covering- plates of
the shoulder-girdle, are more loosely attached to each
other at the middle of the ventral side6, and the first
ventral plate is nearly always wanting*. The first of
the ventral plates that are present (corresponding to
the second in the Great Pipefish), is also, as a rule,
considerably reduced, so that a longitudinal, narrow
strip, covered only with skin, is left behind the junc-
ture on the ventral side of the plates belonging to the
shoulder-girdle- Thus, as the last (and sometimes the
penultimate) ring on the trunk is also without ventral
plate, here as in the Great Pipefish, we find only 17
(sometimes only 16) ventral plates, while the number
of plates in the lower lateral row of the trunk is usu-
ally 18. In the cases where the last-mentioned num-
ber is 19, there are only 18 ventral plates; when it is
17, we find, as a rule, only 16 of the latter. The so-
called occipital plates are only slightly raised, and the

anterior one is still smaller than in the Great Pipefish.
The middle lateral carina of the trunk (the middle la-
teral row of plates) as a rule forms a regular continua-
tion of the carina curving downward in front from
the upper lateral margin of the tail, and in this re-
spect the Deep-nosed Pipefish thus represents the early
stages of the two preceding forms. But it is not un-
usual to find between the said carinee the same break
here as in the Great Pipefish, the middle lateral carina
of the trunk ending in the last ring of the trunk (see,
for example, Plate XXIX, fig. 1), and the correspond-
ing lateral carina of the tail beginning obliquely above
this point, in the first caudal ring. We have never
found the last-mentioned carina intrude within the last
ring of the trunk. The marsupium of the males ex-
tends, when fully developed, along 24 caudal rings,
and attains a length more than double that of the head.

The length of the head, as a rule greatest in young
specimens, measures about 18 — 15 % of that of the
body*7. The contour of the postorbital part of the head
passes evenly into that of the forepart of the trunk.
The occipital carina is not very prominent on the flat
occiput, which slopes evenly and gradually in front into
the slightly concave forehead, but is sharply marked off
from the steeply sloping temples, this sharp break being
formed on each side by the temporal carina running
back from the upper orbital margin. The superior
profile of the forehead and snout is also even, and the
orbital margin rises hardly perceptibly above it. The
praefrontal knob is elongated in an horizontal direction.
The carinat of the snout are the same as in the Great
Pipefish. The distinguishing character of the head of
this species lies in the deep and strongly compressed
form of the snout, when the mouth is closed, and also
in the fact that at the ascending tip of the snout the
lower jaw projects almost as far as the upper. When
the month is wide open, the snout assumes the terete
shape of a straight, cylindrical tube, only a little dis-
tended in front.

The length of the snout measures as a rule 60 — 63
% of that of the head: but in young specimens (be-
tween 6 and 7 cm. long) and exceptionally in adult
ones this percentage may sink to 57 or even to 56.

a 2'8 is the highest percentage we have found in this relation, and this in a male 247 mm. long.

b Now and then these plates are so loosely joined that they are not even contiguous, being united merely by the skin.

c It is present, however, in one of our largest specimens, from Norway.

d According to A. H. Malm ( Om den brednabbade kantndlens utveckling ocli fortplantning , disp. Lund 1874, p. 4) the length of the

head varies in the fry (12 — 43 mm. long) between about 1/6 and */- of that of the body.

676

SCANDINAVIAN FISHES.

Its least- depth, just behind the upward curve of the
tip, is as a rule about 1/5, but sometimes no more
than 15 % of its length. The length of the lower jaw,
which is as a rule about 1/i (varying, however, in dif-
ferent individuals between 30 and 21 %) of that of the
snout, measures in adult specimens, at least after they
have attained a size of 12 cm., distinctly more (gene-
rally V3 more) than the diameter of the eyes", but in
young specimens (6 or 7 cm. long) is about equal to
this diameter*. The eyes are also comparatively smaller
than in the Great Pipefish. In young specimens (6 or
7 cm. long) their diameter is about 74 or even 28 %
of the length of the snout, but with increasing age this
percentage sinks to about 15 or even 14. In the said
young specimens the postorbital length of the head
measures about 38 — 36 % of its entire length, or about
2/3 (at least 64 %) of the length of the snout; but in
older specimens this proportion sinks to about 30 or
29 % of the length of the head, or 50 — 46 % of that
of the snout.

The gill-openings are comparatively somewhat
larger than in the Great Pipefish, their length being
about 2/3 of the diameter of the eyes; but they are
otherwise of the same form and position.

The pectoral fins are of the same obtuse shape as
in the Great Pipefish, or even broader, and generally
contain a greater number of rays.

The dorsal fin is longer and lower than in the
Great Pipefish, in the former respect coming nearer the
Lesser Pipefish. It is also set comparatively further
back, this being due to the shortness of the tail. Here,
however, we find a distinct sexual difference in the
Deep-nosed Pipefish: in the females the trunk is com-
paratively longer than in the males, and the dorsal and
anal fins are consequently situated further back. The
two preceding species represent in this respect that- di-
rection of development which has been fixed by the
preponderant influence of the male characters. In the
Deep-nosed Pipefish the distance between the dorsal
fin and the tip of the snout, to the best of our know-
ledge, is never less than 3972 % of the length of the body,
in the males hardly more than 42 %, in the females
about 44 %, and in young specimens sometimes 47 %.
It generally begins on the last ring of the trunk (the
anal ring), sometimes on the penultimate one. The

base of the fin (12 — 1373 % of the length of the body)
occupies in most- adult specimens 10 rings, in some 9
or partially 11, and in young specimens 8. Its height,
which is only slightly greater at the middle than at
either end, is generally equal to the length of the
lower jaw.

In the Deep-nosed Pipefish as in the Great Pipe-
fish, we have not been able to find more than 3 rays
in the anal fin. According to Krdyer and Day, how-
ever, it may contain 4 rays. It is set- at a distance
from the tip of the snout that measures in the males
about 40V2 — 42Va in the females about 44 72 — 45 7a

% , and in young specimens as much as 4872 % of the
length of the body.

The caudal fin, which is distinguished by the
bluntly pointed form of the hind margin, measures as
a rule about 37 2 01> 4 % of the length of the body,
and thus corresponds most nearly in this respect to
the caudal fin of the Lesser Pipefish. But in con-
sequence of the greater length of the head in the
Deep-nosed Pipefish, the length of the caudal fin is
only about x/5 of that of the head, though it- varies
between 18 and 25 % thereof. In most cases it con-
sists of 10 rays; in one single instance we found 8.

All the fin-rays are of the same type as in the
two preceding species; but in the caudal fin they are
sometimes articulated.

Of the coloration of the Deep-nosed Pipefish Fries
remarks: “In both localities, both in the Baltic and
the Cat-tegat, two colour-varieties occur, the first green
with yellow spots and with the belly shading decidedly
into brassy yellow, the second olive-brown strewn with
numerous whitish dots and spots and with whitish belly.
These two varieties are not constant, however; we find
a series of intermediate forms between them. They
do not stand in any fixed relation either to age or
sex.” The relation of these two colour-varieties to
each other has been explained in recent times by
Heincke0 as follows:

“The power Avhich the Syngnatlii possess of ad-
apting their colour to their environments, is the most
perfect instance of the kind that we know among
fishes. If we place some Deep-nosed Pipefishes in a
large aquarium, together with a quantity of seaweed
( Zostera marina) such as that, which grows in the

“ In a female 255 mm. long, from Bohuslan, the longitudinal diameter of the eyes is only 55 % of the length of the lower jaw.
b In the fry the lower jaw is, of course, considerably shorter — cf. A. H. Malm’s figures (1. c.).
c Schriften des Naturwissenschaftlichen Vereins fur Schleswig-Holstein, Band I, p. 257.

DEEP-NOSED PIPEFISH.

677

native haunts of the fishes, after some time we may
make an extremely interesting observation. The leaves
of the seaweed have partly risen vertically or obliquely
through the water, and stand motionless, slowly sway-
ing to and fro if the aquarium be slightly shaken.
Among them, motionless or swaying with the leaves,
the slender Pipefishes have chosen their positions, and
we can only just see how the gill-covers expand and
contract, or how the perfectly transparent dorsal fin
ceaselessly continues its vibrating and undulating mo-
tion. The colour of the fishes, often down to the most
delicate shades, is exactly like that of the seaweed.
Often we imagine that we are gazing at a blade of
seaweed, and only on closer inspection do Ave discover
that it is a Pipefish, and vice versa. The same light or
dark, green or yelloAvish green hue covers both sea-
Aveed and Pipefishes; the singular, lighter tint of green
Avhich the former sometimes shoAvs in isolated patches,
is faithfully reproduced in the latter. Among the
green, still living seaAveed there lie here and there a
number of partly or entirely dead leaves, in all shades
of colour from green to dirty-broAvn and broAvnish
black. At these spots the Pipefishes assume a different
hue, their colour passing gradually, according to their
different surroundings, into broAvn or broAvnish black,
until, Avhether their position be vertical or horizontal,
they are scarcely to be distinguished from a dead blade
of seaAveed. — The time occupied by this adaptation of
colour varies considerably. Mechanical irritation of the
skin and psychical irritation seem distinctly to ac-
celerate the process. Large Pipefishes, Avhich Avere
quite dark Avhen taken in the hand, at once assume
a pale green colour, all the time struggling violently
to get loose; and if they are transferred to a vessel
Avith darker bottom, their dark coloration returns pretty
soon. Full-groAvn Pipefishes, if left undisturbed, seem,
to require at most an hour to change colour; but in
young specimens, about 21/ 2 cm. long, Avhich have just
emerged from the marsupium, the change is effected
Avith extreme rapidity, in a fraction of a minute. —
The inner layers of the skin contain greenish yelloAV
chromatophores, the outer layers darker ones, black
Avhen contracted, broAvn when expanded. The contrac-
tion of the latter, which affords a beautiful sight under
the microscope, may take place with astonishing ra-
pidity. The handsome, stellate figures of the chromato-
phores, which at many spots appear to lie united by
their projections, distinctly shrink until they form small

dots, Avith one or tAvo grains of pigment detached from
the central mass of colour. I have not been able to
observe the contraction and expansion of the greenish
yelloAV chromatophores, Avhich Avould thus seem to per-
form their alterations much more slowly than the dark
chromatophores. The light cells never expand into
figures of so varied form as the black.”

Our figure (Plate XXIX, fig. 1) represents a spe-
cimen of the green variety.

The geographical range of the Deep-nosed Pipe-
fish extends along the Avhole west coast of Europe,
from Norwegian Finmark to Gibraltar. Tromso Mu-
seum received it in 1881, according to Collett, from
Belstad Fjord, a little south of Tromso; and according
to Lilljeborg Upsala Museum has received it through
Professor Th. Fries from Vadso. On the south coast
of Scandinavia it is very common, and it penetrates into
the Baltic at least to the south of the Gulf of Bothnia
and almost to the head of the Gulf of Finland (Mela).
In the island-belt of Stockholm, according to Sunde-
yall, it is plentiful Avherever Fucus vesiculosus groAvs
luxuriantly. In the island-belt of Morko it is taken
in quantities, according to Ekstrom, Avhen the seine is
drawn for other fish, except during May and June, Avhen
it seldom, if ever, visits the shores or the shalloAvs.
On the coast of Gothland it is common among Zoster a
marina , according to Lindstrom. It is quite as com-
mon, if not more so, in the south and Avest of the
Baltic, on the Avest coast of SAveden, on the Danish
coast, and on the south coast of Norway. Further
north and further south it grows less common, and is
said to be rare in the Mediterranean, Avhere its place
is taken to a great extent by another species, Syn-
gnathus Bondeletii, Avith deeper snout (least depth of
the snout greater than the depth of the body at the
beginning of the tail), shorter dorsal fin (length of the
base less than 1 1 1/2 % of that of the body), and more
rings on the trunk (20 or 21), or thus related in the
most essential respects to our Deep-nosed Pipefish in
the same way as the Great Pipefish to the Lesser
species.

Except in the spaAvning-season the Deep-nosed Pipe-
fish generally keeps close to the shore, in a feAv metres
of Avater or even less, among grass-Avrack {Zoster a
marina ) and bladder-wrack {Fucus vesiculosus ), Avhere
it has shrimps and their young, small crustaceans,
minute mollusks, and Avorms as its usual comrades and
principal diet. Young fishes and tiny Gobies also fall a

678

SCANDINAVIAN FISHES.

prey to it. Though the mouth is small, it is capable
of quite considerable extension, as we have already
remarked of the two preceding species. There is no
doubt, however, that the Deep-nosed Pipefish under-
takes roving expeditions at the surface of deeper wa-
ter: one of the BohusDn fishermen who have collected
marine animals for the Royal Museum, has handed
over to Professor S. Loven a specimen taken “in the
North Sea, off Bergen.”

Ekstrom’s experience in the island-belt of Morko
was that, “though the Deep-nosed Pipefish haunts the
shore, in shallow water, it still repairs to deep water
during the warmer part of the year". In autumn,
winter, and the early part of spring it is met with in
the inlets, in water of little depth. From the end of
April to the beginning of November it is less plenti-
ful in these localities, and during the whole of May
and a part of June it is taken extremely seldom and
invariably in deep water. In temperament it seems to
be sluggish and not very timid. Its movements in the
water are stiff, and betoken but little activity.”

It was the history of the Deep-nosed Pipefish that
gave Ekstrom the clue to the explanation of the hitherto
obscure sexual relations of all the Lophobranchs. “It
has long been known,” he writes in 1831, “that the
Deep-nosed Pipefish carries the roe as well as the new-
hatched young under its tail. Even Aristotle5 re-
marked in this species the peculiarity, otherwise rare
among fishes, that the eggs seem to pass, not as in
most fishes through the vent, but through a slit in the
body. He did not pay any attention, however, to the
organ that envelops the eggs and the young during
their development, but supposed that the eggs were
metamorphosed within the abdominal cavity, and that
the belly itself was opened by the development of the
eggs, at the slit which appears behind the vent. He
thus seems to have been the first to promulgate the
long persistent theory that this slit did not belong to
a distinct organ, but was due to the bursting of the

belly by the growth of the eggs. /Elian c maintained
the same opinion. Pliny quotes the very words of
Aristotle, with the only alteration that the bursting
of the belly was due to the number of the eggs.
Rondelet'7 Avas the first to remark that the eggs are
contained in a special organ.” Rondelet further stated
that the females are distinguished by the possession of
this organ from the males, an opinion Avhich survived
until Ekstrom published his observations at Morko.
“The spawning- season of the Deep-nosed Pipefish,” says
Ekstrom, “occurs here in the month of May, Avhen the
male is forced to seek his female, or vice versa; and
as a regular copulation between the sexes is necessary,
the spawning takes a longer time than among fishes
in general6. At the end of April the females desert
the shore and the shalloAvs to begin their spawning in
deeper water. As the spawning season approaches, the
foliate lids that close the opening of the marsupium of
the male, become tumid, and the marsupium is gradu-
ally filled with a white, clear, and thick mucus, Avhich
serves as a bed for the eggs. The eggs lie imbedded
in this mucus, which decreases in quantity according
to the growth of the young, until little or none of it
remains by the time that the fry are large enough to
SAvim and independently to move through the Avater.
— The eggs, Avhich lie in regular, moniliform roAvs, are
large' in proportion to the fish and, Avhen they are
deposited, yelloAv, but gradually turn Avhite and be-
come transparent Avitli a fine, dark yelloAv point, the
rudiment of the embryo. — At the middle of June the
fish gradually return to the shore from their spawning-
places. At the end of July, in some cases, the young
are so developed that they can leave the marsupium
and folloAv the movements of their father. In other
cases the roe has been only just deposited. In a fe-
male about 2 dm. long I counted 240 eggs in the
ovaries. In the marsupium of the male I never found
a quarter of this number6’. Many of the eggs must,
therefore, be lost during copulation, and Ave must thus

“ KR0YER lias remarked that at this season the shrimps also retire to deep water.

b Hist, anim., lib. VI, cap. XIII. This remark applies rather to the more common Mediterranean species Syngnathus Rondeletii.
c Lib. II, cap. XIII.
d De Pise., lib. VIII, p. 229.

e More recently it has been discovered that the copulation must be repeated several times, for a female never discharges all her eggs
at once into the marsupium of the male, the anterior end of which is open during copulation, and is penetrated by the oviduct, now pro-
jecting to a length of several millimetres. Cf. Lafont, Actes de la Soc. Linn, de Bordeaux, 1871, t. 28, p. 251, Heincke, Arch. f. Naturg.,
1. c., p. 330, and Lilljeborg, 1. c., p. 438.
f About 2 mm. in diameter.

'J In the marsupia of the largest males from Kiel Bay, hoAvever, Heincke found 150 — 200 eggs.

PIPEFISHES.

refrain from inferring the fecundity of this fish from
the number of eggs in the female. To judge by the
numbers among these islands, the males are so few in
comparison to the females that hardly one of the former
can be found among ten of the latter.”

Ekstrom’s observation that during the spawning
the Deep-nosed Pipefish retires to deeper water, at least
two fathoms or more in depth, has been corro-
borated in Bohuslan by A. H. Malm (1. c., p. 18),
who supposes that the spawning is most general
there from the middle of June to the beginning of
August.

Besides the protective likeness which the Deep-
nosed Pipefish attains by means of its changes of colour,
Heinckk has remarked another, which is given to the
males while their marsupium is distended with eggs or
young. The marsupium with its long slit at the middle
then presents a striking resemblance to the spathe of
Zostera, and still further increases the difficulty of

679

distinguishing these fishes from the flowering grass-
wrack.

The young stay in the marsupium or take refuge
therein (see above) until they have attained a length of
about 25 mm. As A. II. Malm and Lilljeborg have
remarked, they are destitute of the embryonic vertical
fin, which is present, however, in the larvte of the
Great Pipefish. They grow rapidly, according to Heincke,
attaining a length of at least one decimetre by the end
of the first year; and the marsupium of the males may
sometimes be fully developed even at an earlier period.
The number of the caudal rings increases during growth
from 32 to 37 or 38, but even in specimens about 6
cm. long the number of rings on the trunk is complete
or at least 17.

The Deep-nosed Pipe-fish is of no greater econo-
mical value than the rest of the Lophobranchs. It may
be employed, however, as food for swine or as bait
for Cod and Bullheads.

Genus NEROPHIS.

Pectoral and anal fins wanting. The males carry tl

side of

Fries bestowed upon this genus, which he charac-
terized, but regarded merely as a subdivision of the
genus Syngnathus, the Swedish name of Hafsnalar (Sea-
Needles)". Rafinesque had indeed established a ge-
nus Nerophisb in 1810, but Kaup was the first (in
1 853 c) to give this name a fixed application. Dume-
rii/ treated this division as a distinct subfamily, Ne-
rophini, which he distributed among three genera.

As we have mentioned above, this group is dis-
tinguished from the preceding Syngnathince not only
by the constant absence of the anal fin and the dis-
appearance of the pectoral fins and, in most cases, of
the caudal fin, but also by the weaker covering of
plates and the less distinct car i rue on the body. The
skin that covers the plates, on the other hand, is more
strongly developed and sometimes elevated at the middle
of the back and the belly in the form of a dermal
carina or even (in Protocampus) of an embryonic, but

“ Syngnatlii opliidii as opposed to the preceding group, which
6 Indice d’ittiologia Siciliana. We have not seen this rare
a minor sea-god) and orptg, snake.

c Arch. Naturg., Jahrg. XIX, Bd. I, p. 234; Cat. Lophobr. 1
,l Hist. Nat. Poiss. (Suit, a Buff.), torn. II, p. 600.

impregnated eggs in a layer of mucus on the ventral
e trunk.

persistent vertical fin. Another essential difference is
that the upper row of plates on the trunk (the dorsal
row) advances on each side of the body even along the
tail, and that the middle lateral row on the trunk passes
in the same manner into the lower caudal row, the lower
lateral rows of the trunk and its ventral row being
thus unrepresented in the plate-armour of the tail.

Fries has remarked the significant sexual distinc-
tions that prevail in the species of this group, partly
in the position of the vent (comparatively further back
in the females), partly in the form of the trunk (deeper
and narrower, with more distinct dorsal and ventral
carinas, in the females). He has also pointed out that
the most trustworthy specific characters must be sought
in the position of the dorsal fin, the number of rings
on the trunk, and the length of the snout in proportion
to its depth and to the length of the head. Guided by
these observations, he arrived at a. safe stand-point for

he called Syngnathi marsupiales.

work. The name is formed from the Greek vrjQog, wet (or NcQEtg,
7ish. Brit. Mus., p. 65.

86

Scandinavian Fishes.

680

SCANDINAVIAN FISHES.

the elucidation of the confusion that had previously
involved the synonymy and definition of these species.
He divided them into two groups:

* Caudal fin rudimentary, with 4 or 5 (6) short rays; greater
part of the dorsal fin situated in front of the perpendicular
from the vent.

** Caudal fin entirely wanting; greater part of the dorsal fin
situated behind the perpendicular from the vent.

The former group received in 1856 of Dumeril
the Elder" the generic name of Pterurusb. As this name
had already been employed in 1810 by Rafinesque
among the Eels, it was exchanged in 1870 by Dumeeil
the Younger0 for Entelurus'1. As a well-marked stage
of development in the gradual disappearance of the
caudal fin, a genus of this nature might well claim a
place within the European fauna, where the character

derived from the rudimentary remnant of the caudal
fin coincides with the above-mentioned character drawn
from the position of the dorsal fin. But Kaup has
described a species, Nerophis Heckelii , which shows
(provided that the description is correct) that the rem-
nant of the caudal fin may be found in conjunction
with the opposite position of the dorsal fin; and the
developmental character loses its validity from a syste-
matic point of view, as it is not based on any actual
divergency.

The genus Nerophis contains 7 known species from
the Atlantic, the Baltic, the Mediterranean, the Black
Sea, and the Indian Ocean. They lead the same life
as the species of the preceding genus, but are still
more whip-like in form, and employ the tail as a more
or less developed prehensile organ.

THE iEQUOREAL PIPEFISH ( SW. STORA HAFSNALEN).

NEROPHIS 2EQUOREUS.

Fig. 173 and Plate XXIX, fig. 2.

Tip of the tail generally furnished with a rudimentary and fragile fin , containing 4 — 6 rays. Vent situated
below the posterior part of the dorsal fin , the distance between which and the tip of the snout is less than 85 %
(81 %?) of that between the vent and the same point and less than 5 times the length of the head.

Fig. 173. Head of a Nerophis cequoreus 2 85 min. long. Natural size. F. v. Wright in B. Fries.

D. (37) 40 — 44 (46); (J. (4—60; Ann. 90 — 95 (100) =
(29— 31)4-

Syn. Acus nostras cauda serpentina, Sibb., Scot. III., pt. 2, tom. 2,
p. 24, tab. 19.

Soe-Naal , No. 2, Str6m, Sondm. Beskr., pt. 1, p. 312.
Syngnathus cequoreus , Lin., Syst. Nat., ed. X, tom. I, p. 337;
Mont., Mem. Wern. Nat. Hist. Soc., vol. I, p. 85, tab. IV,
fig. 1; Pall., Zoogr. Boss. As., tom. Ill, p. 121; Swains.
(Acus) Nat. Hist. Fish. Amph. cett., vol. II, p. 333 ; Jen.
(Syngnathus) Man. Brit. Vert., p. 486 (9); Fr., Vet. Akad.
Handl. 1837, p. 35, tab. Ill, fig. 3; Kr., Damn. Fiske,
vol. 3, p. 705; Kp (Nerophis), Cat. Loph. Fish. Brit. Mus.,
p. 66; Nilss. (Scyphius, ex Risso), Skand. Fn ., Fislc.,
p. 692; Gthr (Nerophis), Cat. Brit. Mus., Fish., vol. VIII,

p. 191; Dum. (Entelurus) Hist. Nat. Poiss., tom. II, p. 605;
(9); Coll. (Nerophis), Forh. Vid. Selslc. Chrnia 1874, Til-
laegsh., p. 202 ; N. Mag. Naturv. Chrnia, Bd. 29, p. 114; Malm,
Gbgs, Boh. Fn., p. 596; Winth., Nat. Tidskr. Kbhvn, ser. 3,
vol. XII, p. 53; Mor. (Entelurus), Hist. Nat. Poiss., Fr.,
tom. II, p. 62; Day (Nerophis), Fish. Gt. Brit., Irel ., vol. II,
p. 261, tab. CXL1V, fig. 4; Storm, N. Vid. Selsk. Skr.
Trondhj. 1883, p. 42; Lillj., Sv.,Norg. Fisk., vol. Ill, p. 465.

Syngnathus anguineus , Jen., Cat. Brit. Vert., p. 30 (cf);
Yarr., Brit. Fish., ed. 2, vol. II, p. 445; Kp (Nerophis),
1. c., p. 65; Dum. (Entelurus), 1. c., p. 606; Mor., 1. c.,
p. 63.

Syngnathus ophidian , Bl. (p. p., nee Lin.), Fisch. Deutschl.,
pt. Ill, p. 115, tab. XCI, fig. 3; Jen., Man. Brit. Vert.,
p. 487; Yarr., Brit. Fish., ed. 1, vol. II, p. 338.

° Ichthyologie Analytique, p. 169.
b Tcxsqov, fin and ovga, tail.

0 Hist. Nat. Poiss., tom. II, p. 605.
d ivvElyg, perfect.
e Sometimes 7, according to Kroyer.

iEQUOREAL PIPEFISH.

681

The iEquoreal Pipefish is the largest of all our
Syngncitliince. It attains a length of at least 6 dm."
The females are largest, their usual size in Scandinavia,
according to Fries, being 45 — 55 cm., while the length
of the males is usually between 32 and 40 cm. The
form of the trunk also shows a considerable sexual dif-
ference. It generally displays more lateral compression
than in the preceding forms, but most in the females,
this being due to the more or less advanced develop-
ment of the dermal carina that runs along the dorsal
and ventral margins of the trunk, but which is want-
ing in the males and young specimens. The tail is
more slender and more terete (its section more circular)
than in. the preceding forms. A section of the trunk
shows an oval, octagonal form, in the females pointed
at both ends; the section of the tail is a rounded square.
The greatest breadth of the trunk measures in the fe-
males about 70 %, in the males about 90 % of its
greatest depth; the young specimens stand about mid-
way between these two extremes. The greatest depth
of the body in the males is about 2 7 2 %, in the females
about 23/4 % of the length thereof. At the beginning
of the tail the depth of the body measures in the fe-
males about 60 %, in the males about 70 % of the
greatest depth, or in the former about 11%, in the
latter about 12 or 13 % of the length of the base of
the dorsal fin.

In addition to the above-mentioned generic cha-
racter that the caudal rows of plates form a continuation
of the two uppermost rows of the trunk on each side
of the body, this species and the following one are
especially remarkable for the great number of rings on
the body, a number which, at least in this species, may
rise to 100. The last caudal rings are so small, how-
ever, that they can scarcely be counted with the naked
eye. The so-called occipital plates are without Carinas,
but are as usual grooved; the anterior is about half
the size of the posterior.

The head is most like that of the Lesser Pipefish, but
is comparatively smaller, with smaller eyes and with less
prominent carina? both on the snout and especially on
the gill-cover, where scarcely a trace of a middle carina
can be found. The snout is terete and straight, ascend-

ing only slightly. The forehead is hollowed into a long,
shallow concavity. The length of the head varies in
adult specimens between about 7l/2 and 9 % (7.6 — 8.8
%, according to our measurements) of that of the body.
In young specimens (between 12 and 17 cm. long) the
length of the head measures about 27 % of the distance
between the dorsal fin and the tip of the snout, in old
specimens 22 — 2472 % thereof. The length of the snout
varies between about 44 and 54 % of that of the head,
and generally begins to exceed V2 of the latter in spe-
cimens 37 cm. long. The postorbital length of the head
varies between about 43 and 38 % of its entire length,
and this percentage generally begins to sink below 40
in specimens 37 cm. long. The longitudinal diameter
of the eyes may measure about 14 % of the length of the
head even in specimens 35 cm. long; in specimens 44 cm.
long it measures about 7m °f the same. The least depth
of the snout varies in different individuals and according
to age between about 29 and 18 % of its own length.

The dorsal fin is of fairly uniform height, this
being always distinctly greater than the depth of the
body at the vent. It begins at a distance from the
tip of the snout that in the males does not exceed
35 % (3l72 — 3472 %•> according to our measurements)
of the length of the body, but in the females (at least
the older ones) does not fall below 37 % thereof6. The
length of its base increases with age from about 12 %
to 1 4 1/3 % of that of the body. The vent lies below
the posterior part of the fin, at a distance from the
tip of the snout that in the males measures about 39 72
— 4372 % (39'6 — 43‘6 %, according to our measurements
of specimens between 12 and 37 cm. long), in the older
females 47 — 49 % of the length of the body. One re-
sult of this is that the length of the trunk behind the
gill-covers, in young specimens less than 17 cm. long
(we have never examined any young females), measures
only slightly more than 1j2 (about 51 %), in the older
males about 3/5 (59 — 64 %), and in the older females
about 74 (74 — 78 %) of the length of the tail. In these
differences, depending on age and sex, lies the most im-
portant character that has been employed as a specific
distinction between Nerophis cequoreus (?) and Ner.
anguineus (cf).

a From Ireland we hear of still larger specimens. “Last winter,’’ writes Blake-Knox in August, 1866 (Zoologist, vol. XXIV, p. 508)^
“I met with an immense fish of this kind: seeing a boy “whacking” a donkey with a gutta-percha stick, as I thought, I asked him where
he got it. “It is only a stalk of a snot (seaweed), sir; see:” and I did see a fine asquoreal, 3 feet 5 inches long. Of its toughness you
may judge. Is not uncommonly taken in the baskets with whelks and crabs.”

b Kroyer, however, mentions a female (sign. B among the specimens mentioned by him) in which this sexual character is absent.

682

SCANDINAVIAN FISHES.

The caudal fin is extremely small, pointed, and so
fragile that it now and then disappears with age.

In the living fish, according to Fries, the colora-
tion is of a handsome flame-yellow or brownish yellow.
Straight across the sides of the trunk and some way
along the tail the body is marked with somewhat un-
dulating, parallel, whitish bands, framed with brown.
These bands are arranged in such a manner that they
lie alternately at the middle of a ring of plates, alter-
nately on the diamond-shaped space between two rings,
the number of the bands thus being double that of the
plates. The dorsal carina of the females is edged with
blackish brown (Lillj.). The ventral side is as usual
lighter, in the males even whitish. A reddish streak
runs from the tip of the snout to each eye and so on
across the temples to the gill-opening.

The iEquoreal Pipefish was known as a Scandi-
navian species to Strom ( Sondm . Beskr., 1762), but
was first introduced into the Swedish fauna by Fries.
It had been described from Scotland by Sibbald in
1684; and the centre of its geographical range seems
to lie on the coast of Great Britain. It is common
among the Orkneys and Shetlands as well as on the
west coast of France. It also occurs in the Mediter-
ranean and the Black Sea (Pallas") ; but it does not
penetrate into the Baltic, though from the Cattegat it
enters the Sound, where Nilsson met with it on several
occasions off Landskrona. According to Winther it
also enters Liim Fjord. To the north, according to
Collett, it is a stationary fish “almost up to Tromso.”
Fries, who has left in the Royal Museum numerous
specimens of the iEquoreal Pipefish from Bohuslan,
found it to occur sparingly, though not rarely, in that
locality, among the seaweeds that fringe the seaward
side of the island-belt. Kroyer says that it is met
with fairly often in the Cattegat, though in compara-
tively deep water. Malm, who obtained his large spe-
cimens in 6 — 14 fathoms of water, and found only
small ones at a depth of 2 — 4 fathoms, makes the
same statement of Bohuslan. At Tylo, off Halmstad,
Ave have found young specimens 14 cm. long, also in
about 2— 4 fathoms of water.

“This species,” says Couch6, “is more especially an
inhabitant of the open ocean, where in summer our

a Rathke did not find it, however, among the fishes of the I
b Fish. Brit. Isl., vol. IV, p. 356.
c ibid., p. 359.

d Andrews, Zoologist, vol. XVIII (1860), p. 7053.

fishermen report that they see it near the surface over
a depth of more than fifty fathoms, at a distance from
land of ten or fifteen leagues.” “Sometimes,” he says
in another passage6, “it abounds in incalculable numbers
from near the shore to several miles in the open sea;
and it is then they appear to perform a perhaps limited
migration or change of quarters; for they swarm at
the surface in fine weather from the early part of sum-
mer to its declension; but after this time they are not
seen, and probably have gone to the bottom, and into
deeper water. When on our coast their actions are
amusing, as with their slender and prehensile tail they
lay hold of some loose and floating object; with the aid
of which, and the anterior portion of the body free,
they steer their wandering course by the waving action
of the dorsal fin.” Their progress thus costs them but
little trouble; but they also run great risk of being-
devoured by fishes-of-prey. The stomach of a Pollack
has been found, according to Couch, to be crammed
with iEquoreal Pipefish.

The iEquoreal Pipefish spawns in summer. The
male and female attach themselves beside each other
to some sprig of seaweed or stalk of grass-wrack d, and
the eggs are imbedded in the layer of mucus deve-
loped on the ventral side of the male, from the vent
to the isthmus. This layer hardens into a solid disk,
which, at least at the beginning of the period of gesta-
tion, may be peeled from the belly, though it then
leaves in the skin traces of the honeycombed depres-
sions which have been occupied by the eggs. The eggs
are considerably smaller than those of the Deep-nosed
Pipefish, but also far more numerous. In a female
44 cm. long the ovaries were 84 mm. in length and /
the eggs about V2 mm- in diameter. On the ventral
side of a male, where at the middle of the length of
the belly the eggs were set in 12 or 13 somewhat irre-
gular, longitudinal rows, the largest eggs were not
much more than 1,/2 mm. in diameter. The narrow
embryos lie coiled in several rings within the egg;
when 11 mm. long they have burst the membrane, but
lie with the head fixed in the cavity where they were
developed. The head is far less developed than in the
larva1 described by Fries of Nerophis lumbriciformis
(Plate XXIX, fig. 4, a); but the form of the body is

:-k Sea.

STRAIGHT-NOSEI) PIPEFISH.

683

still more elongated, the pectoral tins are comparatively
larger, the still rudimentary dorsal tin is hardly so
advanced in development, and the embryonic vertical
tin of the tail, though it is distinct in some of these
embryos preserved in spirits, in others is indistinct,
because it is either shrivelled up, or as yet undeveloped.
I have not succeeded in discovering the least trace of
the future caudal tin.

The food of the dEquoreal Pipefish is probably
the same as that of the other Syngnathince. Still, to
the best of our knowledge, we have no other direct
information on this point than that Andrews (1. c.)
has seen these fishes stripping the stems of Zostera
marina of the young of Antliea cereus , which were
attached in a semiglutinous state.

THE STRAIGHT-NOSED PIPEFISH (sw. tangsnipan).

NEROPHIS OPHIDION.

Fig. 174 and Plate XXIX, fig. 3.

Caudal Jin wanting. Vent situated below the anterior part, of the dorsal fin , the distance between which and the
tip of the snout is more than 90 % ( 92 %?) of that betiveen the vent and the same point and more than 7 (7l/s?)

times the length of the head.

Fig. 174. Head and forepart of a Neropliis ophidion ($) from Morko (C. U. EkstrSm), twice the natural size.

D. 34—40; Ann. 90—100 = (29 — 33) 4- x.

Syn. Syngnathus teres, pinnis pectoralibus candaeque carens, Art.,
Descr. Spec. Pise., p. 1 (excl. syn.); Lin., Fn. Suec., ed.
I, p. 126.

Syngnathus Ophidion , Lin., Syst. Nat., ed. X, torn. I, p.
337; Retz. A. J., Fn. Suec. Lin., p. 312; Ekste., Vet.
Akad. Ilandl. 1831, p. 280, tab. II, figg. 3 et 4; Nilss.,
Prodr. Ichth. Scand., p. 67 ; Retz. A., Vet. Akad. Ilandl.
1833, p. 157, tab. V; Fr., ibid. 1837, p. 36, tab. Ill, fig. 4;
Yare., Brit. Fish., ed. 1, Suppl., part. II, p. 47 ; Ke. ( Ner -
ophis ), Danm. Fisk., vol. Ill, p. 716; Nilss. ( Scyphius ),
Skand. Fn., Fisk., p. 694; Sundev. ( Syngnathus ), Stockh.
L. Hush. Sallsk. Handl., PI. 6 (1855), p. 164; Mgrn, ( Nero -
phis), Finl. Fistcfn. (disp. Helsingf.), p . 70; Lindstr., ( Scy-
phius), Gotl. L. Hush. Sallsk. Arsber. 1866, p. 24 (sep.);
Gthr (Ner ophis), Cat. Brit. Mus., Fish., vol. VIII, p. 192;
Dum., Hist. Nat. Poiss., tom. II, p. 602; Canestr., Fn.
D'ltal., Pesci, p. 145; Coll., Forh. Vid. Selsk. Chrnia 1874,
Tillaegsh., p. 202; ibid. 1879, No. 1, p. 101; N. Mag.
Nattirv. Chrnia, Bd. 29, p. 114; Malm (Scyphius), Gbgs.,
Boh. Fn., p. 597 ; Winth. (Nerojohis) Naturh. Tidskr. Kbhvn,
ser. 3, vol. XII, p. 54; Bncke (Syngnathus), Fisch., Fischer.,
Fischz. W., O. Preuss ., p. 190; Hcke (Neropliis), Arch. Na-
turg., Jahrg. 46 (1880), I, p. 335; Mob., Hist. Nat. Poiss.
Fr., tom. II, p. 68; Day, Fish. Gt. Brit., Irel., vol. II, p.
262, tab. CXLIV, fig. 5; Storm, N. Vid. Selsk. Skr., Trondhj.
1883, p. 42; Mob., Hcke, Fisch. Osts., p. 104; Lillj., Sc.,
Norg. Fisk., vol. Ill, p. 470.

Syngnathus lumbricif ormis , Jen., Man. Brit. Vert. Anim., p.
488.

The Straight-nosed Pipefish never attains so great
a size as the preceding species; even 3 dm. is quite a
considerable length for it, and though the females may
exceed this measurement by some millimetres, it is im-
probable, on the other hand, that the males ever attain
it. The largest male we have had the opportunity of
examining (from Norway, through Marklin), is 2J/a
dm. long".

The form of the body is still more elongated than
in the preceding species, and its greatest depth, even in
the females (excluding their vertical dermal carime),
does not exceed 2 %h of its length, while in the males
this percentage is between Vf and l3/4. At the be-
ginning of the tail (the anal ring), here as in the pre-
ceding species, we find a sharp break, at which the
depth of the body in the young specimens and the fe-
males measures only about l'l % of its length, in the
older males 1‘2 % thereof, and from which the tail
gradually tapers backwards almost to a filament. Both
in the females and the males the tail is terete (of cir-
cular section); and in the latter the breadth of the
trunk may be even greater than its depth, though as
a rule the case is the reverse; but in the older females

“ The largest male from Morko presented to the Royal Museum by Ekste6m is 197 mm. long.
6 The highest percentage we have found is 1'9.

684

SCANDINAVIAN FISHES.

the trunk is distinctly compressed, and its depth in-
creased by two longitudinal dermal folds, one at the
middle of the back and one at the middle of the belly,
both of inconstant height and occurrence, the former
always small, but the latter sometimes almost as deep
as the body. The depth of the body at the middle of
the trunk, including these dermal folds, may rise in
the females to at least 2'8 % of its length.

In adult specimens the middle carina of the plates
on the body is extremely indistinct, and the grooves
that otherwise radiate (vertically) from this carina,
are for the most part effaced and exchanged for
small, round or square hollows, which at the middle
of the central plates of the body, are arranged in
vertical rows, a distinct trace being thus left of the
original grooves. The hind part of the head, the fore-
head, and the opercula, which are without carina?, are
also punctated more or less irregularly with similar
cavities.

The length of the head in proportion to the length
of the body is less here than in any of the other Scan-
dinavian Syngnatliince. In specimens about 2 dm. long
the length of the head may still measure rather more
than 6 % (according to Kroyer even &1/2 %) of that
of the body; but in older specimens this percentage
sinks to 5l/2 or even a little less. While this change
is taking place, the length of the snout increases from
about 41 % to 46 Va 0/0 of that of the head, and the
longitudinal diameter of the eye decreases from about
V2 to about 3/10 of the length of the snout. The form
of the snout is also remarkable for its fairly prominent
dorsal margin and broad ossa symplectica, the muzzle,
when seen from the side, thus showing scarcely any
break from the rest of the head, and also tapering evenly
forwards and being only slightly turned up at the tip.
The least depth of the snout, just behind the articula-
tion of the lower jaw, measures about V3 of its length,
varying, however, between 30 and 38 % thereof. The
tip of the lower jaw falls short of the upper margin
of the tip of the snout, and in this species as in the
preceding and the following ones the length of the
lower jaw7 is about equal to the longitudinal diameter
of the eyes.

The dorsal tin is of fairly uniform height, or a
little higher at the middle, and its height in proportion
to the length of the body is about the same as in the
preceding species (about 2 %); but its base is shorter
(about 10 — ll2/3 % of the length of the body"), and
its position entirely different. In the males the distance
between its beginning and the tip of the snout is about
45* — 48 % of the length of the body, in the females
52 — 53 V2 % thereof. The vent is also situated far back,
though it lies further forward in relation to the dorsal
fin, generally belowr the beginning of the second third
thereof. In the males the distance bettveen the vent
and the tip of the snout is about 48° — 52 %, in the
females 55 d — 58 % of the length of the body. These
proportions bring about a more striking difference be-
tween the sexes, namely that in the males of this spe-
cies the length of the trunk behind the gill-covers is
less, in the females greater than the length of the tail.

The coloration of the living fish, according to Fries,
is olive-green above, with a. dash of yellow7 below7. The
sides are marked with rows of numerous, often round,
small, bluish wdiite spots, and the gill-covers are cross-
ed by fine streaks of a handsome azure blue, wdiich
are interrupted by the branchiostegal membrane, but
continued a little wray along the sides of the body.
The iris is brownish red according to Malm, silvery
wdiite according to Ekstrom, with fine, gray and coarser,
red spots. In coloration the sexes are hardly dis-
tinguishable; but according to Ekstrom the colours
of the male are darker and dirtier.

In Sweden the Straight-nosed Pipefish is quite as
common as the Deep-nosed species, in some localities
even commoner; and both species generally occur to-
gether. In the Baltic this species was known even to

o

Artedi from the coast of Angermanland, and according
to Mela it penetrates to the head of the Gulf of Fin-
land. On the Norwegian coast it is unknown north of
Trondhjem Fjord, but in the south of Norway it is as
common as in Swreden. In Kiel Bay, according to
Heincke, it is less common than the Deep-nosed Pipe-
fish. On the English coast and the north -wrest coast
of France it seems to be less common than in Scan-
dinavia; but in the Bay of Biscay, according to Moreau,

a In the smallest of KrOyer’r specimens, however, this percentage was 12'3.
b According to Moreau, however, 4 1 .

c This percentage may also occur, however, in females of the preceding species, and according to Moreau it may be 45 in this species.
d According to Kr0Yer 54.

STRAIGHT-NOSED PIPEFISH.

it is fairly common. It occurs in the Mediterranean off
Nice, according to Moreau, and on the coast of Algiers,
according to Guici-ienot and Dumeril. According to
Canestrini it lives round all the coasts of Italy and is
common in the Adriatic — assuming that these Medi-
terranean fishes are really identical in species with our
Straight-nosed Pipe-fish". Like the rest of our Pipe-
fishes this species is unknown in the western region of
the Atlantic.

The most important contributions to our knowledge
of the habits and life of this species have been made by
Ekstrom and Fries. As it glides through the grass-
wrack or algte with its serpentine or Eel-like movements,
it shows greater litheness and also greater timidity than
the Deep-nosed Pipefish, but possesses the same pro-
tective likeness to a stalk of seaweed. From its long,
terete, tapering, highly flexible, and finless tail, writes
Fries, it derives little, if any, assistance in its progress
through the water. This organ is generally kept still,
as the fish quietly swims along, and is to be regarded
rather as a rudder than an oar. When the Deep-nosed
Pipefish is stationary or at rest, it sinks, stretched at
full length, to the bottom, and lies on the belly with
tail extended. The Straight-nosed Pipefish, on the other
hand, coils its flexible tail with great skill round the
objects near at hand, and by the help of this organ
preserves an upright position in the water. It may be
seen continually attaching itself in this way, if it can
find anything round which to twine; and when it fails
in this, but has several companions in the same vessel,
one may often see them twist their tails together and
form groups, not unlike tufts of grass-wrack, which
remind us in a manner of the old figures we see of
so-called “Ratzen-Konige.”

The most striking resemblance of the Straight-
nosed Pipefish, however, is its similarity to Chorda
filum , to which it is often found attached, in those
localities where this seaweed floats about at the surface.
Like the rest of the Pipefishes this species lives on mi-
nute marine animals of various kinds, worms, crustaceans,
and mollusks, of a size suitable for its tiny mouth.

“Towards the end of April,” writes Ekstrom from
the island belt of Morko, “the females desert the shore
and the shallows to join the males in deep water and
to perform the operation of spawning. The eggs, which
are fairly large in comparison with the fish, about 1

685

mm. in diameter, are attached to the surface of the
belly from the head to the vent in 2, 3, or 4 rows, not
exactly opposite each other, but so to speak, decussating.
They are somewhat depressed in the skin, united by or
rather packed in tough mucus, and coated with a mem-
brane so extremely fine that it can scarcely be detected
and cannot bear the least touch without breaking. When
this membrane is removed, and the eggs are loosened
from the belly, they hang together like the beads of a
necklace. As soon as the fish is dead, the roe falls
from the body, accompanied, however, by the said layer
of mucus in which the roe-strings have been imbedded.
The eggs seem as though they might easily be detached
from the almost smooth skin of the belly; but they
have a triple fastening, first to the skin, by means of
the glutinous mucus, then to each other, by the union
of the poles, and finally, also to each other, by means of
the said membrane. The laying of the eggs begins at
the end of May, but is not simultaneous in all the spe-
cimens, being considerably protracted: I have seen males
with eggs even on the 11th of August. When the eggs
are deposited, they are golden yellow in colour; but they
gradually fade. At the middle of July most of them
are white, with a yellow spot on the part of the surface
most remote from the body of the male. In some cases
the yelloAv spot is already furnished with two extre-
mely fine, black dots, the first signs of the embryo
with its eyes. We can thus state with certainty that
the spawning-season, which begins during the first,
days of May, lasts throughout this month and also
during June and July. — Neither before May nor after
September have I seen a male with roe.”

i

The new-hatched embryos are about 9 mm. long,
according to Lilljeborg, with the snout turned up like a
pug’s, the eyes and the postorbital part of the head about
equally long, the entire length of the head about ’/g of
that of the body, the vent situated distinctly in front
of the middle of the body (at a distance from the tip
of the snout equal to about 44 % of the length of the
body), and the sides of the trunk furnished with a num-
ber of prominent protuberances. The embryonic vertical
fin runs along the dorsal edge, back from the middle of
the trunk, round the tip of the tail, and along the
ventral edge forward to the remainder of the vitelline
mass, but with a sharp break at the vent. The larval
pectoral fins are comparatively large. All these fins are

Cf. notes b and t’ on the preceding page.

686

SCANDINAVIAN FISHES.

as usual entirely destitute of tin-rays. According to
Mobius and Heincke young specimens less than 1 dm.
long have longitudinal carime on the body like those
of the Deep-nosed Pipefish, and these carime are fringed
with spines growing in a backward direction on the
plates of the body, so that the body, when seen from
the side, presents a serrated appearance. The pectoral
tins do not begin to be reduced until the fish is 9 cm.
long; and according to Collett these fins may still be
persistent, with a length of 1 mm., in specimens 112
mm. long. The Straight-nosed Pipefish thus undergoes
a remarkably complete series of developmental changes,
a retrogressive metamorphosis, with the earlier stages
possessing the structure and external form of more fully
equipped Syngnathince,. The history of its development
is the most distinct and complete illustration we possess
of the development (phylogeny) of the whole family.

The Straight-nosed Pipefish is of no greater im-
i portance in an economical respect than the rest of our
Syngnathince. Whether it can do any harm by de-
vouring the deposited eggs of other fishes, or by reducing
the supply of food for the fry of more valuable species,
is a point on which we have no information. That it
may be used, at least in its younger days, as human
food, is shown, however, by its appearance amongst
whitebait a, for, like the Sticklebacks, it now arid then
forms a part of this favourite English dish, which con-
sists chiefly of Herring- fry. Its common occurrence
renders it well-known to the fisherman, and on the
east coast of Sweden it is this species in particular that
bears the name of liafsnal (Needle-fish), which Fries
adopted for the whole genus Nerophis. In Halland it
is called tangsnipa1 , a reminiscence of the Danish snippe ,
which Krdyer has transferred to the following species.

THE WORM PIPEFISH.

NE ROPHIS L U MBRICIFORMIS.

Fig. 175 and Plate XXIX, fig. 4.

Caudal fin wanting. Vent situated below the anterior part of the dorsal fin , the distance between which and the
tip of the snout is at least 90 % (90 — 92 %) of that between the vent and the same point , but less titan

5 times the length of the head.

Fig. 175. Head and forepart of a Nerophis lumbriciformis , 9) from the Weather Islands (Bohuslan). Twice the natural size.

D. 25—26; Ann. 69— 73 = (17 — 19) + x.

Sjjn . Acus lumbriciformis aut Serpentinus , 1. Ophidion lumbriciforme,
Willughby, Hist. Pise ., p. 160; Ray, Synops. Method. Pise.,
p. 47. The little Pipe-fisli, Penn., Brit. Zool. (ed. 1776),
p. 124, tab. XXIII, No. 62.

Syngnathus Ophidion, Flmng, Brit. Anim ., p. 176 (nec Lin.).

Syngnathus lumbriciformis, Yarr., Brit. Fish., ed. 1, vol. II,
p. 340; Fk., Vet. Akad. Handl. 1837, pp. 38 et 59, tab. Ill,
figg. 5 et 6, tab. IV; Kr. (Nerophis), Danm. Fish., vol. Ill,
p. 723; Nilss. ( Scyphius ) Skand. Fn., Fish., p. 695; Gthr
(Nerophis), Cat. Brit. Mus., Fish., vol. VIII, p. 193; Dum.
TIist. Nat. Poiss., tom. II, p. 604; Coll., Fork. Vid. Selsk.
Chrnia, 1874, Tillsegsh., p. 203; Malm (Scyphius) Gbgs ,
Boh. Fn., p. 598; Winth. (Nerophis), Naturh. Tidskr. Kbhvn,
sev. 3, vol. XII, p. 54; Hcke, Arch. Naturg., Jahrg. 46, I,
p. 339; Mor., Hist. Nat. Poiss. Fr., tom. II, p. 65; Day,
Fish. Gt. Brit., Irel., vol. II, p. 263, tab. CXLIV, fig. 6;
Lillj., Sv., Norg. Fish., vol. Ill, p. 477.

Obs. That it was not this species that Jenyns referred to (Alan.
Brit. Vert. Anim., p. 488), appears partly from his statement that
it may attain a length of 9 inches (228 mm.), partly from his
description, which, as Fries has already remarked, can apply only to
the preceding species.

The Worm Pipefish is one of our smallest Syngna-
thince; its length is seldom more than 15 cm., and’ the
maximum size which the species is known to attain, is
165 mm.c The largest specimens in the Royal Museum
are 141 mm. (c?) and 122 mm. (?) long.

The body is of the same serpent-like form as in
the preceding species, but less elongated and still more
terete. The greatest depth is situated in full-grown
males at the beginning of the trunk, in the females at
the middle of the trunk. In the former it measures

“ Cf. Buckland, Nat. Hist. Brit. Fish., p. 170.

h Tang, lang and snipa, anything long and pointed, e. g. a boat. Tr.
0 Collett (in Lilljeborg, 1. c.). The specimen was a female.

WORM PIPEFISH.

about 2 — 2 V, % of the length of the body, in the latter
at least as much as 3 '7* % thereof. The terete tail
tapers regularly to its end. Here, as in the iEquoreal
Pipefish, the difference between the sexes is most dis-
tinctly expressed externally by the form of the trunk.
In the males this part is somewhat flattened, especially
at the middle of its length, and the breadth of the flat
ventral side, to which the eggs are attached, is greater
than the depth of the body at the same point. In the
females the trunk is laterally compressed, and also fur-
nished, when the fish is in full dress, with a longi-
tudinal carina at the middle both of the back and the
belly, the latter carina being here too the more pro-
minent. Thus the structure of the males in this respect
comes very near the form of the body in the Australian
genus Stigmatopliora , which is furnished, however, with
pectoral fins, and in which the males carry the eggs
under their tails.

In specimens from 10 to 12 cm. long the length
of the head varies between 8 (7‘8) and lxj2 %, in older
specimens between j 2 and 7 % of the length of the

body, or between about 25 1/a and 23 V3 % of the distance
from the dorsal tin to the tip of the snout. The most
characteristic point in the head of this species is the
form of the snout, a reminiscence of the larval stage
of all the other Syngnathincc. The snout is compara-
tively short, measuring in young specimens — between
10 and 12 cm. long — only about 1/4 , in older speci-
mens about V3 (30— 32V2 %) °f the length of the head,
and also turned up like a pug’s nose, with a more or
less selliform depression in the superior profile. The tip
of the lower jaw projects upwards to a level with the
tip of the snout, which is hollowed to receive it. The
longitudinal diameter of the eyes is about half the length
of the snout. The postorbital length of the head
measures about half its entire length.

The dorsal fin is of uniform height. The length
of its base is about 9 or 10 % (8’8 — 10’5 %) of the
length of the body. The distance between it and the
tip of the snout is about 30 % (29'6 — 30'5 %) of the
length of the body. The vent lies below its anterior
part, at a distance from the tip of the snout that mea-
sures about V3 (32'5 — 33'3 %) of the length of the
body. The length of the tail is thus always more than
twice that of the trunk behind the opercula.

The coloration is far from ugly. “Its usual ground-
colour," writes Fries, “is chestnut-brown, in some cases
a Nat. Hist. Irel., vol. IV, p. 242.

087

lighter, in others darker. Along the back lie large,
irregular spots of whitish gray, which are broken up
on the tail into numerous, smaller spots and give this
part of the body a mottled appearance.” The head is
also mottled above and below with whitish gray spots,
framed with darker colour, and two or three of these
spots are especially constant and prominent, obliquely
in front of, below, and obliquely behind the eyes, rising
from the isthmus. On the tail the spots are arranged
more or less distinctly in transverse rows; and in a fe-
male from the Weather Islands (Bohuslan), which lias
lain in spirits for some years, but preserved its colora-
tion remarkably well, this arrangement is distinct on
the forepart of the body also. The dermal carina on
the belly of this specimen seems to have been edged
with darker colour. The iris is yellow, with a ring of
round spots.

This coloration is probably connected with the ha-
bits of the fish. The Worm Pipefish seems to prefer
deep water, where the brown tint prevails among the
algae. It has even been found crawling on a, clayey
bottom, where it lias been taken in the dredge in com-
pany with those marine Annelids, crustaceans, and mol-
lusks whose young and larva? probably form the chief
part of its food. On the Irish coast too, according to
Thompson", this species has been taken in the dredge
in deep water, though it also lives between the tide-
marks, where it hides under stones.

The spawning-season seems to be somewhat later
than in the preceding species. The eggs are set in
four, more regular rows and less firmly attached
(less deeply imbedded in mucus). Collett estimated
their number in different males at 63 and 78; in a
male 135 mm. long we have counted 88 eggs, about
l1/-, nim. in diameter.

It was in this species that Fries made the first
observations of the retrogressive metamorphosis of the
genus Neropliis, observations which have since been ex-
tended, as we have seen, to all the Scandinavian species
of the genus. One of the last days in September, 1837,
Fries had obtained a male specimen to which the eggs
still adhered. He placed the specimen in a vessel of
water, and endeavoured to keep it alive for some time,
in order to observe the hatching of the eggs and the
relations between the young and their father. As the
latter had no marsupium in which the young might
hide at the approach of danger, Fries expected to see

Sea n din a v ia n Fis h es .

87

688

SCANDINAVIAN FISHES.

them attach themselves in some way with the tail to
the parent fish; but in this expectation he was dis-
appointed. After the lapse of six days the fish showed
signs of weakness, and some of the eggs began to
acquire a sickly appearance. Life endured, however, for
some days more, and on the morning of the ninth day
of the captivity Fries observed three young specimens
at the surface. Later in the day the fourth appeared,
and on the following morning two more; but no more
eggs were hatched. The whole mass of eggs now pre-
sented a half-decomposed appearance, became detached,
together with the adhesive layer of cells, from the body,
and broke up piecemeal. The fish died on the same
day towards evening. The larvae lived seven days, and
during this time their size increased from 9 to 151/ 2 mm.

The first point that surprised Fries, was the be-
haviour of the young amongst themselves and towards
the parent fish. They swam in an upright position,
careless of each other, and paying still less attention to
their father, Avho lay at the bottom and was equally
oblivious of them. In order to elucidate this discrepancy
from Ekstrom’s account of the young of this genus,
Fries began to examine the fry more minutely, and
found their structure to present the appearance shown
in our figure (Plate XXIX, fig. 4, a). The whole body
is white and transparent, the spinal column and the
intestine within the abdominal cavity shining through.
The head measures about V6 of the length of the body.
The length of the snout, which curves upwards, is greater
in proportion to that of the head than in older speci-
mens. It is also worthy of remark that, whereas in all
adult Syngncitliince the margin of the opercula is united
by a membrane and by the general dermal covering to
the shoulder girdle, leaving on each side of the occiput
only a small foramen for the passage of the respiratory
water, in the young on the other hand this margin is
entirely free, the gill-openings thus being large, as in
the generality of fishes. The lower figure shows this
point with tolerable distinctness. The vent, though it oc-
cupies its right place in relation to the dorsal fin, lies
nearer the tip of the tail than in older specimens, only

a little in front of the middle of the body. The plates
of the body seem as yet undeveloped; but when the
young specimen is examined from above, we see along
each side of the body a projecting row of fine teeth,
which must be the tips of the transverse vertebral pro-
cesses that in older specimens support the dermal rings
of plates. Fries counted, as far as he could judge, 18 of
these points in front of, and at least 50 behind the vent.
The pectoral fins are small, but distinct, with widened,
somewhat rounded tip. Their motion is extraordinarily
brisk, but the rays are extremely rudimentary. The
true dorsal fin is distinct, but the traces of the inci-
pient rays are only faint. Along the dorsal margin
behind this fin, round the tip of the tail, and along the
ventral margin forward to the vent, runs the embryonic
vertical fin, the chief organ of locomotion in the fry.
Such is Fries’s description of his find; and the equip-
ment of the larvae with comparatively well-developed
organs of motion explains, it is true, their capability
of independent life. But here this explanation stops,
for the larvae of the preceding genus may be equally
well equipped in this respect, though they still take
refuge in the marsupium of the male.

The geographical range of the Worm Pipefish, as
far as we know at present, does not extend so far north
as that of the preceding species. Storm never found it
in Trondhjem Fjord; but on the coast of the Govern-
ment of Bergen it has been met with at several spots,
according to Collett. In Christiania Fjord it is not
rare. Off the Weather Islands and outside Gullmar
Fjord it has been taken on several occasions. Kroyer
and Peter Muller have found it on the north coast
of Zealand (Hornbaek and Hellebaek), but further south
in the Sound and in the Baltic it is unknown. All
round Great Britain and Ireland it is common enough,
according to Day. This is also the case on the north-
west coast of France, according to Moreau, at least
at certain spots; and according to the same author it
occurs, though rarely, even in the Bay of Biscay. No
instance is known of its occurrence in the Mediter-
ranean.

PHYSOSTOMES.

689

TBLEOSTEI PHYSOSTOMI.

Osseous fishes with the air-bladder (if present) furnished with pneumatic duct. Ventral fins (if present) ab-
dominal. No spinous rays proper (unarticulated) in the fins".

Each of these three characters calls to mind an
era in the history of ichthyology. The last was ArtediV
character of the order Malacopterygii (pinnis inermibus);
it was by the second that Linnjeus6 defined the order
Abdominales (pi nine ventrales pone pinnas pectorales);
and the first-mentioned character was Muller’s'* ex-
pedient for the better definition of Malacopterygiens
abdominaux, the order which Cuvier6 had adopted in
his system, and for the combination therewith of Cu-
vier’s Malacopterygiens apodes.

The order or rather suborder which has thus arisen,
is hardly more natural than that of the Physoclysts,
just as the whole Teleosteous order is merely an ex-
pression of the community of those characters which
the fishes belonging to the several directions of deve-
lopment of this type have retained or acquired during
their development from a common prototype, the Ga-
noids. The independence of the directions of develop-
ment asserts itself everywhere, and the characters of the
system must therefore be employed with discrimination.

The Physostomes in general stand nearer the Ga-
noids, which were probably also furnished with a pneu-
matic duct from the air-bladder. This connexion with
the Ganoids appears in the abdominal position of the
ventral tins, which have besides retained not unfrequently
a greater number of rays, just, as the pectoral fins may
display more numerous transverse rows of basal bones,
though the outer (distal) rows are not ossified7. An-
other sign of this connexion is given by the exceptions
which occur in certain Physostomes, to the character
that, otherwise distinguishes the Teleosts with regard

a The rays which in these fishes are termed spines, as being
ture, being- articulated, although the joints are coalesced.

b Gen. Pise., p. 1.

to the structure of the arterial bulb of the heart, a
structure which in this suborder may be almost Ganoid.
These resemblances to the Ganoids range the Physo-
stomes lower in the system, assuming that the system
should be an expression of the development. But for all
this we must not regard the structure of the Physo-
stomes as more primitive; many of them on the con-
trary possess more highly developed organs than the
corresponding organs in the Physoclysts. Among these
organs we find, for example, the stronger supporting-
apparatus which the pectoral fins have acquired in
several Physostomes by the addition to the scapular
disk of a special bone for this purpose (os prcecora-
coideum a). Another example of this is the far more
frequent occurrence in the Physostomes than in the
Physoclysts of the connexion between the air-bladder
and the organs of hearing, a connexion which is ef-
fected in several of them by a special chain of bones7'.

The preponderant part of the Physostomes, as op-
posed to the Physoclysts, is composed of fresh-water
fishes; but the family most important among marine
fishes in an economical respect, that of the Herrings,
is Physostomous. This order also contains the families
of the Salmons, Carps, Pikes, and Eels, all of which
have been of great importance in human economy since
time immemorial.

The Physostomes are not so numerous as the Physo-
clysts— we know about 1,900 species of the former, as
opposed to 3,300 of the latter — but the distinction of
form is sufficiently wide to require the establishment
of several independent series which we here give prin-

hard and stiff, show their Malacopterygian nature in their internal strue.

c Syst. Nat., ed. X, tom. I, p. 241. Cuvier completed the Linniean character by the addition that the ventral fins not only lie be-
hind the pectoral fins, but are also destitute of any immediate connexion with the shoulder-girdle.
d Abh. Akad. Wiss. Berl. 1844, Phys. Abb., p. 175.
e R'egne Animal , ed. 1, tom. II, p. 159; ed. 2, tom. II, p. 269.

f Cf. Gegenbaur, Unters. Vergl. Anat. Wirbeltli., 2:te Heft, p. 153, taf. VIII, figg. 8 et 9.

<J Prtecoracoid, Parker, Shoulder- girdle and Sternum , p. 56, pi. II, fig. 8, p. cr; Knochenspange, Gegenbaur, 1. c., p. 117, taf. VII,
figg- 1 — 5, , v .

h Sagemehl (Morphol. Jahrb., P>d. X, p. 22) gave these fishes the name of Ostariopliysete Teleosts, from oocagiov, a small hone, and
fpvoa, bladder.

690

SCANDINAVIAN FISHES.

cipally after Cope", but with names more adapted to
our previous treatment of the Physoclysts:

1 : The shoulder-girdle suspended from the
head.

A: An internal supporting bone (os prse-
coracoideum) for the coraco-scapular disk
rises from the coracoid bone to the in-
side of the clavicle or of the scapula.
a: Anterior vertebrae more or less con-
fluent; air-bladder connected with
the hearing- apparatus by a row of
ossicles.

a: No distinct os symplecticum; no
suboperculum; maxillary bones
rudimentary, forming in most ca-
ses the base of a maxillary bar-
bel on each side

/?: Os symplecticum distinct; sub-
operculum present; maxillary

bones normal

b: Anterior vertebrae normal; air-bladder
without osseous connexion with the

hearing-apparatus

B: No os praecoracoideum

II: The shoulder-girdle suspended from the
spinal column.

Glanomorphib .

Cyprinorn orpin c .

Thrissomorphid.

Esociformese.

EnchelymorphP.

GL AN OMOKPHI.

The shoulder -girdle suspended from the head (as usual in the Teleosts). Coraco-scapular disk internally strength-
ened by an arch formed by a special bone (os prcecoracoideum) from the coracoid bone to the clavicle or the upper
( anterior ) margin of the scapula. Most of the first four or five vertebrae more or less altered from the ordinary
form and more or less completely confluent with each other or even with the occipital bone. Some of the ribs
belonging to these vertebra metamorphosed into a connecting chain of bones between the air-bladder and: the audi-
tory apparatus. No os symplecticum in the hyomandibular arch. Each palatine arch composed of only two bones ,
os pterygoideum and os palatinum. Opercular apparatus without suboperculum. Maxillary bones reduced, generally
forming , together with their supplementary bones , the mobile base of a maxillary barbel on each side. No true

scales, body naked or covered with plates.

Valenciennes g ranged the Glanomorphs at the head
of the Malacopterygians (nearest to the Acanthoptery-
gians), because they possess the most strongly ossified
fin-rays. This character the possession in the fore-

most pectoral rays and the first (second) dorsal ray of
admirable weapons of defence and attack — is indeed
fairly constant in its appearance, but still variable
enough to render it useless to the systematise The
Glanomorphs also vary so greatly both in the rest of
their structure and in the form of the body that the
distinction between them and the next series of fami-
lies consists principally in the negative characters given
above, the simpler (less complete) structure of the oper-
cular apparatus, which is without suboperculum, of the
hyomandibular arch (the suspensory arch of the lower
jaw and the hyoid bone), which is made up on each side

a Trans. Amer. Philos. Soc., vol. XIV, n. ser. (1871), p. 452.
1 Nematognathi, Cope.
c Plectospondyli, Cope.
d Isospondyli, Cope.
c Haplomi, Cope.
f Apodes, Lin., auctt.

rj Cuv., Val., Hist. Nat. Poiss., vol. XIV, p. 309.
h Atl. Ichth. Lid. Or. Ne'erl., tome II, p. 2.

‘ Cf. Valenciennes in Cuv., Val., Hist. Nat. Poiss., vol. XV,

of only two bones, os hyomandibular e and os quadratum,
and lastly of the palatine arch, which is also composed
on each side of only two bones, os pterygoideum and
os palatinum.

The systematic arrangement of the Glanomorphs
has always presented great difficulty. BleekeiG based
his division of this series into families in the first place
on the singular respiration of certain forms. Apart
from the fact that some ( Plotosus ) are furnished behind
the vent with ramified dermal excrescences', which
Bleeker explains as respiratory organs, others (Hetero-
branclius and Cl arias ) possess similar growths of the
mucous membrane of the branchial cavity, on the se-
cond, third, and fourth branchial arches, this apparatus
being concealed in a cavity behind the branchial cavity
proper. Others again ( Saccobranchus ) are destitute of

p. 411.

GLANOMORPHS.

these growths, but are furnished instead on each side of
the body with a long, cylindrical duct penetrating into
the dorsal muscles, a backward continuation of the
branchial cavity, containing a respiratory sac which is
lined with a continuation of the mucous membrane
of the said cavity. This continuation of the mem-
brane is abundantly furnished with ramifications of the
branchial arteries (the first artery on the right side, the
last on the left"), which convey hither a great portion
of the venous blood of the heart. The blood oxyge-
nated in each of the respiratory sacs by inhaled air* 6 is
partly carried off by arterial ramifications in the sur-
rounding muscles, but most of it is collected in a main
artery, that discharges it into the great dorsal artery.
The whole of this arrangement, it is true, stands in a
physiological respect beside the structure just described
in Heterobranchus and Clarias; but the form of the body
and the structure of the fins range Saccobranchus im-
mediately beside Silurus. An amphibious manner of
life has produced a development of special respiratory
organs for the breathing of air; but nevertheless the
natural kinship may be most safely expressed by the
form of the body and the structure of those external
organs which otherwise afford the most tenable cha-
racters within this series of families.

This is also true of the variations of the air-bladder.
In our Sheatfish it is externally simple, but internally
divided in a longitudinal direction. In Bagrus these
divisions are transverse. In Malapterurus c the anterior
part is externally set off, though not internally divided,
from the posterior, which is partitioned internally, though
incompletely, both in a longitudinal and a transverse
direction; and the anterior part itself is externally di-
vided in a longitudinal direction into two oval chambers,
which communicate, however, with each other internally.
In BMnolepls d the posterior part of the air-bladder has
disappeared, and the two oval, anterior chambers lie
side by side, each enclosed in an osseous capsule. These
among other variations in the structure of the air-
bladder in the different Glanomorphous genera show
that the form of this organ is highly plastic; but up

“ Hyrtl, Stzber. Akad. Wiss. Wien, Math. Naturw. CL, Bd. 1

6 This according to Day. Gunther states, however, ( Introd .
vity receive water.

c Geoffroy, Descr. cle VEgypte , Poiss ., pi. XII, fig. 4.
d Reissner, Arch. Anat., Physiol., 1859, p. 421, Plate XII.
e Stzber. Akad. Wiss. Wien, Math. Nat. Cl., Bd. XI, p. 145.
f Fishes of India , p. 440.

a Weber, De aure et auditu etc., Lips. 1820, p. 53.
h Sorensen, Om Lydorganer hos Fislce, Kbhvn 1884, p. 121

691

to the present the differences are far too little known
to be employed in the system. Kner also came to
the conclusion6 that within the Boras group the diffe-
rences in the form of the air-bladder may well be em-
ployed in the distinction of the species, but not in the
definition of the genera. Day7 has, however, employed
these relations in the classification of the genera; but
he remarks on this head that the marine Glanomorphs
in general have a thick air-bladder, not enclosed in an
osseous capsule, as well as the majority of the fresh-
water forms; but that the further we penetrate into the
highlands of India, the oftener we meet with Glano-
morphs whose air-bladder is cased in this manner. It
would thus seem that here too the amphibious manner
of life exerts a determinative influence.

The incasement of the air-bladder is also merelv a

J

part of the marked ossification that takes place in the
anterior portion of the spinal column. Here, as we have
mentioned, the anterior vertebrae are more or less com-
pletely confluent; and their ribs are transformed into so-
called acoustic bones (Weberian ossicles7), which serve to
connect the air-bladder with the organs of hearing, and
may well deserve their name, though they do not pos-
sess the least homology with the otosteals ( ossicula
auditus ) of the higher Vertebrates. Here this ossifica-
tion of the first vertebrae — conjoined in many Glano-
morphs with occipital plates (interneural growths) —
forms a strong support for the spinous ray of the first
dorsal fin, one of the powerful weapons of these fishes.
Both these structures are also connected, according to
Sorensen, with the power possessed by several Glano-
morphs of producing sounds6. The dorsal spines — the
rudimentary (first) and the great (second) spinous ray —
produce noises (like the spinous ray of the pectoral fins)
by means of the friction between their articular surfaces
and those of the interspinal bones; and the air-bladder
utters sounds partly in the same way as in the Gur-
nards, Filefishes, and others (see Part I), partly by
means of the vibration produced by the violent contrac-
tion of the special muscles running from the occipital
region or the foremost (confluent) vertebrae to the acous-

.1 (1853), p. 305.

'itudy Fish., p. 565) that the lung-like extensions of the branchial ca-

ete.

692

SCANDINAVIAN FISHES.

tic bones, which are attached to the anterior end of
the air-bladder. Several circumstances, of which we
shall give an account below, seem, however, to assign to
the Weberian apparatus its most important function as
a barometrical apparatus for the fish. As in the File-
fishes, we sometimes find here that the air-bladder is
connected with the skin in the scapular region on each
side of the bod}- ; and this is effected either immediately,
lay means of lateral processes from the bladder, or by
a ligamentous connexion between the skin or the first
lateral plate and the acoustic bone that coalesces with
the air-bladder.

In the same parts of the body, the sides of the
scapular region behind the clavicles, we find in some
Glanomorphs ( Plotosus , MaJapterunis") special cavities,
one on each side of the bod}7, more or less completely
closed bulbs of the abdominal cavity (secondary abdo-
minal cavities), in which a part of the liver and (on the
right side) the gall-bladder or a portion of the kidneys
may find room. In several Glanomorphs {Doras, Arias,
Aspredo, Silurus etc.) we meet with a singular sac be-
neath the skin behind each of the clavicles, usually
opening into a hole ( poms lateralis 1. pectoralis ) at the
upper angle of the pectoral fin, somewhat above or a
little behind it. We have already remarked a similar
cavity in the case of the Batrachoids (Part I, p. 133).
It has been interpreted as a saccate poison-gland; but
in neither case has its signification been yet explained.
In our Sheatfish the pore leads upwards to the cavity
under the flat, backward process from the posterior
corner of the angular clavicle.

These fishes are known in English as Catfishes on
account of their long and sometimes numerous barbels,
sensory threads (corresponding to whiskers) round the
mouth. These barbels, accompanied sometimes by large

labial flaps, are by their extraordinary development
especially characteristic of the Glanomorphs.

The variety of form among the Glanomorphs (Ne-
matognates) is fairly great. About 660 species have
been described, and Bleekee has distributed them among
185 genera, arranged in 6 families. The form of the
body varies from the ordinary piscine form, usually
with flattened head and well-separated caudal fin, to the
Anguilliform type, with the vertical fins continuous
round the tip of the tail. The plated Glanomorphs, as
Cope has remarked, present in their armour an obvious
resemblance to the Sturgeons, to which the Glanomorphs
possess another likeness in the barbels on the jaws. That
the resemblance between these two groups of fishes is
not merely external, has been already shown by Ban-
ker6 in his account of the analogy between the different
parts of the scapular arch in the armoured Glanomorph
Callichthys and in a Sturgeon, with posttemporale, supra-
clavicula, clavicula and inter clavicula equally distinct as
dermal growths. Thus the Glanomorphs — in the ma-
jority of which, however, the structure of the scapular
arch reminds us most strongly of the Carps — rank as
the most distinct intermediate forms in this respect
between the Sturgeons and the Teleosts.

Most of the Glanomorphs belong to fresh water,
especially to the lakes and rivers of the Tropics, with
their deltas and brackish water; but many live in the
sea, where they keep, however, to the littoral region.
To the natives, especially within the Tropics, these fishes
are in many localities of great economical value; but
they are generally despised by Europeans on account of
their repulsive appearance. Many of them too, especially
in the tropical regions, are dreaded for their formidable
spines, which are said to be poisonous, though as yet
we have no trustworthy evidence on this head.

Fam. SILURIDiE.

Body naked {without plates on the sides); head also covered with skin. Only one {if any) dorsal fin furnished with rays,
short and situated on the abdominal part of the body. Caudal part of the body and the anal fin long. Branchial
arches simple {without appendicular branchice). Gill-openings large ; branchiostegal membranes free from the isthmus.

Thus defined, the family of the Sheatfishes corre-
sponds to Bleeker’s subfamilies Siluriformes and Ailice-
formes — the former without, the latter with a small
adipose fin on the posterior part of the dorsal margin

a Cuv., Val., Hist. Nat. Poiss., vol. XV, pp. 415 and 525 a
h Shoulder-girdle and Sternum (Ray Soc. 1867), p. 23.

— as well as to his Saccobranchi formes, with their sin-
gular, tubular, respiratory cavity along each side of
the body, and contains about 50 known species from
the Old World and the Sunda Islands. In Gunther

1 Hyrtl, ]. C., p.

302.

SHE ATFI S H E S .

693

the family is represented chiefly by the subfamily Silit-
ridce Heteropterce , with the single exception that Blee-
ker has referred the Pangasince, with stronger spinous
ray in the comparatively longer dorsal tin and with
comparatively shorter anal fin, to the Bagriform division,
which in conjunction with a more advanced development
of the dorsal spine possesses a support for this spine in
a distinct cuirass on the head and the anterior part of
the back. The essentia] character of the Siluroids pro-

per thus lies in the reduction of the dorsal tin (or tins,
both of the rayed, anterior tin and of the adipose tin),
combined with the elongation of the anal tin. Gun-
ther’s Heteropterce (with different fins) expresses this
character fairly well. The whole Glanomorphous series
is so nearly continuous, however, that up to the pre-
sent the limits of the families which we are compelled
to establish by the great number of the species, are in
a measure arbitrary.

Genus SILURUS.

No adipose fin. Dorsal fin soft-rayed ( without spinous ray). Dense cards of pointed teeth in the lower jaw, on
the intermaxillary hones, and on the head of the vomer. Barbels 4 or 6, two of which belong to the upper jaw ,
the rest to the lower. Anal fin united at the base by a membrane to the rounded caudal fin, but at the margin

separated from the latter fin.

The genus Silurus, the name of which occurs even
in ^Elian and Athenasus (in the second century A. D.),
Avas in Artedi “ a combination of the Burbot and the
Sheatfish, though the former Avas also included in the
genus Gadus. In Linnaeus6 it embraced, together Avith
the genus Loricaria , the Avhole of that series of families
Avhich Ave have iioav called Glanomorphs. At present,
Avith the definition given it by Gunther, it contains

5 known species, 3 from India, 1 from Dauria, China,
and Japan, and 1 from Europe. An East Indian ge-
nus of small fishes, Silurichthys, comes extremely near
Silurus, but is distinguished therefrom by the oblique
shape of the caudal fin, Avith the upper corner con-
siderably longer than the loAver, a character of Avhich
Ave find at least traces in Silurus , and by the closer
union of this fin to the anal fin.

THE SHEATFISH (sw. malen).
SILURUS GLANIS.

Plate XXX, fig. 1.

Six barbels. Dorsal fin situated in front of the middle of the abdomen, about half-way between the vertical lines
from the anterior ends of the insertions of the pectoral and ventral fins. Vomerine card of teeth continuous, with
only a slight indentation at the middle of the hincl margin. Lower jaw projecting beyond the tip of the snout.
Coloration olive-green, above dark, almost blackish, below pcde, fading even into whitish, on the sides spotted;
all the fins reproduce the coloration of the body ( darker base, lighter margin), except the dorsal fin, which is plain,

of the same colour as the back.

V.

R. br. 15 I. 16; D.
1

1

(83) 87 (89)’ P' (14) 15 (17)’

(10) 11 (12)

2—3

; (J . x + 1 7 + x ; Vert. 20 + 51 (54)

Syn. Gadus, Lin., Fn. Suec ., ed. 1, p. 109, No. 291; Silurus, It.

Scan., p. 62. Silurus cirris duobus ad maxillam snperiorem,
quatuor in mento, Gronov., Mus. Ichth. (1754), p. 6; Mai ,
Osb., Vet. Akad. Hand]. 1756, p. 34. Weis, Bl., Fisch.
Deutschl., part. I, p. 242, tab. XXXIV.

Silurus Glanis, Lin., Syst. Nat., ed. X, tom. I, p. 304; Holm,
D. Vid. Selsk. Skr., XII (1779), p. 133, tab. I et II; Retz.,

Fn. Suec. Lin., p. 343; Pall., Zoogr. Ross. Asiat., tom. Ill,
p. 82; But, Rtzb., Medic. Zool., part. II, p. 31, tab. V,
fig. 2, C, D, E, tab. VI, figg. 1—4; Nilss., Prodr. Ichthyol.
Scand., p. 38; Cuv., Val., Hist. Nat. Poiss., tom. XIV, p.
323, tab. 409; Kr., Panin. Fiske, vol. Ill, p. 120; Nilss.,
Skand. Fn., Fisk., p. 359; Lloyd, Ant. 20-dr. vist. Skand.,
p. 40; Hckl., Kn., Siisswasserf. Oestr. Mon., p. 308, fig.
165; Sieb, Siisswasserf. Mitteleur., p. 79; Mgrn, Finl. Fiskfn.
(disp. Helsingf. 1863) p. 35; Gthr, Cat. Brit. Mus., Fish.,
vol. V, p. 32; Seidl., Fn. Balt., p. 100; Buckl., Nat. Hist.
Brit. Fish., p. 357; Mor., Hist. Nat. Poiss. Fr., tom. Ill,
p. 439; Mela, Vert. Fenn., p. 310, tab. X; Bncke, Fisch.,

a In Artedi the Sheatfish is mentioned only in the Appendix {Gen. Pise., p. 82 and Syn. Pise., p. 110), and has probably been in-
troduced thither by Linnaeus daring his revision of Artedi’s Ichthyology. See Skdnska resan, p. 62.
h Syst. Nat., ed. X. tom. I, pp. 304 and 307 ; ed. XII, tom. I, pp. 501 and 508.

694

SCANDINAVIAN FISHES.

Fischer., Fischz. W '., 0. Prenss., p. 103. fig. 83; Grimm,
Fish., Hunt. Russ. Wat., pp. 12 et 19; Apostol., Peche en
Grece, p. 31; Norback, Handl. Fiskev., Fiskafv., p. 373;
Bncke, Hanclb. Fischz., Fischer. (M. v. d. Borne), p. 142,
fig. 150; Lillj., Sv., Norg. Fisk., vol. Ill, p. 358.

The Sheatfish is the largest osseous fish in the fresh
wafer of Europe. According to trustworthy verbal sta-
tements" it attains in the south of Russia, in the Dnie-
per, a length of at least 48 dm. and a weight of about
295 kgm. Kramer (1756), quoted by Heckel and Kner.,
mentions a still larger Sheatfish from the Danube, “so
large that two men could scarcely join hands round its
body,” which thus might have measured between 6 and
7 metres in length. The Sheatfish described by Osbeck
(1. c.) was from Lake Bafven in Sodermanland and
weighed 103/5 kgm. at a length of 1 1 1/s dm. In Swe-
den the species seldom attains a length of more than
2Yg metres, though we sometimes hear of specimens
half as long again h

The form of the body reminds us strongly of the
Burbot, with flattened head, terete trunk, and laterally
compressed tail, though these three parts pass evenly
into each other. Lithe as the Burbot — or even more
so — and still more like the Eel in its movements, with
loose and slippery skin, the body of the Sheatfish may
assume different forms in different position; but when
extended and at rest it presents the appearance given
in our figure. The dorsal profile is almost straight,
only slightly elevated at the end of the first quarter,
where the dorsal fin is situated, from this point slightly
curved towards the tip of the snout, and with a down-
ward curve just behind this fin. The ventral profile is
more regularly curved, more sharply ascending in front
when the mouth is closed. The flat snout, the convex
forehead, and the forepart of the back, which is also
convex, but furnished at the middle with a more or less
deep, longitudinal groove all the way from the occiput,
are continued behind by the narrow, but terete dorsal
edge of the tail. The ventral side, on the other hand,
is indeed terete in front and along the belly itself more
or less tumid; but behind the vent it is so sharp that
the interhaemal spines (supporting bones) of the anal fin
are even externally perceptible beneath the skin. The
greatest depth of the body, which occurs at the dorsal
fin, measures in young specimens, 2 or 3 dm. long,
about 19 % of its length; while in older specimens 2

metres long this percentage usually sinks to 13c. The
thickness at the same spot is in young specimens usually
distinctly less than, in old specimens generally about
equal to the depth. The least depth of the body, just
in front of the caudal tin, measures in the said young
specimens about 5 % of its length, in the older ones
about 4 — 3 7 2 % thereof.

The length of the head (from the tip of the snout
to the extreme end of the branchiostegal membrane)
measures 7s — Vis of that of the body (from the tip of
the snout); but in old specimens the lower jaw projects
in front of the tip of the snout for a distance that
sometimes measures at least 1/9 of the length of the head.
The broad, flat snout, with its sides sharply rounded
(forming a broad ellipse) in front, passes very gradually
into the terete form of the occiput and the forepart
of the trunk, without any sharp lateral break, the
head thus resembling a broad wedge with rounded sides.
The breadth of the head across the opercula is the great-
est breadth of the body (in old specimens about 14 %
of the length of the body); but we are most struck by
the great breadth of the gape, the width of which,
measured straight across the corners of the mouth when
closed, is about 4/5 of the greatest breadth of the head,
and occupies almost the entire breadth of the snout.
The lips are fleshy, the dermal folds at the corners of
the mouth being especially thick. In young specimens
the mouth shuts so tightly that the upper and lower
lips are contiguous; but in older ones the lower jaw
projects so far that a considerable portion of its card of
teeth is left bare. The eyes are round and small, and
sometimes even difficult to distinguish, as their colour
differs only slightly from that of the head. Their rela-
tive size varies considerably according to age. In spe-
cimens between 1 and 2 dm. long the longitudinal dia-
meter of the eyes measures about 10 % of the length
of the head, 20 — 19 % of the breadth of the inte rorl iital
space, or 30—27 % of the distance between them and
the middle of the tip of the snout (the length of the
snout). In specimens between 1/2 and 1 in. long these
percentages have sunk, the first to about 6, the second
to about 13 or 12, and the third to about 18 or 17.
In a Sheatfish 19 dm. long (the original of our figure)
the longitudinal diameter of the eyes was 5 % of the
length of the head, only slightly more than 9 % of the

a Kessler, Bull. Soc. Natur. Mosc., tom. XXIX (1856), I, p. 350.

b On the lOlli of August, 1870, a Sheatfish 12 Sw. ft. (3 '/3 m.) long is said to have been brought to market at Eskilstuna.
c In gravid females, or when the stomach is distended with food, the belly of course becomes both deeper and broader.

SHEATFISH.

695

breadth of the interorbital space, and also only a little
more than 14 % of the length of the snout. As appears
from these measurements, the breadth of the low, but
rounded interorbital space measures about l/2 (varying,
however, between 55 and 48 %) of the length of the
head, and the length of the snout usually a little more
than V3 (varying between 38 and 33 %) of the same
length. Here, however, as usual, the length of the snout
is measured obliquely (from the anterior margin of the
eyes to the middle of the tip of the snout): its real
length (the distance between its tip and the middle of
the line joining the anterior margins of the eyes) is
in young specimens rather more than 1J2, in specimens
1 in. long rather less than 1/s, and in our largest spe-
cimen about a/5 of the breadth of the interorbital space.
In proportion to the entire length of the head the real
leno’th of the snout decreases, during the growth of the
fish from a size of 3 dm. to one of 2 in., from 1/4 to
only J/ 5 of the said length of the head, and at the same
time from about 41 to 28 % of the postorbital length
of the head. The eyes thus lie invariably in the fore-
most third of the head, just behind the line between the
hind margins of the corners of the mouth. Just in front
of this line, or exactly in it, lie the posterior pair of
nostrils. These nostrils are small and round, and are
situated so far inwards that the distance between them,
which is relatively less in old specimens, measures only
about 66 — 56 % of the breadth of the interorbital space.
The anterior pair of nostrils lie far out on the snout,
in the line between the maxillary barbels, and at a
somewhat -smaller distance from each other. The margin
of all the nostrils is tubular, the posterior pair being
more elevated in a funnel-shape.

Of each of the maxillary bones there remains, as
Ave have indicated above, hardly more than the articular
knob itself, a small, triangular or scaphoid bone, holloAv
on the outside and articulating on the inside, which is
button-shaped, with the lateral ethmoid (prefrontal) bone
and the intermaxillary bone. But on the lower anterior
side of this bone Ave find another, still smaller, supple-
mentary bone, forming the base of the maxillary barbel,
Avhich is long and narroAV, but compressed, and may be
moArecl in all directions. The length of this barbel is,
hoAvever, extremely variable. Noav and then the barbel
on one side of the body differs from that on the other.
As a rule too, these barbels are comparatively shorter
in old specimens than in young: in a specimen 1 metre
long the longest barbel measured someAvhat more than

7i0 °f the length of the body, in a specimen 2 metres
long not quite l/6 of the same length or, in other Avords,
a little more than the length of the head. The tAvo
pairs of barbels on the loAver jaw are considerably shorter,
the anterior pair being also shorter than the posterior,
which generally extend to about a line Avith the hind
margin of the preoperculum or with the articulation of
the operculum. These barbels are not set on the loAver
jaAv itself, but a little farther inwards, on the skin be-
tAveen the branches of the jaw, the anterior pair nearly
beloAv the anterior nostrils, the posterior pair beloAv
the anterior margins of the eyes.

The surface of the head is smooth, and the bones
and cavities Avhich othenvise give many Glanomorphs
one or more important characters, are perceptible only
to the touch. Within this series of families Ave rather
constantly find, in the forms that have a more or less
fully armoured head, a longitudinal groove, or an oblong,
naked patch, along the middle of the forehead and snout.
Here, as in the other forms the head of which is des-
titute of dermal ossifications, no groove of this kind is
externally perceptible; but in the skull Ave find a long
opening (fontanel) between the anterior parts of the
frontal bones proper, and this opening is continued
betAreen the posterior parts of the ethmoid bone.

In the same Avay Ave find no external trace of the
suborbital ring; but in the skeleton it displays some
striking peculiarities. The foremost bone in this ring
(the preorbital bone) is comparatively small, flat, tri-
angular, and acute-angled in front. One side of this
bone lies on the outer top of the lateral ethmoid bone,
and the point in front is united by ligaments to the
top of the long lateral process of the ethmoid bone
proper. With its inside this bone forms the outer mar-
gin of the nasal cavity, the inner margin of Avhich is
formed by the long, but also fairly broad nasal bone Avith
its someAvhat crescent-like shape. Both the nasal bone
and the preorbital bone are furnished on their upper
surface Avith hollows and grooves for the ramifications
(muciferous ducts) of the lateral line. The second sub-
orbital bone forms the anterior margin of the orbit and
occupies a singular position, straight outwards and down-
Avards from the lateral ethmoid bone, Avhile its length
is so great that the tip lies above the loAver jaw, though
Avithout coalescing Avith the latter, just in front of the
articulation thereof. At the distal part of this sub-
orbital bone, Avith its anterior side united to the pos-
terior side thereof, lies the third suborbital bone, Avhich

Scandinavian Fishes.

88

696

SCANDINAVIAN FISHES.

is angular, and forms with the shorter (inner) arm a
part of the outer margin of the orbit. These two sub-
orbital bones thus form with their outer parts an osseous
ridge projecting from the orbital ring, a structure re-
minding us of that which in the Cottoids and the Lump
Suckers unites the orbital ring to the preoperculum.
The fourth • suborbital bone, which is the longest, is
curved in an S-shape. With its lower curve it forms
a part of the lower (outer) and the whole of the pos-
terior margin of the orbit. With its upper curve it
bends backwards to be united by ligaments to the out-
side of the anterior extremity of the posterior frontal
bone, which at this point, outside the frontal bone pro-
per, meets the hind extremity of the lateral ethmoid
(prefrontal) bone.

The crescent-shaped, obliquely set preopercula are
also externally indistinct, being entirely covered by the
skin; but a distinct dermal fold on each side marks the
position of the margins of the operculum and interoper-
culum. The former is triangular, with the upper margin
convex and the lower posterior margin slightly concave.
The interoperculum too, which also replaces the suboper-
culum, is triangular in form, but forked behind, with
the upper posterior corner projecting obliquely under the
operculum. The anterior extremity of the interoper-
culum is united by a long and strong ligament to the
hind extremity (angular part) of the lower jaw; but this
ligament runs along the inside of the interoperculum as
Avell, and is attached here and also, no less firmly, to
the outside of the upper part of the epihyoid bone®.
Behind the said dermal fold marking the hind mar-
gin of the opercular apparatus proper, the head is con-
tinued on each side by a broad branch iostegal membrane,
which lies in a rather indistinct flap above the upper
angle of the base of the pectoral fin. Both of the
branchiostegal membranes are entirely free from the
isthmus and united to each other only at the extreme
front for a very short distance, nearly below the corners
of the mouth; but the left branchiostegal membrane
overlaps the right under the isthmus to a considerable
extent. The branchiostegal rays are numerous; 13 or
14 of them are set on the ceratohyoid bone and are

fairly uniform in thickness and terete, but narrow and
pointed. The last two of these rays are set on the epi-
hyoid bone, and the last in particular is flat towards
the tip, externally grooved (hollowed in a longitudinal
direction), and fairly broad.

The cards of teeth in the lower jaw, on the inter-
maxillary bones, and on the head of the vomer are of
similar form and parallel, curved like a crescent and of
fairly uniform breadth, but growing narrower as usual
behind (distally on the vomer). They are composed of
dense, pointed, and somewhat recurved teeth of uniform
size, which are comparatively small, but in so large a
fish still form a powerful weapon. In the lower jaw
and on the intermaxillary bones the halves of the cards
are separate in front; but the vomerine card of teeth
presents an unbroken surface6. Of the three upper pha-
ryngeals on each side only the hindmost is furnished
with teeth, and this bone is also the only one of the
three that is visible in the pharynx, having extended
under the other two, which are small and serve to sup-
port it and to attach it to the under surface of the skull.
The patch of teeth on the first-mentioned pharyngeal is
fairly large, convex and elliptical. Each of the two
lower pharyngeals is furnished with an oblong patch of
teeth, broader in front, narrower behind (above), and
concave on the insideb All the pharyngeal teeth are
pointed and resemble the jaw-teeth, but are smaller,
smallest on the lower pharyngeals and in the outer pos-
terior corner of the upper cards. The gill-rakers are
scattered and fairly short, but pointed like teeth. On
the first and second branchial arches they are set in a
single row, and on the front of the first arch we find
9 or 10 on the ceratobranchial bone and 2 on the lower
part of the epibranchial bone. In some cases the lower
pharyngeals are also furnished on their outer margin
with a row of short, similar spines. The palatal folds
within the cards of jaw-teeth are well-developed, but
the upper is quite shallow. The upper jaw is not ca-
pable of the slightest protrusion, for the intermaxillary
bones are firmly united by ligaments not only to the
large anterior process of the ethmoid bone proper, but
also, though at a somewhat greater distance, to the

a Cf. above, p. 664, note A, on the singular position of the interoperculum in the Syngnathince.

5 In a Sheatfish 2 metres long the patch of teeth on each half of the lower jaw is 29 mm. broad in front and 118 mm. long, though
the posterior (outer) part for a distance of 40 mm. is considerably narrower than the anterior part; the patch on each intermaxillary bone is
85 mm. long and 22 mm. broad, a breadth also attained by the vomerine card of teeth, which measured across the palate is 96 mm. in length.

c In a Sheatfish 2 metres long each of the cards of teeth on the upper pharyngeals is 47 mm. long and 30 mm. broad, on the lower
pharyngeals 85 mm. long and 24 mm. broad at the broadest point. In a Sheatfish 1 metre long each of the two cards of teeth on the upper
pharyngeals is 30 mm. long and 21 mm. broad, on the lower pharyngeals 47 mm. long and 14 mm. broad at the broadest point.

SHEATFISH.

697

lateral ethmoid bones. The tongue is wanting. The
urohyoid bone is of a. singular form. Its anterior, shor-
ter part (corpus) resembles a parallelopiped. Its posterior
part is expanded into a rhombic, horizontal disk, with
the two anterior sides curved upwards and the two
posterior sharp, but deeply concave. Above this disk,
and at right angles to it, rises a strong, elevated ridge,
from the extreme beginning of the bone.

The form of the dorsal tin is especially characteris-
tic of the Sheatfish and reminds us strongly of the
adipose tin in our Salmonoids, resembling an obliquely
linguiform flap on a fairly narrow stalk, snapped, as
it were, and bent backwards at the middle. The fin
lies at a distance from the tip of the snout that mea-
sures in young specimens about 27 %, in old about 23
% of the length of the body", or in the former about
69 % , in the latter about 63 % of the distance between
the anal fin and the tip of the snout. Its four inter-
spinal bones coalesce into one supporting bone, the lower
extremity of which is attached between the tops of the
fifth and sixth neural spines (belonging to the sixth
and seventh vertebrae). The first ray is simple but
distinctly articulated, the posterior two6 or three rays
are branched and articulated.

The anal fin, the length (base) of which usually
measures more than half (about 53 or 54 %c) of the
length of the body, is of almost uniform height through-
out its length, though its height increases behind in the
same proportion as the depth of the body decreases.
Its greatest height (longest ray) is as a rule about 1/1G
— 7,3 of the length of the body, but may sometimes
rise to about l/a thereof. The fin begins just behind
the vent with its fairly large genital papilla, at a di-
stance from the tip of the snout that measures in young
specimens about 40 — 43 %, in old about 36 % of the
length of the body. Its last ray is united by a fin-
membrane to the lower margin of the caudal fin, as
far as the black colour extends over its base.

The caudal fin, which during life seems always to
be directed downwards, sometimes almost at a right
angle to the longitudinal axis of the bodyd, may, how-

ever, be raised and brought without violence into the
position which it occupies in our figure. In this posi-
tion, however, the skin forms a protuberance at the
dorsal margin just in front of the fin. The length of
this fin generally measures in young specimens about
11%, in old about 9 — 872 % of that of the body; but
the variations in this respect are quite as great as in
the case of the height of the anal fin, and in a speci-
men 1 metre long we have found the length of the
longest caudal rays to be 1472 % of that of the body.

The pectoral and ventral fins are the only ones in
whose various relations we apparently find perceptible1
external differences between the sexes. These fins are
relatively larger in the males than in the females, to
judge by our measurements of two males respectively
about 1 and 2 metres long and two females respectively
57 7 2 and 737 2 cm. long, our only specimens in which
the viscera were sufficiently well preserved to enable
us to decide the sex with certainty.

The pectoral fins are broad, but obliquely oval.
Their insertion is partly covered at the upper angle by
the broad branchiostegal membrane. Their first- (upper-
most) ray is strong and spinous, though articulated at
the tip, and forms a weapon which in old specimens
develops a number of spines on the inside of the outer
(distal) part. When this weapon is employed, the spi-
nous ray is erected in an outward direction, a creaking
sound is heard, a sound which the fish probably uses
to inspire terror, and the articulation is locked, so that
the ray cannot be forced back. This is a faculty espe-
cially common among the fishes of this family, and the
manner in which this result is attained is as follows.
The base of the spinous ray is not only furnished as
usual with two articular knobs — one for each of the two
articular surfaces which lie on the hind margin of the
firm osseous connexion between the scapular and cora-
coid parts of the shoulder-girdle, and of which the outer
(the one nearer to the clavicle) is an obliquely-set and cir-
cularly hollowed groove — it is also furnished on the out-
side with a tumid, pulley-shaped, articular knob, which
fits into a corresponding ear-shaped groove on the in-

“ To judge by these changes of growth, as well as by the vomerine cards of teeth, which in the Indian Silurus ivynaadensis (Day,
Fish. Ind., p. 480, pi. CXI, fig. 6) are separated from each other, this Indian form seems to stand nearer the original type common to both
species, though we must not forget that the Indian species is known only in specimens at most 12 in. (305 mm.) long. Besides, external
differences of sex are unknown in these species.

b In old specimens we have only found two branched rays in the dorsal fin.
c According to Kroyeu, however, sometimes (in young specimens) 49 or 50 %.

d This downward curvature of the caudal fin and its union to the anal fin are given by Sagemehl (Morphol. Jalnb., X 1884), p. 5)
as one of the proofs of the close relationship between the Gymnotoids (the so-called Electric Eels) and the Siluroids.

698

SCANDINAVIAN FISHES.

side of the angle of the clavicle. When the spinous
ray is erected, it is simultaneously twisted in a forward
and downward direction — the result of the above-
mentioned oblique position of one of the two articular
surfaces on the scapulo-coracoid bone — and during this
twisting process the last-mentioned, pulley-shaped knob
rubs against the wall of the clavicular groove — thus
producing the creaking noise — until this articular knob
is so firmly wedged in the lower end of the groove that
the ray cannot be bent without being twisted back at
the same time. Another consequence of the erection of
the spinous ray is that the insertion of the fin, which
previously occupied an oblique position, in a backward
and downward direction, now becomes horizontal. The
other rays of the pectoral fins are repeatedly branched,
the first being sometimes twice as long as the spinous
ray, the first (uppermost) three or four gradually in-
creasing in length, the lower ones from this point very
gradually decreasing in length down to the last two or
three, which grow rapidly shorter, one after another.
The last ray is not even half so long as the spinous ray.
The entire length of the fins, from the upper angle of
the insertion to the tip of the longest ray, is in our
males more, in our females less than 2/3 of the length
of the head".

The ventral tins are of a broad oval shape. Their
first ray is simple but short. They are inserted at about
the end of the first third of the body, so far back that
the distance between the dorsal fin and the tip of the
snout measures only about 75 — 70 % (in young speci-
mens 80 %) of the distance between the ventral fins and
the same point. Their length is in the males more, in
the females less than 2/5 of the length of the head* 6.
The pelvic bones are of a shafted spade-shape, with the
outer posterior margin forming an arc. They meet each
other partly in a fairly firm suture between the pos-
terior (extended) parts, partly in a ligamentous con-
nexion between the anterior ends.

The lateral line follows a straight course along the
middle of the sides of the tail (sometimes a little higher,
sometimes lower), but in front it rises in a greatly elong-
ated arch up to the temporal region, and may often
be traced distinctly in a straight line across the cheek
below the eye, as shown in our figure. The slimy

a In the former 72 — 81 %, in the latter 54 — 55 % of the li

6 ii ii ii 48 53 %, ,, ,, ,, 34 36 % ,, ,,

c In a male nearly 2 metres long the total length of the
pylorus and the bottom of the stomach about 1 dm.

surface of the skin, as we have mentioned above, is
entirely scaleless.

The coloration seems to vary considerably according
to local circumstances. The ground-colour is above
olive-green, below white. The back is always darker,
sometimes shading even into black. The sides are ir-
regularly spotted (mottled), in light specimens with
large and distinct spots of the dark colour of the back
on the ground-colour of the belly, which grows lighter
and lighter below. The dorsal fin is of the same dark
colour as the back, olive-greenish or black. The other
fins in common repeat the coloration of the body, the
base bearing the colour of the back, the margin or outer
part that of the belly. Such was at least the appearance
of the large Sheatfish, a male, from which our figure
is taken. It reached Stockholm alive, and was kept alive
while the artist painted it. But as its strength began
to fail, the lighter parts, especial])' of the pectoral, ven-
tral, and anal fins, became more and more suffused with
blood, and finally, when the fish was in a moribund
state, blood oozed in drops from the margins of these
fins. The maxillary barbels are on the upper side of
the same colour as the back, underneath paler or even
of the same colour as the belly. During life the outer
parts of the eyes were blackish blue, the inner parts
lighter blue, with the iris yellow but punctated with
dark dots and set off by a sharply-defined, yellow inner
margin from the pupil, which was black.

The length of the abdominal cavity measures in
young Sheatfish about 1/3, in older ones about 1/i of
that of the body. The peritoneum is white. The oeso-
phagus is continued straight backwards by the large and
muscular stomach, the pointed or rounded bottom of
which lies almost above the middle of the length of the
pelvic bones. Rather far forward, at about the middle
of the stomach and on its under surface, the pyloric
part projects in a forward direction. This part is bounded
from the intestine by a more or less strong contraction,
and internally divided from the same by a valve". The
duodenum, which is without appendages, is of about
the same thickness as the pyloric part, runs first for-
ward nearly to the diaphragm, then bends upwards to
the right of the oesophagus, and curves backwards, de-
creasing in thickness and thus passing imperceptibly

igth of the head.

n ?? n

esophagus and stomach is about 35 cm., and the distance between the

SHEATFISH.

699

into the small intestine, often in several small curves, above
and behind the stomach. Here the intestine again bends
forwards, below and to the left of the stomach, to about
a line with the pylorus, then returns in a sharp curve,
and proceeds to the vent. The liver is large, though
its length varies: the left lobe is the longer, extends to
about a line with the bottom of the stomach, sometimes
not quite so far, sometimes much farther, and is divided
at the tip into two secondary lobes of varying length “.
The gall-bladder, which lies between the duodenum and
the anterior part of the left lobe of the liver is saccate
and large6. The pancreas0 lies nearer the duodenum,
in the shape of a horse-shoe with the prongs directed
forwards and surrounding the gall-duct. The spleen is
situated above the stomach, between this organ and the
air-bladder, to the left of the first coil of the intestine
and sometimes entirely in front of, sometimes half in
front of and half behind and above the anterior angle
of the last coil of the intestine. It is rounded (ellip-
tical) and flattened'6. The structure of the large air-
bladder is mentioned above0. The under surface of its
posterior division is furnished with a longitudinal groove,
at the anterior end of which the pneumatic duct de-
scends into the oesophagus. The anterior end of the air-
bladder is attached partly to the lower surface of the
tip of the foremost, transverse processes (belonging to
the second abdominal vertebra), partly to the anterior
part of the side of the large vertebra which, as we have
mentioned above, is produced by the coalescence of the
original second, third, and fourth vertebrae. Within the
latter fastening of the air-bladder lies the descending
blade of the hindmost and largest, so-called acoustic
bone ( malleus ), which is loosely folded in beside the
large, composite vertebra. The second blade of this
acoustic bone lies horizontally, pointing in a forward
direction, at right angles to the descending blade, and
glides along the under surface of the base of the first
transverse process. This acoustic bone is thus both bent

and twisted; and within the angle of the bend the base
of the descending blade forms a free margin, which, when
the bone is in its natural position, lies as a continuation
of the upper margin of a lateral groove on the body of
the vertebra. As only the third (hindmost) of the coa-
lescent vertebrae is furnished with ribs, the so-called
malleus thus corresponds to a rib of the middle ver-
tebra (the third abdominal vertebra). The malleus co-
heres at its inner anterior corner with the vertical, flat
incusf, which should thus correspond (if this homology
with the ribs should receive the corroboration it still
requires from the history of development) to a rib
of the foremost of the coalescent. vertebrae (the second
abdominal vertebra). The first abdominal vertebra, which
in the Sheatfish is separated both from the following
vertebrae and the occipital bone, is without either trans-
verse process or neural arch, unless these be represented
on each side by the angular stapes , which is united by
ligaments to the subjacent incus, and by the flat,
but oblong claustrum, which lies above this point, is
also united to the stapes, and covers the atrium sinus
imparls on this side of the body. This atrium , which
lies beside the spinal cord, is a lateral extension of the
sinus impar situated in the occipital bone, a mem-
branous, saccate extension of the cerebral membrane
that lines the labyrinth. The anterior end of this sinus
is furnished in the Carp-fishes5’ — and probably here as
well6 — with a connecting duct between the sacculi of
the labyrinth. The air-bladder is thus connected by
means of the acoustic bones, not indeed immediately
with the true hearing-apparatus, but with the lymphatic,
subdural chamber that surrounds the latter. Modern
scientists have therefore adopted the opinion1 first main-
tained by Hasse;, and regard this connexion less as a
conductor of sound than as a barometrical apparatus
which conveys to the consciousness of these fishes a sense
of the varying atmospheric pressure to which their air-
bladder is exposed at different moments.

“ In the above-mentioned specimen the length of the left lobe of the liver is 36'/2 cm. and its greatest breadth nearly 8 cm. The
length of the right lobe is 14x/2 cm., its breadth nearly 5 cm.

* In the specimen just mentioned the gall-bladder is about 14 cm. long and, when collapsed, about 61/, cm. broad.

c The length of this gland from the middle of the round posterior margin to the end of one of the prongs is (in the same speci-

men) about equal to the breadth of the gall-bladder.

d In the same specimen the spleen is 11 cm. long, 5 cm. broad, and about 8Y2 mm. thick.

e In the same specimen the air-bladder is 25 1/9 cm. long and rather more than 9 cm. broad. In a female 575 mm. long it is 89

mm. long and 46 mm. broad.

f Cf. Weber, De Aure et Auditu Hominis et Animalium , tab. V, fig. 30.
g Cf. Nusbaum, Zoologischer Anzeiger, 1881, p. 553.

h Cf. Retzius, Das Gehororgan der Wirbelthiere, I, p. 77, taf. XIII, fig. 1, si.

1 Cf. Sagemehl, Morphol. Jahrbuch, X (1884), p. 14.
j Anatomiscbe Studien , No. XIV, p. 596.

700

SCANDINAVIAN FISHES.

The ovaries are fusiform or cylindrical sacs with
fairly thick walls; the testes are flat, lobate, and thin-
walled. The kidneys lie as usual between the air-
bladder and the spinal column, forming an anterior
division and a larger, posterior one, beside and behind
the air-bladder. The urinary bladder is large, and is
most developed on the right side.

The principal habitat of the Sheatfish lies in the
south-east of Europe, the species being commonest in
Russia and Austria. It is there too, in the Caspian and
Black Seas and the rivers that flow into these waters,
that it attains its maximum size. According to Pallas
and Grimm the Sheatfish occurs throughout European
Russia, with the exception of the basin of the White Sea
and its rivers. In the Danube too, especially towards
the mouth, the species is plentiful and of great size. In
the Greek Peninsula it is common, according to Apo-
stolides, in the River Peneus, off Larissa, and in Li-
vadia. In Italy and the Iberian Peninsula it is wanting,
and also in France and Belgium, except in the River
Doubs in the extreme east of France, where it has been
caught occasionally near the town of Dole. In Switzer-
land its occurrence is confined to the Lakes of Constance
(the basin of the Rhine) and Morat (Murten), a small
lake east of Neuchatel, where it lives, strange enough,
outside its strict geographical range, which lies on the
other side of the Rhine. In England, as well as in
Scotland and Ireland, it is wanting, though repeated
attempts have been made to plant it there. Before
Haarlem Meer was drained (1836 — 53), the Sheatfish,
according to Gronovius, was very common in this lake,
which was chiefly formed, however, by an inroad of the
sea in the fifteenth century. It is still found in several of
the small lakes of Holland. In the rivers and lakes of
Germany the Sheatfish is widely spread, and also in the
Baltic Provinces of Russia, according to Seidlitz. Nor
does it shun the brackish waters of the North, for it
occurs, according to Benecke, in the Haffs of Northern
Germany. In Finland, according to Malmgren, it is
found only in the lakes near Tavastehus and is very
seldom caught, though, according to Gadd", it attains
so great a size “that a yoke of oxen are required to
move it from the spot". Nor is the Sheatfish a common
species in Sweden, though it occurs at many scattered
spots in the south-east of the country, principally in
three separate districts: one to the north, including

Lakes Malar, Hjelmar, Borsjo, and Bafven in Soderman-
land and Lake Hunn in Ostergothland ; one more to the
east, the neighbourhood of Oscarshamn, where it is known
from Lakes Humel, Nejer, Versjo, Tvinger, Storutter,
Greater and Lesser Ramsjo, Goten, Maren6, Bodasjo (Fli-
seryd), and the River Emm; and one to the south, the
neighbourhood of Christianstad, where it is found in
Lakes Immel, Ifosjo, Ousby, and the River Helge. Nils-
son also quotes a doubtful newspaper statement of the
occurrence of the Sheatfish in Lake Bolmen, in Smaland.
In Denmark the Sheatfish was common at the end of
the last century, according to Holm, in Lake Soro, where
it had probably been introduced from Germany; but it is
now exterminated there. It has also been taken once in
a river near Kjoge, into which Kroyer supposed it had
wandered after some roving excursion in the Baltic.
In Norway the Sheatfish has never been found.

The Sheatfish is a sluggish but voracious fish-of-
prey. Its very appearance is repulsive. Slimy and slip-
pery as an Eel, with its broad gape, small, blinking
eyes, and long, warily plied barbels, it looks as though
fashioned especially to lie in wait for the destruction of
others. Its size too has rendered it an object of dread
even to man. An old Bohemian proverb says, “One
fish is another’s prey, but the Sheatfish eats them all;”
and we have numerous accounts of the Sheatfish attack-
ing domestic animals and children. Hidden among the
rushes or in the mud, the tints of which are reproduced
in its coloration, it lies motionless the greater part of
its time, only waving its barbels to and fro, until some
victim approaches so near that only a sudden movement
is requisite to seize and devour its prey. Or one of
its senses tells it that a dainty morsel, some decom-
posing body or baited hook, is not far off, and it wrig-
gles thither. During a part of the year, the cold sea-
son, it retires to deeper water; in spring and summer,
til! the spawning is over, it keeps to shallower spots
and the shore.

The Sheatfish generally leads a solitary life, though
not to such a degree that it does not seek company.
One of the authors who have most contributed to our
knowledge of its habits, the Dane Holm, who published
in 1779 his observations of the Sheatfish in Lake Soro
in Zealand, even tells us: “The Sheatfish is fond of
company and is therefore seldom found alone; but only
three or four, and these always of about the same

a Abo Tidningar, 1 772, p. 366.

b From these nine lakes the observations were collected in 1890 by Mr. E. Svedjviark the geologist.

SHEATFISII.

701

size, have ever been seen together at Soro. Of them
we are told that they pursue and drive their flying
victims in common, until, sure of their prey, with one
consent and with open mouth they dart about with in-
credible speed, seizing and devouring their victims. In
order to attain this great velocity the Sheatfish chiefly
employs its tail, by whose serpentine movements and
quick blows it sculls itself along like the Eel. Fortu-
nately for the fishes it pursues, the Sheatfish by its
violent movements and rapid pace sets the water in
commotion and thus betrays itself.” When the Sheatfish,
on the other hand, lies lazily at rest and allows small
fishes, frogs, crustaceans, insects, and even worms to
glide down its maw, we may well conjecture that here
too (cf. above, pp. 263 — 264) the inner transverse cur-
tains of the mouth act as valves, so to speak, in pump-
ing the tiny victims into its gape. However, the
Sheatfish is not exclusively carnivorous; vegetable sub-
stances also enter into its diet.

From older times we have accounts of assaults
made by the Sheatfish upon higher animals as well, even
upon man, or at least of its devouring the bodies of
drowned persons. Gesner tells us that the stomach of
a Sheatfish was found to contain a human head and a
right hand with two gold rings, and that geese, ducks,
and animals that were being watered, have been seized
by the same fish. Pallas says of the Sheatfish in Russia
that it is so voracious that, it does not fear to seize
bathers by the legs. Valenciennes quotes several accounts
from Hungary of its attacking children. Heckel and
Kner state that at Vienna on one occasion a poodle was
found in the stomach of a Sheatfish, and that at Press-
burg a similar discovery was made of the remains of a
boy. E. Svedmark, the geologist, who has collected the
instances of the occurrence of the Sheatfish in the Di-
strict of Calmar, was told there that “long ago” a person
had stood on the shore of Fake Versjo and watched a
Sheatfish seize a lamb. The Sheatfish is therefore feared,
this being also due in all probability, as Heckel and
Kner have remarked, to the bitter experience that a
few may have had of the obstinate wounds caused by
the spines on the first pectoral ray of the Sheatfish.
Specimens of large size may also prove no easy booty
to their captors. “On Lake Storutter,” writes Svedmark,
“a fisherman speared in the back a monster that drew
the punt all round the lake before it finally surren-

dered.” Lilljeborg tells a similar tale of a Sheatfish
that was caught on a longline in Lake Immel.

The Sheatfish is eminently sensitive to changes in
the weather, especially to thunder, when it may often
be seen at the surface, as well as at other times on
warm summer daj^s with drizzling rain (Holm). In many
places too the Sheatfish passes for a weather-prophet.
“When it breathes air,” says the fisherman — and this is
a necessity to most fishes, but above all to the Physo-
stomes — “it raises itself out of the water like a big,
black man” (Svedmark). It is extremely tenacious of
life. A male 19 dm. long was sent by rail on the 30th
of May, 1889, in a Avooden box containing straAv and
a little ice, from Lake Bafven to Stockholm. It Avas still
living on its arrival, and was kept alive for several days in
a vessel Avhere the water Avas just sufficient, to cover it.

The spawning-season occurs in the summer (May —
July), generally about midsummer. “At this season,”
says Holm, “the Sheatfish is sluggish and lethargic, being
easy to surprise and capture. This applies in particular
to the females, which for some time previous to the
spaAvning lie* among the rushes, Avhere, it is said, the
female deposits her roe, after first providing a secure
and convenient receptacle for it by scooping Avith her
tail a hole in which the young subsequently stay for
some time.” The ancient account of Aristotle, that
the male watches the roe, is now regarded as dubious.
The number of eggs varies as usual according to the
size of the fish. In a female 1'87 kgm. in Aveight Be-
necke counted more than 60,000 eggs. The eggs are
slightly yelloAv, 3 mm. in diameter, and are hatched in
8 — 14 days. The fry grow quickly, Avhere food is abun-
dant, attaining in the first, year, according to Brehm°,
a weight of 3/4 kgm., in the second a weight of 1 ’/2
kgm. A fisherman of Strasburg, Baldner by name,
Avho has left a manuscript written in 1666 and con-
taining an account of all the birds, fishes, and other
animals he had taken, tells us that, a Sheatfish caught
in the. River 111 near Strasburg Avas kept in captivity
from 1569 to 1620 and during this period attained a
length of about lV2 metres.

Young Sheatfish, less than 1 metre in length, are
quite eatable, though not exactly easy of digestion. The
tail is the best part, of the body. The flesh is soft,
white, and rather sweetish in taste. Valenciennes com-
pared its flavour most nearly to that of the Eel, but

Thierleben , Abtli. Ill, Bd. 2, p. 201.

702

SCANDINAVIAN FISHES.

found it much inferior to the latter. The flesh of older
specimens is tough and rank. “As the Sheatflsh has ;
no scales,” says Holm, “the Jews are not allowed to eat
it — a privation which, in my opinion, they need not
regret. To render the flesh eatable, the fat must first
be boiled away, and the tish then boiled again in wine
and with plenty of spices to remove its nauseous flavour.”
In Sweden, however, the Sheatflsh is so seldom caught
that it is of little economical importance. To the fish-
mongers of Stockholm it seems to be almost unknown.
In Russia the case is different. From Astrachan for
example, according to Grimm, about 1 2/3 million kilo-
grammes of salted Sheatflsh are annually exported. In
1881 three-fifths of this quantity was taken in the River

Koora and one-fifth in the Black Sea. The total annual
catch of Sheatflsh in the Black and Caspian Seas, in-
cluding the nearest parts of the Russian rivers that flow
into them, is estimated by Grimm at 41/ 5 million kgm.
of flesh and 16,000 kgm. of fish-glue, the latter manu-
factured from the air-bladder. This is a tough glue, but
inferior to (more impure than) genuine isinglass. The
skin of the Sheatflsh is used by the Russians and Tar-
tars to glaze their windows; and they boil down the
fat into lamp-oil (Pallas).

The usefulness of the Sheatflsh is counterbalanced
by the damage caused by its voracity, and Day con-
gratulates his country on the failure of the attempts to
introduce the Sheatflsh into England.

CYPMNOMORPHI.

Physostomes with the shoulder-girdle (as usual among the Teleosts ) suspended to the head. Coracoscapular disk
strengthened internally by an arch formed by a special bone (os prcecoracoideum) from the coracoid bone to the
clavicle or the upper (anterior) margin of the scapula. The first four abdominal vertebrce deviating more or less
distinctly from the normal vertebral form , and partly metamorphosed into so-called acoustic bones. ITyomandibular
and ptery go-palatine arches as well as the opercular apparatus complete. Maxillary bones fully developed. No
jaw-teeth. Body naked or covered with cycloid scales. Head generally naked , at least on the sides and snout".

This series of families was established, as Ave have
mentioned above, by Cope under the name of Plecto-
spondyli , but according to his definition thereof is far
less natural than the preceding series. On the one hand
it includes in his Avorks the so-called Salmon-Carps
(Characinidce), inhabitants of the tropical regions of
Africa and America, with the posterior part of the edge
of the mouth formed, as in the Salmonidce , by the ma-
xillary bones and, in the great majority of the genera,
Avith an adipose fin on the posterior part of the dorsal
margin, as in the Salmons. On the other hand the series
excludes the so-called Toothed Carps (Cyprinodontidce),
a tropical and subtropical Carp-like family Avith range
extending round the globe, destitute, it is true, of the
osseous connexion between the air-bladder and the sac-
cate extensions of the cranial cavity and also Avithout
praccoracoid bone, but with the edge of the mouth
formed, as in the Carp-fishes, by the intermaxillaries
alone, though these bones, in contradistinction to those
of the Carp-fishes, are furnished Avith teeth. If we add

to this that the Salmon-Carps, like the Salmons, have
Avell-developed pyloric appendages, Avhile the Toothed
Carps, like the Carp-fishes, are destitute thereof, Ave
have sufficient anatomical grounds for breaking up the
series Plectospondyli. The series then acquires, with the
above characters, the same limitation as Gunther’s fa-
mily Cyprinidce , its members being restricted to the
chiefly Asiatic Loach-fishes (Cobitidce) and Stone Carps
(Homaloptericlce), the Carp-fishes (Cyprinidce) of the
Old World and North America, and the Suckers (Cato-
stomidce), chiefly from North America. All these fishes
have the edge of the mouth formed, at least for the
most part, by the intermaxillary bones, Avith the maxil-
laries lying above and behind them, and all are Avith-
out jaAV-teeth. Only tA\ro of these families are re-
presented in the Scandinavian fauna.

A: Air-bladder entirely or partly enclosed in

an osseous capsule; at least six barbels... Fam. Cobitidce.
B: Air-bladder without osseous capsule; at

most four (or no) barbels Fam. Cyprividce.

a The genera Lepidocephcdus and Lepidocephalichthys , Cobitidce from India and Further India, are furnished, however, with scales on
the sides of the head. See Bleeker, All. Ichth. hid. Or., tome III, p. 12 and Day, Fish. Ind ., p. 601*.

LOACHES.

703

Fam. cobitid^e.

Air-bladder more or less reduced , entirely or partly enclosed in an osseous capsule. Mouth fringed with, six or

more barbels. Pseudobranchice wanting.

This family was established under the name given
above by Bonaparte" in 1846, but subsequently re-
ceived of Heckel and Inner/' the name of Acanthopsi-
des, derived from the generic title which Agassiz had
supposed to be deserved by our most common species.
By most authors, however, the family has been regarded
as a subfamily among the Cyprinoids. Still it possesses
so many distinctive characters that it may well main-
tain its position in the system. The most prominent
among these characters are those which suggest an
alliance with the Glanomorph series. Among these we
find externally the slimy skin, generally naked, other-
wise with small scales, and the comparatively numerous
barbels round the mouth; while internally the capsule
of the air-bladder reminds us of the corresponding
structure in many of the Glanomorphs, and the ossifica-
tion of the head shows a fronto-parietal fontanel simi-
lar to that we have remarked above in the Siluroids.
The intestinal respiration of the Cobitoids may also
be in some degree a trace of their connexion with the
preceding series of forms, where we have seen the
respiration of air accomplished in various ways, and
where the respiration of water seems in certain fishes
to be assisted by an apparatus consisting of ramified
appendages in the anal region. In the present series,
on the other hand, it has long0 been known that under
certain circumstances, where there is scarcity of water
or of the oxygen contained therein, our European Co-
bitoids ascend to the surface and swallow air, which
they audibly eject after a time through the vent, with
the oxygen now changed to carbonic acid gas.

The Cobitoids are distinguished from the following
family not only by the capsulate air-bladder and the
greater number of the barbels, but also by the absence
of pseudobranchige — this is also the case in the great
majority of the Glanomorphs — and the comparatively
small size of the gill-openings, the branchiostegal mem-

branes being united underneath for the greater portion
of their length to the isthmus. The lower pharyngeals
are destitute of the strength which they possess in the
following family, are more like branchial arches, and
are armed with a row of weaker, but in general more
numerous teeth.

The air-bladder is partly free, it is true, in several
fishes of this family, a posterior division thereof lying
free within the abdominal cavity. But in others, and
in particular in all our European species, it is com-
pletely enclosed in a porous or retiform, osseous cap-
sule, or only a small, rounded expansion of the air-
bladdeW juts out at the hind orifice of the capsule,
where otherwise only the pneumatic duct descends to
the oesophagus. The osseous capsule of one of the
European species, Cobitis ( Misgurnus ) fossilis , is excel-
lently described and figured by Weber (1. c.). Its
morphological explanation consists in a tumidity and
transformation of the osseous structure originally be-
longing to the lower and inner parts of the transverse
processes of the third vertebra and to its pair of ribs.
The upper and outer (dorsal) part of this transverse
process on each side may still be distinguished in the
wall of the capsule; and in our Spined Loach ( Cobitis
taenia ;) the top of this process, as tvell as each of the
ribs behind it, bears a scleral (epipleural, muscular)
bone branched at the top. The osseous point that, pro-
jects downwards on each side of the capsule, also pre-
sents an unmistakable resemblance to the ends of the
ribs behind this point. The same explanation is given
of the hollow bone extending from each side of the
body of the second vertebra and from the lower part
of its neural arch, backwards, outwards, and down-
wards, above the anterior upper part of the surface of
the osseous capsule. Within this hollow bone are con-
tained the so-called acoustic bones. In the Spined Loach
this hollow bone — the outcome of the material of the

a Cat. Met. Peso. Europ. pp. 5 and 26.
b Siisswasserf. Oesterr. Mon ., p. 296.

c See Erman i Gilbert’s Annalen der Physik, Bd. XXX (1808), p. 140. Cf. also Siebold, Siisswasserf. Mittelenr., p. 340.
d See Weber, De Aure etc., tab. VI, fig. 48, sign. 8.

Scandinavian Fishes.

89

704

SCANDINAVIAN FISHES.

lateral parts of the second vertebra — bears on its top
a scleral bone, the second in order from in front: —
the first (sometimes double) lies on each of the lateral
occipital bones.

Such is the osseous capsule in Colitis fossilis and
C. tcenia, in form a transversely-set ellipsoid, entire, and
contracted only at the middle of its ventral side by a
shallow groove in the longitudinal direction of the body;
but in our second species, the Loach ( Colitis larlatula ),
this contraction has gone so far that the two lateral
parts of the osseous capsule are separated from each
other, though posteriorly united on the ventral side by
a hollow osseous bridge. In this species we also find,
strange to say, a well-developed pair of ribs on the
first vertebra.

On each outer (lateral) side of the osseous capsule
(or on the outside of its divisions in the Loach) we
find two holes, one of which leaves room for the union
of the hind extremity of the so-called malleus with the
air-bladder, while the second, posterior, and larger hole
admits into the osseous capsule, to the lymphatic cham-
ber surrounding the air-bladder®, a membranous duct
from the subcutaneous connective tissue. A communi-
cation ( introitus capsules vesicce ) is thus formed between
the air-bladder and the side of the body exactly above
the insertion of the pectoral fin and on a level with
the upper angle of the gill-opening* 6, where this duct
passes between the dorsal and lateral divisions of the
great lateral muscle. The latter communication has its
analogue, as we have seen above, both in Batistes and
the Glanomorphs; but here we have no evidence to
show that it is connected in any way with the pro-
duction of sound. On the other hand, the signification
which Hasse and Sagemehl have attributed to the con-
nexion between the air-bladder and the apparatus of
hearing, that of a barometrical apparatus, here finds
perhaps its strongest corroboration, for we know that
these fishes are so sensitive to changes of atmospheric
pressure that they have long been trusted as weather-

prophets. One species ( Misgurnus fossilis ) has received
in Germany the name of Wetter fisclC . “It has been
observed,” says Bloch d “that at the approach of storm
this fish rises from the bottom towards the surface and
is particularly restless in its movements. It may there-
fore be used instead of a weather-glass, if set in a glass
vessel containing river- water or rainwater and a little
rich soil at the bottom. Twenty-four hours before a
storm of wind or thunder it invariably grows restless,
disturbs the water till it becomes turbid, and rises and
sinks up and down in the vessel. In settled weather,
on the other hand, it usually lies still at the bottom.
The fish may be kept alive almost a whole year, if
the water and soil are changed only twice throughout
the summer and once a week during the winter. In
winter, however, it must be placed in a warm room
and at a window.”

The Cobitoid family consists exclusively of fresh-
water fishes, the great majority belonging to Asia and
the neighbouring islands. According to Bleeker the
number of species exceeds 80; Day knew 46 from In-
dia. The former distributes these species among 12
genera, Gunther c recognises 11. The three species that
occur in Europe belong, according to these opinions,
each to a distinct genus, the first ( Misgurnus ) with bar-
bels on both jaws — this character is possessed by Mis-
gurnus fossilis, an inhabitant of the regions south and
east of the Baltic — , the second genus ( NemacMlus )
without erectile spine below the eyes (like Misgurnus),
but without barbels on the lower jaw — this genus is
represented by the Loach ( NemacMlus larlatulus), which
has been observed in several localities in Scandinavia — ,
and the third ( Colitis ) containing our common Spined
Loach ( Colitis taenia ), without barbels on the lower
jaw, but with an erectile spine (at other times directed
backwards) below each eye. It has already been re-
marked, however, by Valenciennes7 that on a very
minute examination of Misgurnus fossilis we discover
a small crevice in the skin below each eye, and that

a Cf. Hasse, Anatomische Studien, Bd. 1, p. 595.

6 See Weber, ]. c., tab. VI, figg. 45 and 46.
c Heckel and Kner. Susswasserf. Oesterr. Mon., p. 300.
d Naturg. Fisch. Deiitschl., Th. I, p. 218.
e Introd. Stud. Fish., p. 604.

f Cuv., Val., Hist. Nat. Poiss., vol. XVIII, p. 6. In our specimens of Misgurnus fossilis we easily discover a longitudinal groove below

the eyes; but the crevice in the skin is impossible to detect, and the spine that may be felt at the lower anterior corner of the eye, lies

in front of the said groove. On dissection, however, it appeared that Weber ( De Aure etc., figs. 43 and 44) was perfectly correct in re-
presenting in his figure a preorbital spine in Misgurnus fossilis. This spine lies hidden among the muscles, but is movable as in Cobitis
taenia and has the same form, the analogy being so complete that the other spine, which is more easily felt externally, corresponds to the

lateral spine on the true preorbital spine of Cobitis taenia.

LOACHES.

705

in this crevice may be felt the tip of a spine, which
does not, however, admit of erection. Again, Day ob-
serves of the Indian species of the genus Nemachilus
that “sometimes the preorbital is raised and with a free
lower edge, while this may not occur in all examples
of the same species”. Several authors too, Moreau and
Lilljeborg for example, have abandoned the attempt to
distinguish the genera within this family exclusively by
the presence or absence of the preorbital spine. On the
other hand, it seems highly probable, to judge by our
knowledge of the remaining genera, and also when we
consider the analogous relations within the Glanomorph
series, that the presence or absence of barbels on the

lower jaw affords a constant and more useful generic
character, though in many species that are otherwise
without them, these barbels are indicated by contrac-
tions and interposed swellings on the underlip. Whether
we choose on this ground to retain the genus Misgurnus
for the European Misgurnus fossilis and a few other
species, or whether we regard this group as a subgenus
of the genus Colitis, a course for which we shall find
strong reasons below, is a matter of no great im-
portance in the Scandinavian fauna, for though Misgur-
nus fossilis has once been planted in Sweden — in ponds
at Ulriksdal, according to Linnaeus — it has not spread,
so far as is known, to any extent in this country.

Genus COBITIS.

Six barbels", none on the lower jaw. Body elongated, terete , or compressed. Head naked ( without scales). Dorsal
fin short and situated above the ventral fins. Caudal fin rounded , truncate, or slightly concave.

This genus comprises the great majority of the fa-
mily. Including the species that have previously been
referred to the genus Nemachilus, as being without
movable preorbital spine below the eyes — by far the
greater number belong to this class — 67 species have
been adopted and described by Gunther, Day, Herzen-
stein, and Bleeker. Day enumerates 31 species from
India. Herzenstein assigns 17 species to the highlands
of Central Asia (Tibet). From Syria and Palestine 7
species are known, described by Heckel and Gunther.
Bleeker cites only two species from Java and Sumatra,
Castelnau two from Cape Town* 6. Europe also pos-
sesses two species of the genus. It thus appears that
the genus thrives best in the rivers and brooks of the
Asiatic Highlands; and it is probably thence that it
has spread to the lowlands.

In one of our European species ( C . taenia) Ca-
nestrinP remarked in the structure of the pectoral

fins an external difference between the sexes, an ob-
servation which Herzenstein has subsequently verified
with one or two modifications, in the majority of the
species that inhabit Central Asia. In the males of these
species the second ray of the pectoral fins is more or
less thickened and broad, and furnished during the
spawning-season with tubercles or spines.

The members of this genus are in general of in-
significant sized, though, as Valenciennes has pointed
out, not to such an extent that we are justified in
assuming that it was to these fishes that Aristotle
referred when he included Kcofiirig among the Aphyce
(cf. above, p. 264). The genus Colitis of modern
ichthyology dates from Artedi.

In Scandinavia only two species are found:

1: Preorbital bone furnished with a spine... Colitis taenia.

2: No extensile spine below the eyes Colitis larlatula.

a In one Indian species there are 8 barbels; but all of them belong to the snout and the upper jaw.

6 These two species are recognised by Gunther, however, merely as doubtful.
c Fn. Ital., part. Ill, Pesci, p. 21.

d One species, Colitis yarkandensis from the Asiatic Highlands, attains, however, a length of at least 3 dm., and is thus only slightly
inferior in size to the European Misgurnus fossilis, which sometimes measures 3 1 / 2 dm.

706

SCANDINAVIAN FISHES.

THE SPINED LOACH (sw. nissogat).
COBITIS TtENIA.

Plate XXXI, fig. 4.

Body laterally compressed. Breadth ( thickness ) of the head at the eyes less than 2/5 of its length; breadth of
the interorbital space less than the diameter of the latter. Below each eye a movable preorbital spine

pointing in a backward direction.

R. br. 3 ;

(2) + 1 . (2) + l (1) + 1. T7 (1)+1.

7—8 ’ " 5 ’ ' 8 ’ ' 6" ’

C. «+l + 14 + l+«; Vert. 45 b.

Syn. Cobites aculeata, Rond., Pise., part. II, p. 204. Tcenia cor-
nuta, Schonev., Ichthyol. Slesv. Hols., p. 74. Cobitis , aculeo
bifurco infra utrumque oculum, Art., Gen. Pise., p. 2;
Syn. Pise., p. 3; Spec. Pise., p. 4.

Cobitis Tcenia, Lin., Syst. Nat., ed. X, torn. I, p. 303; Fn.
Suee., ed. II, p. 121; Bl., Naturg. Fisch. Deutschl., I, p.
221, tab. XXXI, fig. 2; Pall., Zoogr. Ross, ,4s., tom. Ill,
p. 166; Flmng ( Gobitis ), Brit. Anim., p. 189; Nilss. {Co-
bitis), Prodr. Ichth. Scand., p. 35; Jen., Man. Brit. Vert.,
p. 417; Agass. ( Acanthopsis ), Mem. Soc. Neuch., tom. I, p.
36; Yarr. {Botia ex Gray), Brit. Fish., ed. I, vol. I, p.
381; I\r., Damn. Fiske, vol. Ill, p. 564; Cuv., Val. (Cobitis),
Hist. Nat. Poiss., vol. XVIII, p. 58; Nilss., Skand. Fn.,
Fisk., p. 345; Kessl., Bull. Soc. Natural. Mosc., tom. XXIX
(1856), p. 352; Hckl, Kn., Susswasserf. Oesterr. Mon., p.
303; Costa, Fn. Regn. Nap., Pesci, Abdom. Malacott. Cypr.,
p. 31; Sieb., Siissiuasserf. Mitteleur., p. 338; Mgrn, Finl.
Fisk. (disp. Helsingf.), p. 37 ; Steind., Stzber. Akad. Wiss.
Wien, Math. Naturw. Cl. LIE, i (1865), p. 490; Blanch.,
Poiss. d. eaux. donees Fr., p. 285; Canestr., Arch., Zoo!.,
Anat., Fisiol., vol. IV, fasc. I, p. 146; Gthr, Cat. Brit.
Mus., Fish., vol. VII, p. 362; Canestr., Fn. Ital., part. Ill,
Pesci, p. 20; Cederste., Ofvers. Vet. Akad. Forh. 1874, No.
9, p. 47, tab. XI; Fedders. (Botia), Naturh. Tidskr. Kbhvn,
ser. 3, vol. XII, p. 92; Mor. (Cobitis), Hist. Nat. Poiss. Fr.,
tom. Ill, p. 434; Bncke, Fisch., Fischer., Fischz. O ., W.
Prenss., p. 147 ; Day, Fish. Gt. Brit., Irel., vol. II, p. 201,
tab. CXXXVII, fig. 3; Mela, Vert. Fenn., p. 314, tab. X;
Norback, Handl. Fiskev., Fiskafv., p. 444; Lillj., Sv., Norg.
Fisk., vol. Ill, p. 345; Fatio, Fn. Vert. Suisse, vol. IV,
tab. V, fig. 2; vol. V, p. 10.

Cobitis caspia, Eichw., Bull. Soc. Nat. Mosc. 1838, II, p. 133.
Cobitis tcenia japonica, Schl., Fn. Japon., p. 222, tab. CIII,
figg. 3 et 3. a.

Cobitis elongata , Hckl, Kn., 1. c., p. 305.

Cobitis larvata, De Fil., Mem. Accad. Torin., XIX, p. LXXI,
vide Canestr. Arch. Zool., Anat., Fisiol., vol. IV, fasc. I,
p. 150, tav. VI, fig. 7.

The Spinecl Loach is in general the smallest of the
European Cobitoids, its ordinary length being between
1/2 and 1 dm., though according to Moreau it may
sometimes attain a length of 12 cm. From Krain
(Austria), however, Heckel and Kner have described
examples of this species, under the name of Cobitis
elongata, that measured nearly 16 cm.

The body is shallow and compressed, but of fairly
uniform depth. The greatest depth of the body, which
generally lies about half-way between the head and the
dorsal fin, is about 1/s of its length0. The thickness at
the same point is about 7/n of the depth in our most
terete specimen; but as a rule the body is still thinner,
the greatest thickness being sometimes no more than
half the greatest depth. The body is of almost uniform
thickness throughout its depth, with obtusely rounded
dorsal and ventral edges, at the tip of the tail ribbon-
shaped, in front with the snout obtusely rounded at
the sides. This species, like the next one and the
majority of the species within the family, is charac-
terized in general by a slightly marked (comparatively
deep) peduncle of the tail, the least depth thereof vary-
ing between 7 and 8 % of the length of the body. But
in the aforesaid form ( Cobitis elongata) described by
Heckel and Kner, this depth seems from their figure
to have been only 5 % of the length of the body.

The length of the head measures 1 6 1/2 d — 1472 %
of that of the body. It gives the fish a singular ap-
pearance on account of its sharply arcuate upper pro-
file, with the deep snout projecting beyond the mouth
and lower jaw, which lie on the same plane as the
lower profile, which is generally straight. The specific

length,

8 Sometimes 5, according to Heckel and Kner.

b 41, according to Valenciennes. 40 — 42, according to Canestrini. 43 — 44, according to CederstrSm.

c According to Canestrini’s measurements of 23 specimens the greatest depth of the body varies between 10 and 1 6 1 / 2 % of its
increasing generally with age.

d In young specimens (less than l/, dm. in length) even 18 ft, %, according to Canestrini.

SPINED LOACH.

707

character also appears with considerable distinctness in
the compressed form of the head, the thickness at the
eyes being less than 2/ 5 of the length of the head", or
than 3/4 of the length of the base of the dorsal tin''.
The deep form of the snout is due to the marked de-
velopment of the vertical plate of the ethmoid bone in
front of the narrow frontal bones, which are arched
above the orbits. The small round eyes are set high
and turned somewhat upwards, above the more or less
tumid cheeks, the distance between them and the upper
protile of the forehead being about half their diameter6'.
Their position is such that the length of the snout/7
(from their anterior margin obliquely downwards to
its tip) is less than the postorbital length of the
head6 (from the posterior margin of the eyes ob-
liquely downwards to the hindmost part of the margin
of the branehiostegal membrane). As in Misgurnus
fossilis they are entirely covered with a thin skin
( oculi velati, Bleeker), the orbits being externally de-
stitute of any sharp limits. The anterior part of the
orbits is coasted below by a longitudinal slit in the
skin, out of which slit the fish can erect the preorbital
spine, which otherwise points in a backward direction.
This spine is formed, as Lilljeborg has pointed out,
by the anterior frontal (lateral ethmoid) bone on each
side. This circumstance may be most easily elucidated
by a comparison with the following species, where the
anterior frontal bone occupies exactly the same position
as the extended (vertical) base of the spine in this spe-
cies, forming the anterior margin of the orbit in a ver-
tical position between the frontal and parasphenoid
bones, but with a firm osseous connexion (suture) with
the anterior outer margin of the orbitosphamoideum,
which even separates the frontal bone proper from the
upper anterior corner of the triangular anterior frontal
bone. Here, in the Spined Loach, the said osseous
connexion is loose and transformed into an articulation.
rfhe bone is also of much harder texture, white and
firm as dentine. Its base, which forms the long, ver-
tical articulation with the orbitosphenoid bone, is almost

terete and curved in a rounded obtuse angle, but is set
in a somewhat oblique position, the lower (shorter) arm
of the angle running obliquely inwards and forwards,
while the upper arm is vertical. The tooth projects
backwards in a direction approaching to that of the
lower arm, at least below the middle of the basal part.
It is curved backwards and inwards and is generally
bicuspid, a shorter, but equally pointed tooth being set
at the middle of the outer surface of the main tooth7.
The base of the tooth is united by a strong ligament
to the hind margin of the orbit. The suborbital bones
seem otherwise to lie entirely wanting both in this spe-
cies and Misgurnus fossilis. In front of the preorbital
spine a dentiform protuberance belonging to the palatine
bone may be felt beneath the skin, at the point where

Fig. 176. Right preorbital spine of a Cobitis tcenia , seen from without.

About 15 times the natural size.

this bone is elongated in a backward direction to meet
the entopterygoid bone. The nostrils are set rather high
on the sides of the snout, somewhat nearer to the eyes
than to the tip of the snout. The posterior nostril on
each side is round and somewhat larger than the anterior,
which lies just in front of it, and the hind superior
margin of which is canaliculate or elevated in an obli-
quely-cut, tubular form. The mouth is small and tooth-
less; it lies on the under surface of the head, fringed
by a comparatively thin upper lip, thickened only at
the corners of the mouth, and small barbels, which
comprise one pair in front and one on each side at the
middle of the upper jaw, belonging to the dermal fold
that runs from the cheeks over the mouth and forms
the tip of the snout. The hindmost (largest) pair of
barbels, on the other hand, lie at the corners of the

“ 26 — 36 %, according to our measurements of specimens between 1/2 and 1 dm. long. The greatest thickness of the head measures,

according to Canestrini, 27 — 40 % of its length in the typical Cobitis tcenia ; but in the form which has received the name of C. Icirvata ,

it is said sometimes to measure 47 *L % thereof.

b 48 — 69 %, according to our measurements.

c The diameter of the eyes is about 20 — 15 % of the length of the head.
d About 38 — 43 % of the length of the head.
e About 49 — 56 % of the length of the head.

f G. C. CederstrOm (1. c.) has remarked as an external sexual difference that in the males the preorbital spine is furnished with

several lateral denticulations. In the males examined by us, however, this character does not hold good.

708

SCANDINAVIAN FISHES.

mouth and belong to the thick part of the lip. On the
lower jaw too, the lip runs back in a dermal flap on
each side. This flap, as Kroyer3 has already pointed
out, may now and then be denticulated or even pro-
duced at some spot so as to resemble a barbel, thus
depriving the above-mentioned generic difference be-
tween Cobitis and Misgurnus to some extent of its
validity. The intermaxillary bones are small and nar-
row, with the main branches only slightly longer than
the straight nasal processes. The maxillaries, on the
other hand, are high and of singular form. Behind the
articular knob they expand into a square, but again
contract, and at the hind extremity once more expand
into a rounded lobe curved in a downward direction.
In the upper jaw we find a well-developed palatal fold.
There is no free tongue. On the first branchial arch
the gill-rakers are set in a single row (containing 11 —
14) corresponding to the posterior row on the other

Fig. 177. Right lower pharyngeal of a Cobitis taenia , seen from
within and above. About 15 times the natural size.

arches, where they are set in a double row, and on the
lower pharyngeals in one row (containing 7) on the outer
anterior margin of these bones. They are short and
depressed. The pharyngeal teeth are subulate, pointed,
and curved; they are set in a row (10 or a few more,
3 of which lie on the upper arm) on the inner (posterior)
margin of the lower pharyngeals, which are geniculate,
like branchial arches. Sometimes, however, we find one
or two smaller teeth (supplementary teeth?) beside the
principal row. The gill-openings are merely vertical
slits, the branchiostegal membrane on each side of the
body coalescing with the skin at the anterior end of
the insertion of the pectoral fin. The height of the
opening is about equal to the thickness of the head at
the eyes; but the branchiostegal membrane is loose, the
gill-openings being thus capable of expansion by means
of the three long, sabre-like branchiostegal rays on each

side. The preoperculum extremely narrow, the other
opercular bones well-developed. The hind inferior
margin of the operculum proper is curved in an S-shape,
with the anterior lower corner produced in a downward
direction, but the breaks are filled by the suboperculum.
As we have mentioned above, the head is scaleless; but
the ducts belonging to the system of the lateral line
are usually quite distinct on its surface, being marked
by rows of small pores, often raised in a tubular form,
along the temples, straight across the occiput, along the
preoperculum and the branches of the lower jaw, on
the forehead and at the upper orbital margin on each
side, and along each of the cheeks, below the slit con-
taining the preorbital spine and forward on the snout
below the nostrils.

The body, on the other hand, is densely covered
with small, round, thin, imbricated, cycloid scales. The
lateral line is distinct only at the very beginning, where
it forms a backward continuation of the temporal canal,
first sloping downwards and then straight, for a distance
of about twice the longitudinal diameter of the eyes.

The dorsal fin is obliquely rectangular, with the
upper angles rounded, the upper posterior margin being
thus more or less convex. Its base is somewhat elevated
in front, forming a slight break in the otherwise straight
contour of the back. Its beginning lies at a distance
from the tip of the snout that measures about 41 — 43
% of the entire length of the body, 47 — 50 % of the
length of the body minus the caudal fin, or 67 — 65 %
of the distance between the anal fin and the tip of the
snout, and is generally situated somewhat in front of
the perpendicular from the insertions of the ventral fins.
At its origin we find two rudimentary, unarticulated
rays (supporting rays), the first extremely small, the
second about one-third as long as the third ray. The
third ray is simple (undivided) but articulated, and
only slightly shorter than the fourth, which is the
longest ray in the fin and, like the remaining rays,
branched. The last ray is about half as long as the
longest one, and there is no fin-membrane behind it.
The shape of the fin is highly inconstant, the length
of its base (on an average about 872 % of the length of
the body6) varying between 64 and 76 % of its height
(the length of the longest ray).

The anal fin is of the same structure and shape as
the dorsal, only that the branched rays are fewer and

a Damn. Fislce, III, p. 568.

b Varying, according to Canestrini’s measurements, between 7‘2 and 10 % of the length of the body.

SPINED LOACH.

709

the base shorter, on an average about 6 % of the length
of the body®. Its shape varies otherwise exactly as that
of the dorsal tin. The lips of the vent protrude in a
tubular form within a triangular depression, which is
situated just in front of the beginning of the anal tin,
or at a distance in front of this point hardly as great
as the diameter of the eyes.

The pectoral tins consist of 8 or 9*, sometimes 10
rays. In the last case we tind two simple rays (the
first, only rudimentary) at the anterior margin of the
tins; while otherwise only the first ray is simple and
not much shorter than the second, which is branched
and either longest or about equal in length to the third
ray. The remaining rays gradually decrease in length
and are all deeply branched, except the last, which is
simple and about 1/3 as long as the longest ray. The
fins are set low down and when expanded occupy a
horizontal position. Besides the above-mentioned sexual
difference first remarked by Canestrini in the structure
of the pectoral tins, we also tind in the Spined Loach
a second one, which was pointed out by Bonizzi to
Canestrini. At the base of the posterior (upper or
inner) side of the pectoral tins we find a cartilaginous
lobe, set in an obliquely transverse position. This lobe
is large and of more general occurrence in the males0,
smaller, rudimentary or, most commonly, entirely want-
ing in the females. Cederstrom noted a third external
difference between the sexes, consisting in the fact that
in the males the pectoral tins are as a rule longer, and
therefore more pointed, than in the females. In 7 males,
between 50 and 73 mm. long, the length of the pectoral
tins varied between 15 and 18 % (on an average 1 6 V2
of that of the body; in 5 females, between 88 and 96
mm. long, this percentage varied between 10 and 1 1 1/2
and was on an average Ilk

The ventral tins are of the same structure as the
pectoral, but shorter and more rounded. In the males
their length is about 11 — 13 %, in the females about

9—10 % of that of the body. In this relation, however,
we must also take into consideration the changes of
growth, for the relative length of the ventral tins de-
creases with increasing age. At the outer angle of the
insertion of each ventral tin — where in the Teleosts in
general a triangular dermal flap, free at the top and
pointing in a backward direction, is furnished with sin-
gular, generally elongated scales — we find a, small, ve-
siculate, oblong and posteriorly blunt, dermal swelling,
which is, however, without scales of a special type0.

The caudal tin, which in specimens between and

1 dm. long occupies about 13 — 12 % of the length of
the body, is truncate with rounded corners. The middle
14 rays are branched. The small, short, supporting
rays generally number 3 or 4 (sometimes 5f) above and

2 or 3 (sometimes 4f) below. The base of the tin ad-
vances, in the form of a thin ridge, forward along the
upper and lower margins of the peduncle of the tail.

The digestive canal is extremely simple, consisting
of a straight tube of almost uniform thickness from the
pharynx to the vent, with only a slight expansion in
the anterior half to indicate the presence of a special
stomach. The peritoneum is silvery white. The liver
is long, with two lobes, the right lobe being generally
the longer*7. The ovary is simple, but the testicles are
paired. At the spawning-season both the ovary and the
testicles extend forward along almost the whole length
of the abdominal cavity*. When the ovary is as full
of eggs as possible, it is deeply forked underneath, thus
appearing double when seen from below.

The ground-colour of the body is yellowish, some-
times even orange, on the back more or less dashed
with gray, on the belly white. The singular markings
consist of dark gray or even blackish spots and streaks.
Large spots are set in a longitudinal row at the middle
of the back, along the dorsal sides, and along the middle
of the sides of the body, where they sometimes coalesce
into bands, especially behind; and between these rows

“ Varying, according to Canestrini’s measurements, between 5'4 and 7'1 % of the length of the body.
b Sometimes only 6 or 7, according to Heckel and Kner, Canestrini, and Benecke.
c Cf. the similar growth in the Bramoids and Blennioids; see above, pp. 76 and 219.
d According to Canestrini’s measurements this sexual difference is less marked.

e In the following species this dermal swelling is equally insignificant, in Misgurnus fossilis we have failed to find it at all.
f According to Lilljeborg.

'J As usual, however, varying in length. In a gravid female 108 mm. long the right lobe of the liver extends to a line with the tip

of the pelvic bones, the left to the end of the second third of the distance between the insertions of the pectoral and ventral fins. In a

male almost ready to spawn and 69 mm. long, the left lobe of the liver is somewhat longer than the right and extends about as far back

as the left lobe in the female just mentioned.

h In the female just mentioned the eggs were about 3/4 mm. in diameter. In another female, which had partly deposited its spawn,
eggs l3/4 mm. in diameter were found.

710

SCANDINAVIAN FISHES.

lie smaller, somewhat lighter spots arranged in a net-
work or scattered. Similar small spots of an oblong
shape or even vermiculated also appear on the top of
the head and on the cheeks. The most prominent and
most persistent parts of this design, even in specimens
preserved in spirits, are two black markings, the one on
the head, the other on the peduncle of the tail. From
the anterior margin of the eye a blackish streak runs
obliquely downwards to the tip of the snout, and a
similar streak often appears behind the eye horizontally
across the temple. At the upper corner of the base of
the caudal fin (sometimes at the lower corner as well)
we line! a black spot, edged during life, like the spots
in the large, inferior row on the sides of the body, with
a lighter colour. These two markings recur, however,
though they are generally less sharply defined, in other
species of the genus. The tins are more or less trans-
parent, with lustrous rays, which in the dorsal and
caudal fins, sometimes in the pectoral and anal fins as
well, are marked with dark spots, distinctly arranged,
at least on the first-mentioned fins, in transverse bands.
The iris is yellow.

The Spined Loach occurs in the brooks, rivers, and
lakes of almost all Europe, and is probably spread
throughout Northern Asia, except perhaps the colder
regions, for according to Schlegel (1. c.) it lives in
Japan. Pallas also quotes special names for this fish
not only from the Ostiacs of Siberia, but also from the
Tungs of the River Katunga and Lake Baikal. The Spi-
ned Loach goes westward at least to England and Scot-
land; but in Ireland, according to Thompson, it is want-
ing. Valenciennes knew it as a Spanish species. Ac-
cording to Canes trini it also occurs in Sicily. In Scan-
dinavia the Spined Loach is common enough in scattered
localities in the south and east of Sweden, at least up
to Lake Wener and the Malar Valley. It seems to be
commonest in Oster Gothland and in the basin of Lake
Malar. To the best of our knowledge it is wanting in
Halland and Bohuslan", as well as in Norway. It is
assigned to the Danish islands of Fiinen, Laaland, and
Zealand by Feddersen, who states that it is especially
common in Lake Mariebo (Laaland). The Royal Mu-
seum possesses specimens from the River Kopinge in
Scania and from Jonkoping (C. Sundevall), from Lilia

Halsviken (Wetter, N. W. of Motala; H. Widegren),
from the River Skeninge (numerous specimens; Dr.
Cnattingius), from the River Lida in Wester Gothland
(J. W. Dalman), from Lake Wener (1836, Mr. Goebel* * 6),
from Lake Softer in Nerike (1842, Colonel Ankarsvard),
from Lake Malar off Flottsund (Mesch), from Helene-
borg (Liljeholmen), and from Hammarby Lake near
Stockholm. In the Norrstrom (Stockholm) a Spined
Loach was taken on the 16th of April, 1846, among spe-
cimens of the Smelt. In 1869 Cederstrom caught a spe-
cimen 42 mm. long in the channel off Beatelund in the
island-belt of Stockholm, a proof that the Spined Loach
can also live in the brackish water of the Baltic. Ce-
derstrom found the Spined Loach especially plentiful in
the River Orsunda in Westmanland. Lilljeborg assigns
it to several other localities in Sweden, and surmises
with reason that the Spined Loach is fairly common in
Scandinavia, though on account of its insignificant size
and its manner of concealing itself it easily escapes notice.
This is probably true of Finland as well, though both
Malmgren and Mela state that the Spined Loach is ex-
tremely rare in that country and is known with certainty
only from the neighbourhood of Viborg and Lake Vuoxen.

The Spined Loach prefers running water, in small
streams with a stony bottom, where it can conceal itself
under the stones, or with a bottom of gravel, sand, or
even mud, in which it can bury itself with only the
head visible and ready, when danger threatens, to hide
itself entirely or even to creep some distance, burrowing
its way through the loose ground. The fish also occurs,
however, in still water, as we have seen, in lakes and
meres with a suitable bottom. It is oftenest found in
company, though not in shoals properly so called. It
generally lies still; but when disturbed or when shifting
its position it is rapid in its movements. Cederstrom
saw “these fishes dive into the thick, moist ooze and hide
themselves there as speedily as a Sand-Eel plunges into
wet sand to conceal itself. When they felt themselves
prisoners, they at once bent down the head and pressed
the fiat part of the cheek firmly against the skin of my
fingers, thus causing in some instances a slight but
disagreeable irritation, most like that of sucking (cupping),
on the part of the skin affected.” The fisherman may
sometimes complain with reason0 that “when pursuing

a Holmberg’s statement ( Boh . Hist ., Beskr., p. 30) that the Spined Loach occurs in Ramtvet Mere (Bullar), lias received no subsequent

confirmation, according to Malm. See Gbgs , Boh. Fn ., p. 569.

6 The Spined Loach has since been taken in Lake Wener (Hamrnarovik), according to Dr. S. W. Tenow, Verml., Dais Ryggr. Dj., p. 106.
c See Lloyd, Scandinavian Adventures , vol. I, p. 71. On the 11th of June, 1858 I took some specimens of this species in the River
Motala, not far from its outlet in Lake Roxen. There I was told that the Spined Loach was dreaded “for its bite”, which was said to be incurable.

SPINED LOAC'II.

711

liis avocation barefooted in the summer the fish not
unfrequently wounds his feet;” and if the spine be
broken off and left in the wound, it may well cause
some trouble. But, as Blanchard" has already re-
marked, these spines cannot serve as any formidable
weapons of defence, even if they are emplcqed in this
capacity in case of need. Cederstrom assumes their
function to be the protection of the eyes, as the fish
burrows along in the sand or gravel. Perhaps they may
also be organs of adhesion, like the opercular and inter-
opercular spines of the Stegophilidce, those small Brazi-
lian Glanomorphs6 which usually take refuge in the
branchial cavity of larger fishes, and retain their posi-
tion there by means of the said spines, but which also
try to force their way into other cavities, or to attach
themselves with the spines to the skin of larger aquatic
animals or of persons bathing or wading.

The spawning-season occurs in spring and early
summer, from April to July; but as yet we know no-
thing of its course. It is generally stated that the males
are far rarer than the females; but in a consignment
of some thirty specimens from the River Skeninge there
were as many males as females. The former being,

however, as a rule smaller than females of the same age,
they probably manage more easily to escape observation.

The food of the Spined Loach consists principally
of small crustaceans, Entomostraca ( Lynceidce , according
to Cederstrom); but the fish also preys on all other mi-
nute animals that come in its way. According to Ivroyer
it is “fairly voracious and lives on worms, but also on
fish-roe, small fry, and other minute aquatic animals.”
The Spined Loach shows the same propensities in aquaria,
where it is “interesting by reason of the eagerness with
which it roots up the bottom in quest of food, casting
sand and gravel in pellets out of its gill-openings”
(Benecke).

The Spined Loach is of little use as human food,
the flesh being dry and tough; but as bait for Eels,
Pike, and other predatory fishes it may be employed
with advantage c.

In the Malar Provinces the Spined Loach is known
by several names: tanglake (Tang Burbot), according to
Artedi; ormfish (Snake-fish), according to Gyllen-
stjerna (in Nilsson); stenlake (Stone Burbot), according
to Iverus (in Lilljeborg). In Scania Lilljeborg heard
it called stenbit (Stone-biter).

THE LOACH (sw. gronlingen).
COBITIS BARBATULA.
Plate XXXI, fig. 5.

Body rather terete. Head somewhat depressed: its breadth ( thickness ) at the eyes more than 2/5 of its length;
breadth of the interorbital space greater than the longitudinal diameter of the eyes. No preorbital spine below the eyes.

V.

E. br. 3; D.
1

(3 1. 2)+ 1 (2 1. 1)+ 1 1_

5 1.6 ’ ' 111.12’

(6) 7 1. 8

6 1. 7

; C. *+1 + 16+1+*; Vert. 41 A

Syn. Cobitis barbatula, Rond., Pise., part. II, p. 204. Cobitis tota
glabra maculosa, corpore subtereti, Art., Ichthyol. , Gen. Pise.,
p. 2; Syn. Pise., p. 2; Lin., Fn. Suec., ed. I, p. 125.
Cobitis Barbatula, Lin., Syst. Nat., ed. X, tom. I, p. 303 ;
Penn., Brit. Zool., vol. Ill (ed. 1776), p. 247. tab. LVIII,
No. 142; Bl. Naturg. Fish. Deutschl., part. I, p. 224, tab.
XXXI, fig. 3; Pall., Zoogr. Ross. Asiat., tom. Ill, p. 164;
Nilss., Prodr. Ichth. Scand., p. 35; Cuv., Val., Hist. Nat.
Poiss., vol. XVIII, p. 14, tab. 520; Kb., Damn. Fisk., vol.
Ill, p. 539; Nilss., Skand. Fn., Fisk., p. 343; Ivessl., Bull.
Soc. Natural. Mosc., tom. XXIX, p. 350; Tiiomfs., Nat.

Hist. Irel., vol. IV, p. 139; Sund., v. Wr., Skand. Fisk.,
ed. 1, p. 207, tab. 53; Hckl, Kn., Stisswasserf. Oesterr.
Mon., p. 301; Mgrn, Finl. Fisk. (disp. Helsingf.), p. 36;
Canestu., Arch. Zool., An at., Fisiol., vol. IV, fasc. I, p. 144;
Steind. Stzber. Akad. Wiss. Wien, Math. Naturw. CL, LIII, I
(1866), p. 203; Blanch., Poiss. cl. eaux douces Fr., p. 280;
Gthr ( Nemachilus ), Cat. Brit. Mus., Fish., vol. VII, p. 354;
Sundstr., Fn. Sverig. Ryggradsdj., p. 278; Fedders. (6V
bitis), Naturb. Tidskr. Kbhvn, ser. 3, vol. XII, p. 92;
Mor., Hist. Nat. Poiss. Fr., tom. Ill, p. 432; Bncke,
Fisch ., Fischer., Fischz. O., IV. Preuss., p. 145; Mela,
Vert. Fenn., p. 313, tab. X; Day ( Nemacheilus ), Fish.
Gt. Brit., Irel., vol. II, p. 203, tab. CXXXYII, fig. 2;
M6b., Hcke (Cobitis), Fisch. Osts., p. 123; Lillj., Sv.,
Norg. Fisk., vol. Ill, p. 332; Fatio ( Nemachilus ) Fn.
Vert. Suisse, vol. V, p. 19.

° Poiss. cl. eaux douces Fr., p. 287.

1 See Reinhardt, Vid. Meddel. Naturb. For. Kbhvn, 1858, pp. 83 et seqq. LUtken, ibid. 1891, p. 55, note.
c BLANCHkEE, La peche et les poissons, p. 449.
d Sometimes 39 (Lilljeborg), 40 (Kroyer), or 42 (Valenciennes).

90

Scandinavian Fishes.

712

SCANDINAVIAN FISHES.

The Loach is generally a little larger than the pre-
ceding species. In Scandinavia and Western and North-
ern Europe it seems never to attain a greater size than
10 — 13 cm. In the Crimea, however, according to
Pallas, it may sometimes measure nearly half a foot
(about 16 cm.); and in the mountain brooks of Persia,
lie says, it attains a still greater size. The largest
specimens we have examined were from Finland and
measured 112 mm. in length.

The body is in front more or less terete, behind
more and more compressed. The head is entirely diffe-
rent in form from that of the preceding species, but
the rest of the body is of almost exactly the same shape,
the tail being only slightly shallower than the forepart
of the body. The greatest depth of the body measures
between 13 and 11 % (10 1/2 %) and the least depth of
the peduncle of the tail between about 7 and 872 %
of the length of the body.

The length of the head is about 20 — 18 %a of that
of the body. Seen from above it is almost parabolical.
It is also broader than in the preceding species, some-
what depressed, flattened at the top, with sloping facial
line and blunt snout. Underneath it is broad, flat, and
even. The entire head is scaleless; but the skin is here
finely granulated, the canals of the system of the lateral
line being marked by several rows of small, raised
tubercles, each pierced with a duct. One row, thin and
almost double, runs below the eye from the upper
rostral barbel to the lateral line. Another roAv, con-
taining some few pores, runs above each eye and the
nostrils. A third row coasts the margin of the pre-
operculum and follows the under surface of the lower
jaw. A transverse canal across the occiput may also be
traced in three scattered pores, set in a row. The eyes
are small and somewhat oblong, their longitudinal dia-
meter in specimens between 87 and 112 mm. long being
about 16 % of the length of the head. The orbits are
distinctly bounded by a dermal fold ( oculi liberi, Blee-
ker). They are set high, at the sides of the flat fore-
head, separated by a distance of about three times their
vertical diameter, and at a distance from the tip of the
snout at least (in our specimens) a little greater than
the postorbital length of the head, which length in young
specimens is almost equal to the breadth of the head
at the eyes, in older ones perceptibly less. The cheeks
are soft and fleshy, without spine or hollow depression.

The nostrils lie just in front of the eyes, each lateral
pair being so close together that the anterior nostril
does not extend half-way from the eyes to the tip of
the snout. The posterior is the larger and simple, the
anterior has a raised, tubular margin, which projects
behind in a point and thus resembles in form a quill
pen. The mouth lies, as in the preceding species, under
the tip of the snout and is very small and transversely
set. The lips are thick, and the upper seems double,
as in so many other fishes, this being due to the for-
mation of an upper fold by the projecting skin of the
snout itself, in addition to the fold starting from the
intermaxillary bones. These two folds send out, as in
the preceding species, 6 thick, soft barbels, so arranged
that two pairs originate from the under surface of the
upper fold, the smaller pair at the very tip of the snout
and the other pair just behind this point; while the
third pair starts from the very corners of the mouth
and contains the largest barbels, their length being a
little more than V3 of that of the head. The underlip
is more fleshy and at the middle — often on the sides
as Avell — cloven or divided into lobes. The gill-open-
ings are small, the branchiostegal membrane passing, in
the same Avay as in the Spined Loach, into the skin of
the belly just beloAv the base of each pectoral tin, so
that the openings are separated by the entire breadth
of the broad, flat breast. This membrane contains 3
long, but not broad rays, Avhich extend almost below
the tip of the operculum. The operculum itself as Avell
as the small, oblong suboperculum is distinct; the other
opercular bones are Avell covered by the thick skin.
The apparatus of the branchial arches exhibits in the
form of the urohyoid bone a striking resemblance to
the corresponding bones in the Sheatfish.

The dorsal tin begins someAvhat in front of the middle
of the entire length of the body, or about half-Avay —
sometimes a little farther back — along the body minus
the caudal fin: the distance betAveen this tin and the tip
of the snout measures in our specimens about 42— 48 %
of the entire length of the body, 48 — 55 % of the length
of the body excluding the caudal fin, or 65 — 71 % of the
distance betAveen the anal tin and the tip of the snout.
The length of its longest ray is about equal to the
greatest depth of the body or someAvhat greater than
this depth, and is much greater than the length of the
base of the tin, Avhich seems as a rule to vary betAveen

19 — • 1 6 ' G %, according to Canestrini’s measurements.

LOACH.

713

about 9 and 11 %a of the length of the body. The
rays are actually 11 in number, but the first two are so
small and line as to be difficult of detection; the third
ray is also quite short; the fourth ray does not extend
quite to the top of the fin and, like the rays in front
of it, is simple; the fifth and sixth rays are the long-
est and, like the following ones, branched.

The anal fin is much smaller and generally contains
8 or 9 rays, among which the first branched ray (the
3rd or 4th from the beginning) is the longest. The
vent occupies the same position as in the preceding
species.

The caudal fin is truncate at the hind margin, with
rounded corners. Of its 18 articulated rays the 14
middle ones are equal in length, but the two uppermost
and the two lowest somewhat shorter. The outermost
of these rays above and below is simple, the others are
branched. Besides these there are a few quite short,
unarticulated, supporting rays. The median length of
the fin varies between about 1 1 1/8 and 14 % of that of
the body or between 60 and 75 % of that of the head.

The pectoral fins are fairly large — their length
greater than the greatest depth of the body — broad,
and rounded, with 11 — 13 rays (usually 12), among
which the first is simple, articulated, and rather short,
the others are multifid and deeply branched, the second
and third being the longest, the last two very small
and difficult to distinguish with accuracy.

The ventral fins are much smaller than the pec-
toral, their length being only slightly greater than the
height of the anal fin. They begin almost vertically
below the beginning of the dorsal fin, are oval in shape,
and contain 7 — 9 rays (usually 8), among which the
outermost is simple, articulated, and rather short, the
others are deeply branched, the innermost two closely
united, and the second and third the longest.

The scales are rather small — in specimens 12 cm.
long their breadth is less than 1 mm. — and circular,
densely set, but not imbricated, numbering about 15
from the lateral line to the dorsal edge. On the fore-
part of the body, on the back and belly, we find only
a few, scattered here and there; on the hind part of the
body they are firmly inserted and set close together. The
lateral line is straight. Neither here nor in the Spined
Loach does it pierce any scales. It starts from the upper

angle of the gill-opening and then follows the middle
of the side. In front it is furnished with distinct, pro-
minent pores, but behind these pores become indistinct,
being hidden by the scales.

The coloration is of a fairly bright, grayish brown
with darker, large spots along the back and the lateral
line; but these spots are not so distinct and constant
as in the Spined Loach; below they are broken up into
an inconstant pattern, with which the sides of the body
as well as the cheeks are mottled. As a rule we find
a black streak on the sides of the snout as in the Spined
Loach; and an oblong, black spot usually appears at the
lower corner of the base of the caudal fin; but even
these markings are less defined than in the Spined Loach.
The dorsal, caudal (sometimes the anal as well), and
pectoral fins bear transverse bands of dark spots. The
ventral fins as well as the anal fin are in most cases
plain, in others marked with a few dark spots.

The internal organs differ from those of the Spined
Loach partly in the liver being somewhat shorter and
behind not divided into lobes but collected in a mass,
which envelops the stomach below and on the sides,
partly in the stomach being furnished with a pyloric part
pointing in a forward direction; the intestine also forms
a coil, which runs forward beside the posterior part of
the stomach. The ovary is forked in front.

According to observations made in the vicinity of
Kuopio in Finland and communicated by W. vox Wright
to Sundevall, the Loach is there common6 in streams
and brooks with fairly clear water, a gravelly bottom,
and a depth of a foot or two. It is said to keep close
to the bottom and generally to hide under stones, but
not to burrow in the bottom, as the Spined Loach does.
In the said locality it spawns at the beginning of June,
about a fortnight after the breaking up of the ice.
In Germany, according to Bloch and other writers, it
spawns in March and April, or early in the spring,
there as in Finland. Its habits are otherwise not un-
like those of the Spined Loach.

Linnaeus tells us in the Fauna Snecica that this
species was introduced from Germany into Lake Malar
by King Frederick I; and the Royal Museum possesses
two specimens with no assigned locality, but marked
in the same way as the other fishes from the Museum
of Drottningholm, where Linnaeus examined and cle-

“ Sometimes about 8 %, according to CanestrinTs measurements.

b According to Qvensel’s catalogue one specimen in the Royal Museum was taken in 1800 at Uleaborg. Mela also believes that the
species occurs in the extreme north of the Gulf of Bothnia.

714

SCANDINAVIAN FISHES.

scribed them. These specimens may thus be ascribed
with reason to the said transplantation. However, the
Loach does not seem to have continued to multiply in
Lake Malar; Sundevall at all events never succeeded
in obtaining the slightest evidence of its occurrence
there. On the other hand, it has been found in recent
times both in Scania and the south of Halland. In Scania
it was discovered in 1864 by Lector FIultmark in a
brook at Trollenas and by Mr. C. Moller at Gisleberga
and Bosarp; in 1869 it was met with by A. Nathorst
in the river at Andrarum. In 1888 the Royal Museum
received through Mr. Trybom, Assistant Inspector of
Fisheries, two specimens which had been taken by Mr.
F. Svenonius in the Laga River at Laholin. It may
possibly be found on closer investigation to occur in a
few more parts of Sweden; but a fish of this small size
may easily escape observation. In Norway it is wanting,
to the best of our knowledge; but in Jutland it was
found in 1878 by C. E. V arming in the River Nebels.

— It has been met with throughout the rest of Europe
as well as the north of Asia. In Germany it is said
to be taken at several places in great numbers with
nets constructed for the purpose. Its flesh has always
been praised as highly delicious and so easy of digestion
that even a weak stomach has nothing to fear from it.
Bloch describes at length the method of constructing
holes or small pools for the cultivation of this fish.

The Swedish name ( Grouting ), which we have applied
to this species, is given by Linnaeus in his Fauna
Suecica. It is now unknown Avith Avhom the name ori-
ginated — Ave are also ignorant of the source from Avhich
Nilsson clreAv his name of nissoga (BroAvnie-eye) for the
preceding species — but it is obviously a corruption
of the German Griindling (Eng. Groundling), a name
Avhich has been given to the fish because it keeps close
to the bottom. There is nothing green in its coloration.
Another Scandinavian name for the Loach is Smarting,
from the German Schmerling.

For the elucidation of the natural relations be-
tween our two Colitis species Canestrini’s numerous
measurements afford interesting materials of com-

parison. Taking into account the measurements given
by him Ave obtain the folloAving results:

Average

i n

Cobitis barbatula.

Cobitis

tcenia.

4

specimens.

3

specimens.

9

specimens.

9

specimens.

Length of the bodv expressed in millimetres

53.6

92

56.2

85.6

(1) Lateral length of the head

.. in

% of

the length

of the body

18.3

17.3

17.i

15.8

(2) Length of the head to the occiput

-- »

55 55

55 55

55 55

55

16.6

15.8

15.2

14.9

(3) Diameter of the eyes

-- j?

55 55

55 '5

55 55

„

4.i

3.4

3.i

2.3

(4) Preorbital length of the head

-- 55

55 55

55 55

15 55

„

7.9

7.7

7.3

6.8

(5) Breadth of the head

5 J 55

55 55

55 5>

„

9.9

9.i

5.4

5.2

(6) Depth of the body

-- 55

55 55

55 55

55 55

55

12.o

12.4

13.o

14.3

(7) Length of the dorsal fin

-- 5>

55 55

55 55

55 55

55

9.5

8.5

8.5

8.4

(8) Height ,, „

55

55 55

55 55

55 55

55

14.3

12.7

14.3

13.5

(9) Length of the pectoral fins

*- 55

„ „

55 55

55 55

„

16.2

14.2

13.8

11.9

(10) .. ,, ventral „

- »

55 55

55 55

55 55

55

12.3

12.2

11.6

10.3

(11) anal fin....

55 55

55 55

55 55

55

6.2

5.4

6.3

5.9

(12) ,, caudal

-- 55

55 55

55 55

>5 55

„

16.4

16.2

—

—

(13) „ ,, „ head and trunk

-- 55

55 55

55 5 5

55 5 »

55

57.5

60.2

62.4

64.6

species belong to a common direction of development;
and in this direction of development Colitis tcenia
evidently occupies the more advanced rank.

(Sundevall, Smitt.)

The correspondence in the changes of development
is here so great — the percentages of the two different
ages in both species rise or fall Avith so great similarity
— that Ave are fully justified in assuming that these tAvo

CARP-FISHES.

715

Fam. CYPRINIDtE.

Air-bladder free ( not enclosed in a capsule).

The family of the Cyprinoids, the largest in the
whole class of fishes, contains about 1,000 described
species, most of them, however, from Asia and North
America, so that only a hundred species occur in Europe,
and only a score belong to the Scandinavian fauna. The
majority of our fresh-water fishes are comprised, how-
ever, in this family. All the Cyprinoids prefer fresh water
and are most commonly found in lakes, rivers, ponds, and
fens. Still most of our species also occur in the brackish
water of the Baltic. Not a single member of the family
is a predatory fish properly so called ; most of them live
chiefly, though not exclusively, on vegetable substances.
They may with every reason be called omnivorous. In
a certain sense they may be regarded as Ruminants:
the Carp, the Tench, and the Bream are adduced as
examples of this; but as a rule food passes rapidly
through their intestinal canal: a Goldfish fed with wheat-
bread passes after some minutes a white, vermiform mass
of excrement, which hangs from the vent. They are
sensitive in a high degree to atmospheric influences;
but some of them, though they do not properly belong
to an Arctic temperature, can in a torpid state survive
the process of being frozen". Even in a milder climate
they are known in cold weather to collect in dense
shoals, which lie still in the deepest parts of the water;
and Valenciennes6 states of the Barbel of Southern
Europe that he once found a company of this species
which during the winter had packed themselves together
in the hollow trunk of a tree.

Among the peculiarities in the reproduction of
these fishes we shall here remark only one, which is of
general interest, but belongs to a species foreign to our
fauna, the little Bitterling ( Rhodeus amarus ), a form
fairly common in Eastern and Central Europe and at
most about 9 cm. long. It has long been known that
the eggs of some fish are found among the branchial
lamellae of the painter’s mussel ( Unio pictorum) ; but until

Mouth fringed with at most four (or no) barbels.

1869 it was a matter of doubt to what species these
eggs belonged. Before this time Krauss (1858''), Kess-
ler (1860d), Dybowski (1862"), and Siebold ( 1868')
had described an external oviferous tube, sometimes 30
mm. long, which in the female Bitterling is developed
during the spawning-season from the margins of the
urogenital opening just behind the vent, and into which
the comparatively large, ellipsoidal eggs — sometimes 3
mm. long — force their way and arrange themselves in
a single row. Noll£/ at last discovered that the said
eggs in the branchial cavity of the painter’s mussel be-
long to the Bitterling. By observations which he has
since completed, he showed that, when the eggs are ripe,
the female Bitterling applies the oviferous tube to the
inspiratory opening of the painter’s mussel, into which
the male at, the same time emits his seminal fluid. In
this Avay about 40 eggs may be forced one by one into the
branchial cavity of the mussel, Avhere they attach them-
selves to the branchial lamella? and are developed until
the fry have attained a length of about 11 mm. The
young of this fish thus lead a kind of parasitic life, a
commensalism from Avhich they liberate themselves,
when capable of an independent existence, by making
their Avay out through the expiratory tube of the mussel.

The spawning-season of the Cyprinoids occurs in
spring and summer, Avhen both sexes assume a brighter
and more handsome dress, and the males develop sharp,
tubercular excrescences on the scales, Avhich excrescences
fall off, hoAvever, simultaneously Avith the fading of the
coloration, as soon as the spawning-season is over.

To man these fishes are of no inconsiderable value
and utility. Most of them have a soft, Avhite, and pa-
latable flesh, and in Scandinavia, as on the Continent,
are the objects of lucrative fisheries. This is not the
case in England, Avhere fresh-Avater fishes in general are
little esteemed6, and the Cyprinoids in particular (Avith
the exception of the Carp) have a bad reputation.

“ Pallas (of the Crucian Carp), Zoogr. Ross. Asiat ., tom. Ill, p. 298.
b Cuv., Val., Hist. Nat. Poiss., torn. XVI, p. 13.

c Jahreshefte d. Ver. f. vaterl. Naturk. in Wurttemberg, 14 Jahrg., p. 121.
d At the Conference of Naturalists at Konigsberg, according to Siebold.
e Cyprinoiden Livlands, p. 87.
f Susswasserfische Mitteleuropas, p. 118.

g Zoologisclier Garten 1869, p. 257; 1870, p. 237; 1877, pp. 351 — 362.

h Day lays the blame of this on the English cook’s ignorance of the proper method of dressing fresh-water fish for table.

716

SCANDINAVIAN FISHES.

The form of the body is generally regular, showing
the compressed, fusiform outline most typical of the
Teleosts, whether it extends in a longitudinal direction,
as in our Gudgeons and Minnow, or rises vertically, as
in our Breams. In the great majority of the fishes of
this family" the body is covered with dense and firmly-
attached, more or less hard, cycloid scales, large or
small. The greatest external differences which here, as
in the Glanomorphs, afford the most useful characters
for the subdivision of the family, consist in the position
and dimensions of the fins, especially the dorsal and
anal. The back possesses only one fin (adipose fin
wanting), situated as a rule at the middle of the trunk,
but of a greater or less extent forwards or backwards,
in front of or behind the region of the ventral fins,
and sometimes set on the tail, above the anal fin,
which in its turn shows varying dimensions, greater
or less than those of the dorsal fin.

The Oyprinoids in general have a rather small
mouth, the position of which may vary from the tip of
the snout to the under surface thereof. The lips of
some (several East Indian) forms are strongly and sin-
gularly developed, sometimes funnel-shaped and fringed,
continuous or divided into lobes; but in our forms they
are smooth and hardly more fleshy than usual. In a
West Asiatic genus, Chondrostoma, which also occurs in
Southern and Central Europe, a cartilaginous sheath is
developed on the lower lip. In a North American ge-
nus, Acrochilus, a similar sheath appears both on the
upper jaw and the lower; and in the genus Labeo of
the Old World the sheath may appear on either or both
of the jaws. In many Cyprinoids the mouth is fur-
nished, as in the preceding family, with barbels, which
always belong, however, to the upper jaw and never
exceed two pairs; the most common and usually most-
developed is the barbel at each corner of the month.
lfhe margin of the upper jaw is formed by the toothless
intermaxillary bones, which are generally at least to
some extent (sometimes highly) protrusile. This does
not depend, however, as usual on any elongation of the

nasal processes, which are here comparatively short —
though often prolonged upwards by a cartilaginous con-
tinuation, which, when the mouth is closed (the upper
jaw drawn up), folds into a cavity between the tip of
the ethmoid bone and the rostral cartilage h — but in
most cases only loosely united to the rostral cartilage
in front of the ethmoid bone and above the more or less
cartilaginous head of the vomer. The intermaxillaries
are without erect, lobate process (cf. above, p. 463), a
growth which is all the more developed on the maxillary
bones behind them. The maxillary bones are besides
remarkable in most cases for their detorted form and
the short and generally thick head of their articular
knob. A distortion of the toothless dental part of the
lower jaw that reminds us of the Mugiloids (see above,
p. 330), occurs in several Cyprinoids (e. g. in the Bream
and Barbel, but not in the Ide), in which the upper
dental margin is thus turned outwards. We are also
reminded of the Mugiloids by the upright protube-
rance frequently present at the symphyseal tip of the
branches of the lower jaw. The most striking resem-
blance to the Mugiloids — depending on a similarity of
diet — - belongs, however, to the palate. The palatal
curtains (vela transversa) of the Cyprinoids are well-
developed, at least- in the upper jaw. The palatal roof
is lined with a mucous membrane, thickly covered with
papillae and arranged in longitudinal folds, which is
continued backwards, smooth but- with large gustatory
papillae, on the tumid, soft, cushion-like mass of muscles
and fat — the Carps tongue so highly prized by the
epicure — situated under the posterior part of the
cranium. Backwards and downwards from the body of
the occipital bone runs an osseous (pharyngeal) process0,
pierced at its base for the passage of a blood-vessel
( aorta abdominalis), the under surface of which process,
just at, the end of the said cushion, is shod in a de-
pression with a cartilaginous, more or less hard and
tumid disk, the so-called Carp-stone or pharyngeal car-
tilage ( Karpfen stein , la meule). Against this disk the
lower pharyngeal teeth, which are highly characteristic

" In a Southern European genus, Aulopyge , however, the body is entirely naked. It is also entirely or partially naked in three
Asiatic genera.

0 This cavity sometimes conduces in a remarkable way, even externally, to the singular form of the snout, which in an Indian ( Labeo
iiukta, Day, Fish. Ind ., p. 543, pi. CXXVIII, fig. 5) and a Sumatran species {Labeo — Scliismatorhynchus — heterorynchus, Blkr, All. Ichth.
Ind. Or. Neerl ., Cypr., p. 50, tab. IV, fig. 4) acquires a monstrous appearance, owing to the presence of a deep, horizontal, transverse
hollow in front of the nostrils.

c This process was formerly regarded by some as an hypapophysis, by others as an entire haemal arch; but- it arises, as Sage-
mehl (Morphol. Jabrb., Bd. XVI, p. 516) has at least shown to be probable, in a totally different way from these parts of the skeleton,
namely by the ossification of connective tissue which Sagem ehl regards as a remnant (trace) of the ligament (also pierced by the aorta) which
in the Characinoids unites the occipital and parasphenoid bones to the air-bladder. This same connexion we shall also find in the Clupeoids.

CARP-FISHES.

717

of the Cyprinoids, work in the mastication or dilaniation
of food. The form of the carp-stone, as well as the form,
number, and position of the teeth on the lower pha-
ryngeals, afford in most cases the most tenable charac-
ters for the definition of the Cyprinoid species and also,
to some extent, of the genera, and have been generally
employed for this object ever since Agassiz" and Hec-
kel* directed attention to this point. We shall often
return to these structures; here we need only remind
the reader that the said teeth on the lower pharyngeals,
which latter may be more or less falciform or more
like branchial arches0, are shed or renewed annually,
as JuRiNEd and (subsequently) Siebold0 have pointed
out. The upper pharyngeals are small, less remarkable,
and toothless. They lie above the sides of the aforesaid
cushion in the posterior part of the roof of the pharynx;
and in front of and outside them we find, in most
cases, but with great irregularity, distinct or covered
(glandular) pseudobranchia?.

The short oesophagus is recognisable internally by
the longitudinal folds of the mucous membrane. The
stomach is also short and shows very little, if any,
expansion. Its mucous membrane lies in zigzag folds
or is downy like velvet. These fishes have no dis-
tinct pyloric part or pyloric appendages, and the sto-
mach passes gradually into the intestine, the length of
which is exceedingly variable, for it sometimes forms
only two coils, sometimes five, before it runs back
to the vent. The intestine ends in a more or less
defined rectum, which is marked by longitudinal folds
of the mucous membrane. Often the whole digestive
canal lies imbedded in a more or less lobate and sub-
divided mass of liver and in a thick layer of fat. The
variations in the length of the intestinal canal are con-
nected with the diet: a greater proportion of animal
food is accompanied by a shorter intestinal canal. Hec-
kel divided the Cyprinoids7 on this account into two
groups: Macroentri , with long intestine — represented
in the Scandinavian fauna by Cyprinus and Carassius
— and BracJiyentri, among which we find all the rest
of our Cyprinoids. The gall-bladder lies on the right

side of the stomach and sends out a gall-duct into the
same; the spleen lies above or to the left of the be-
ginning of the intestine. The largest organ in the ab-
dominal cavity under ordinary circumstances (when the
ovaries are not extraordinarily swollen) is the air-
bladder. This organ is externally double, but internally
continuous, consisting of an anterior, shorter part and
a posterior, longer one, tapering behind, which usually
follows the curve of the upper Avail of the abdominal
cavity back to the very end of the cavity, and Avhich
communicates in front by means, of the pneumatic duct
with the oesophagus. The ovaries as avcII as the testes
are paired and closed, their discharging ducts sharing
as usual Avitli the urethra a common aperture just be-
hind the mouth of the rectum. The kidneys lie along
the under surface of the spinal column and are generally
most developed at the contracted part of the air-bladder.

The most remarkable peculiarities of the skeleton
are the alterations Avhich the first four abdominal ver-
tebrae undergo in connexion Avitb the development and
function of the so-called acoustic bones. The bodies of
these vertebrae coalesce, as Ave have mentioned above,
more or less firmly with each other, the boundaries
between them being usually marked, hoAvever, by distinct
sutures, even if the intervertebral cartilage has dis-
appeared. Thus, in the Tde for example, all four are
divided from each other Avith almost equal distinctness;
and the intervertebral cartilage' is still present between
the bodies of at least the third and fourth vertebra?,
the neural arches of Avhich, on the other hand, are more
firmly united by a suture. In the Chub as Avell as in
the Bream the bodies of the second and third vertebrae
are almost perfectly confluent. In the Barbel the body
of the first vertebra is confluent Avith that of the se-
cond, and that of the third Avith that of the fourth.

The body of the first vertebra is always the small-
est. Its transverse process, which is also considerably
smaller than that of the second vertebra, stands, like
the latter, straight out in a horizontal direction and
originates from the body, also like the latter, Avithout
perceptible suture. The third vertebra is apparently

° Mem. Soc. Sc. Nat. Neuchatel, vo], 1 (1836), p. 36.

b Abbildungen und Beschreibungen der Fisclie Syriens nebst einer neuen Classification irac] Karakteristik sammtlicher Gattungen der
Cypfinen, Stuttgart 1843 (reprinted from Russegger’s Reisen, B. 1, Tli. 2).

c Cf. above, p. 631, on the lower pharyngeals and teeth of the File-fishes.

d Mem. Soc. Phys., d’Hist. Nat., Geneve. Tom. I (1821), p. 20. Bloch ( Fisclie Deutschland s, Th. 1, p. 47) also suggested the
possibility of this.

e Siisswasserf. Mitteleur ., p. 82.

1 Russeggers Reisen , 1. c., p. 1001.

718

SCANDINAVIAN FISHES.

without transverse processes; but the fourth, when its
body is distinct from that of the third, not only pos-
sesses a rib-like transverse process on each side, united
by a suture to this side and both set and curved in
the same direction as the ribs, only shorter and more
terete than they, but also bears on its under surface
an inner, lamellar process, set transversely and united
to the body of the vertebra and to the transverse
processes, pierced at the base (like an hypapophysis),
and curved backwards and downwards behind the tip of
the large pharyngeal .process of the occipital bone. To
the posterior surface of this inner process of the fourth
vertebra is attached the centre of the anterior end of
the air-bladder, and the passage in its base receives the
aorta and the anterior end of the kidneys. According
to Sagemehl" this process also belongs to the fourth
vertebra in Hydrocyon (a Characinoid) and originates
from the ventral and partly from the lateral sides of
the body of this vertebra6. In the Barbel, where, as
we have mentioned, the body of the fourth vertebra
coalesces with that of the third, we see, however, that
the roots of this process lie on the anterior part of this
composite body, thus within the limits of the third ver-
tebra. A removal in a backward direction thus seems
to have taken place in the rest of our Cyprinoids, an
assumption which finds further ground in the insertion
of the hindmost so-called acoustic bone ( malleus ) between
this process and the true transverse process of the fourth
vertebra to join the wall of the air-bladder. Here, as in
the Sheat-fish (see above, p. 699), we explain the malleus
on each side as the transformed rib of the third vertebra.
In the Cyprinoids it is crescent-shaped or, rather, like
the blade of a headsman’s axe, and at the inner (con-
cave) margin is set the process whereby it articulates
with the side of the body of the third vertebra. Its
anterior end, which is united by a ligament to the so-
called stapes, projects above the base of the transverse
process of the second vertebra. This last vertebra sends
out to the ligament just mentioned the bone which has
been called the incus , a bonelet bifid at the base and
with one branch articulating in a hole in the body of the

second vertebra and the other united by a ligament to the
same bone. The incus on each side is explained as the
transformed ribc and neural arch of the second vertebra.
The tivo anterior among the so-called acoustic bones,
the stapes and claustrum, Avhich lie close to each other
on the covering membrane of the atrium sinus imparls
(see above, p. 699), are partly foliate in form, and are
explained as representing on each side the othenvise Avant-
ing neural arch and spine of the first vertebra. The se-
cond vertebra is apparently Avithout neural spine (upper
spinous process), but its place is taken by a covering
bone above the spinal canal betAveen the occipital bone
and the large neural spine of the third vertebra, Avhich
spine is usually strengthened by coalescence Avith the
neural spine of the fourth vertebra.

In the structure of the head Ave shall here remark
only the comparatively perfect development of the or-
bital ring. Not only do the ordinary (here 4 — 8) sub-
orbital bones surround the eye behind, beloAv, and in
front; the eye is also protected above by a supraorbital
bone, a covering bone on the frontal bone of each side.

The skeleton of the Cyprinoids, Avhich as a rule
contains a moderate number of vertebrae (40d — 50), is
further distinguished by the high and upright, anterior
and upper, articular processes (zygapophyses) of the ab-
dominal vertebrae, especially in the forepart of the trunk,
where the top of each of these processes meets the base
of the neural spine of the vertebra immediately in front
or the upper part of the neural arch of this vertebra.
The shoulder- girdle is strong. The incurved anterior mar-
gin of the clavicle may sometimes, as in Labeoe, be de-
veloped into a disk so broad that only a narroAv passage
is left for the oesophagus. The coracoid bone is also as a
rule comparatively broad, and the precoracoid bone
ascending from the upper margin thereof, is bifid at the
top, one branch meeting the clavicle, the other the sca-
pula. The pelvic bones are elongated, in front bifid,
sometimes for the greater part of their length, behind
united by cartilage or a suture. From a morphological
point of vieAv the articulation of the ventral fins is in-
teresting, as Davidoff has shoAvn7, on account of its

“ Morphol. Jalirb., Bd. X (1884), p. 55.

b Sagemehl explains it as a transformed pair of ribs belonging to this vertebra. Sorensen, who calls it os suspensorium , ascribes it
to an ossification of the wall of the air-bladder. Cf. Sagemehl’s explanation of the pharyngeal process of the occipital bone (see above).

c Ligamentous ossification, according to Sorensen, Om Forbeninger i Svommeblceren etc., Dsk. Vid. Selsk. Ski-., 6:te Rsekke, Naturv.,
Math. Afh., B. 6, No. 2, p. 41 (sep.). The said paper did not reach me, unfortunately, until this sheet was in the press.
d In Barbus maculatus , according to Gunther, 30.
e Gunther, Cat., VII, p. 47.

f Morphol. Jalirb., VI (1880), p. 404, taf. XXI, fig. 4.

CARP-FISHES.

719

retention in a cartilaginous form of a row of four basal
bones" — a point which reminds us of more primitive
types (the Ganoids) — the innermost of which even pos-
sesses the same form as in Amia and Lepidosteus. Be-
hind, from the symphysis, each pelvic bone projects on
the inner side of the insertion of the ventral tin in a
process, directed straight back or curving outward to
the side and serving as a point of attachment for a
muscle coming from the base of the anal fin. The three
branchiostegal rays are as a rule strong, of a broad,
ensiform shape; and the urohyoid bone, a triangular,
horizontal disk with the point turned forward and 'with
a strong and high osseous ridge, highest behind, on the
upper surface, reminds us most strongly of the cor-
responding bone in the Siluroids.

The systematic arrangement of the Cyprinoids in-
volves many difficulties, partly on account of the great
wealth of forms, partly in consequence of the more than
usually strong proclivity of these fishes for cross-breed-
ing between different species, even between species be-
longing to genera recognised as distinct. The Cyprinoids
have been named after the island of Cyprus, the ancient
sanctuary of Venus6, and this name has arisen from
observations of the fecundity and vivacity of these fishes
when spawning. They crowd together in wild tumult
to spawn and mingle with each other, seeking the same
spawning-places and often having the same spawning-
seasons. Cross-breeds between the Carp and the Crucian
Carp have long been known. Schonevelde (1624) speaks
of such hybrids in the Elbec, Marsigli (1726) tells us
from the Danube that the fishermen of this river, who
called these forms Sittich karp f en, explained them as
hybrids'* *, and according to Borner (1781) the fishermen
of Silesia had made the same observation", namely that
these forms are produced by cross-breeding between the
Carp and the Crucian Carp, when Crucian Carp are
incautiously allowed to enter the culture-ponds for Carp,
and that for this reason the breeders of Carp carefully

avoided buying fry from such ponds as were suspected
also to contain Crucian Carp. These observations were
regarded, however, by zoologists in general merely as
surmises — ichthyologists described these hybrids as a
distinct species, and Hecket/ in 1853 made them the
type of a distinct genus — until DybowskC in 1862 and
Sieboli/ in 1863 gave the question a different turn.
Siebold also showed' that there was good reason to
regard the Letter of German writers (Bloch’s Cyprintis
Buggenliagii) — a fish which was said to lead the way
for the shoals of Bream, and whose capture was there-
fore supposed to be a good omen of an abundant take
— as a hybrid of two kinds between the Bream genus,
Abramis ( brama and blicca ), on the one hand and the
Roach and Rudd genera, Leuciscus and Scardinius, on
the other; and also to interpret the Hacliette ( Alburnus
dolabratus ) of French authors as a hybrid ' between the
Chub ( Leuciscus ceplialus ) and the Bleak ( Alburnus Juci-
dus). Jack el 6 has added several forms from Bavaria
to this list of hybrids, one of which, that between the
Roach and the Rudd, may possess a special interest for
our fauna. As yet, however, to the best of our know-
ledge, only the first of these hybrids, that between flic
Carp and the Crucian Carp, has been proved by breeding
experiments really to be of a hybrid nature. A great
held of investigation is still open here to the piscicul-
turist, with whom it lies to solve these questions; but
we know enough already to prevent any surprise at
the uncertainty in the systematic arrangement of the
Cyprinoids.

The Scandinavian fauna, it is true, contains only a
few of the variations that have ranked as types for the
subfamilies and genera hitherto established within the
Cyprinoid family. Still, we possess a sufficient number
to enable us to see among them the most important
extremes in the differentiation of the family. From the
most harmonious Roach type the form-series within the
Scandinavian fauna proceed in two directions, on the

a Davidoff calls these bones radii.
b Kurcoig was also a name of Venus.

c Ichthyol. Slesv. Hols., p. 34: “medii Carasi ob id dicti, halb Harass vel Karpffkarass, quod e Caraso et Carpa compositi videantur.”
d “Similitudine inter Cyprinum et Carassium mediat, narn ex ovis Cyprini, quantum Piscatores asserunt, et semine, ved lacte Carassi,
aut e contra progeueratur,” Danubius Pannonico-mysicus, tom. VI, p. 61.
e Zoologies Silesiacce Prodromus, p. 205.
f Verh. zool. bot. Ver. Wien, Bd. II, p. 29.
g Vers. Monogr . Cypriniden Livlands, p. 55.
h Siisstvasserf. Mitteleur., p. 94.

* L. c., pp. 145 — 152.

J Sieb., 1. c., p. 167.

k Abb. zool. mineral. Ver. Regensb. 1864 and 1865; Zool. Garten 1866; Fische Bayerns.

Scandinavian Fishes.

91

720

SCANDINAVIAN FISHES.

one hand to an elongation of the base of the dorsal tin, but
with this base more and more reduced the nearer the
series approaches to Leuciscus, on the other hand to almost
as great an elongation of the base of the anal tin. By
combining the expressions of these changes we find one
division, the subfamily of the Carps, corresponding to
Klein’s" genus Cyprinus , in all the forms of which the
dorsal fin is at least 1/3 (up to 4 1/2 times) longer than
the anal, another division, within which the base of the
dorsal fin measures at most about 130 %h , but some-
times no more than 70 % of the length of the base of
the anal fin, and a third division, within which this
percentage lies between 60 and 23. The last two di-
visions, which comprise the majority of our Cyprinoids,
may also be defined by a comparison between the lengths
of the lower jaw and of the base of the anal fin. In
the former group, the subfamily of the Roaches, Klein’s0
genus Leuciscus , the length of the lower jaw measures
as a rule perceptibly more than half of the base of the
anal fin; but the proportion is subject to a change of
growth ,l which approximates this group to the latter
one, essentially corresponding to Klein’s0 genus Brama ,
the subfamily of the Bleaks and Breams, within which
division the length of the lower jaw may indeed sink
to about Vs of the base of the anal fin, but may also
rise at least to 48 % thereof7. On these grounds we
therefore distinguish among all the forms that in this
respect have attained a fixed character77 — which is the
case in all the Scandinavian species — three subfamilies:

A : Dorsal fin considerably longer than
the anal, length, of the base of the
latter at most 75 % (sometimes only

21 %) of that of the base of the former Subfamily Cyprinince.
B: Dorsal and anal fins of fairly equal
length, minimum length of the base
of the latter at least 77 % of that of
the base of the former, minimum length
of the base of the former at least 70 %

of that of the base of the latter Subfamily Leuciscince.

C: Anal fin considerably longer than the
dorsal, length of the base of the latter
at most 60 % (sometimes only 23 %)

of that of the base of the former.. Subfamily Abramidince .

The knowledge of the Scandinavian Cyprinoids has
been of fundamental service in the determination of the
other European forms of this family, and the first edition
of Scandinavian Fishes was of great importance from
this point of view. Fries and Ekstrom there prefaced
their diagnosis by a general summary of what their pre-
decessors, from Artedi, had accomplished to the same
end, and this summary is even now not without interest.

Artedi, they wrote, who described with an accuracy
unparalleled in his time all the indigenous piscine spe-
cies which he had the opportunity of seeing and ob-
serving in nature, adopted h sixteen Swedish species of
the genus Cyprinus:

Iden (the “Ide”), in modern times Leuciscus idus ,

Sarfven (the Rudd), ,,

Morten (the Roach), ,,
Stdmmen (the Dace), ,,

Sandkryparen (the

Gudgeon), „

Aspen (the “Asp”), ,,
LAjan (the Bleak), ,,

Virnban (the

“Zarthe”), ,,

Bjorknan (the

White Bream), ,,

Braxen (the Bream), „
Fliran (the “Zope”), ,,

Scardinius ei \j tin -
oplitlialmus,

Leuciscus rutilus ,
Leuciscus grislagine ,

Gobio fluviatilis ,
Aspius rapax ,

Alb ur nus lucid us,

Abramis virnba ,

Abramis blicca ,
Abramis brama,
Abramis ballerus,

Faren, which has proved to be merely a nominal species,
Karpen (the Carp), in modern times Cyprinus carpio,

Sutaren (the Tench), ,, ,, ,, Tinea vulgaris,

Rudan (the Crucian

Carp), ,, ,, ,, Cyprinus carassivs ,

Elritsan (the

Minnow), ,, ,, ,, Phoxinus aphya.

a Hist. Pise. Nat., Miss. V, p. 58.

h According to our measurements at most 123 % ; but Kr0yer mentions a Roach in which the base of the dorsal fin measured 129 % of that of the anal.

c L. c., p. 64, with the exclusion, however, of the Bleaks.

d In large Rudd the percentage may sink to 51.

e L. c., p. 61, with the exclusion, however, of the Rudd and Tench.

f In the above-mentioned hybrid, the German Leiter , which also occurs in Scandinavia, the length of the lower jaw is sometimes
rather more, sometimes less than half the length of the base of the anal fin.

<J In genera which presumably stand nearer the original forms of the family, this character has not acquired the same degree of con-
stancy as in those which have advanced further in the differentiation of form. So, for example, the above taxonomic relation seems to give
a sexual difference in the Bitterling ( Rhodens amarus ), to judge by the 6 specimens which the Royal Museum has received through Professor
Siebold from the River 111 at Strasburg. In the four females among these specimens the length of the base of the anal fin is at most 75^
(70 — 75 %) of that of the base of the dorsal fin, while in the two males this percentage is respectively 77 and 81 %.

h Descr. Spec. Pise., pp. 6 — 31. Together with the excellent descriptions the Swedish names may serve in the first place to deter-
mine these species.

CARP -FISHES.

721

All these forms had retained their position of recog-
nised species until the time when the first edition of
Scandinavian Fishes appeared. Fries and Ekstrom
then advanced the opinion that the so-called Faren was
a young- Bream" to which Artedi had applied a Swe-
dish name that is used throughout the Sodermanland
coast of Lake Malar for the “Zope”.

In the second edition of his Fauna Suecica Linnaeus
adopted as Swedish all Artedi’s species with the excep-
tion of the Gudgeon, and added only one new, but very
characteristic species, the Skdrbraxen , Cyprinus (now
Pelecus ) cultratus. We must not suppose, however, that
Linnaeus understood by his Cyprinus BiorJma the same
species as Artedi: — -this Linnsean form, as Fries and
Ekstrom have shown, is a nominal species made up of
the White Bream and the Zope. Linnaeus is also the ori-
ginator of another nominal species, briefly alluded to in
the Sy sterna Natures'1 under the name of Cyprinus Idharus,
as occurring in the Swedish lakes, and subsequently
introduced by Retzius into the Scandinavian fauna.
From the short diagnosis given by Linnaeus it is hardly
possible to decide with absolute certainty whether this
nominal species is identical with the Ide or the Roach,
though it is most probably the same as the former.

In the third edition, begun by Retzius in 1800,
of the Fauna Suecica , 20 species of the Carp genus are
adopted, namely the same 16 as in the second edition (the
Bjorkna and Faren thus in their Lin mean signi fication,
and not as in Artedi), together with Artedi’s Sandkry-
■ pare ( Cyprinus Gobio), and three new species are added:

Cyprinus Dobula,

,, Idharus ,

,, Phoxinus.

Among these three Fries and Ekstrom recognised
the specific rank of the first alone; of Idharus they had
already declared their opinion, and with regard to Pho-
xinus they pointed out that the specific character as-
signed to it did not justify its recognition as a distinct
species from Linnaeus’s Cyprinus Aphya.

In 1832 Nilsson gave in his Prodr omus Ichthyo-
logies Scandinavian a scheme of a Scandinavian piscine
fauna, which possesses the merit, in addition to several
others, of having established the definitions of the spe-
cies on more trustworthy diagnostic grounds. Within
the Carp genus ( Cyprinus ) the said author includes 20
species, namely the original 1 6 of Artedi, which we
there find restored to their first signification and more
fully diagnosed than before; furthermore C. cultratus ,
Dobula , Phoxinus , and a species now introduced for the
first time into the Scandinavian fauna, Bloch’s Cypri-
nus Gibelio. The last-mentioned species was, however,
shown by Ekstrom to be merely a variety of the
Crucian Carp.

In the description published by Ekstrom of the
piscine species of MorJco there also occurred a mistake
which he hastened to correct in Scandinavian Fishes , as
it had given rise to another nominal species, Cyprinus
microlepidotus. On continued observation of the growth
of the Cyprinoids he had convinced himself that this
fish, though very different both in form and coloration,
Avas nothing more than the Ide at an early stage of
groAvth .

The first edition of Scandinavian Fishes thus con-
tained 17 Scandinavian species, determined with cer-
tainty, Avithin the Cyprinoid family. In Kroyer’s me-
ritorious Danmark's Fiske (vol. 3, pp. 289 etc.) and in
Nilsson’s Skandinavisk Fauna ( Fiskarna , pp. 281 etc.)
the number of species is the same. This number Avas
first increased in 1871 by a neAV addition to our fauna,
Avhen Lilljeborg'- described SAvedish specimens of Leu-
caspius delineatus, specimens to Avhich his attention had
been called by Mr. Ahlbom, Collector of Customs at
Landskrona. In 1877 Doctor Feddersen found in Jut-
land a species neAV to Denmark, Abramis bipunctatus.
We thus knoAv at present 19 or — as the Goldfish may
claim a place in the character of a domesticated spe-
cies — 20 Scandinavian species of Cyprinoids, together
Avith the above-mentioned hybrids.

a As we shall show later on, Artedi (1. c., p. 23) has really described an Abramis ballerus or “Zope" under the Swedish name
of 11 Faren".

6 Ed. X, tom. I, p. 324.

c Of vers. Vet. Akad. Forh. 1871, No. 7, p. 815, tab. XVII, A.

722

SCANDINAVIAN FISHES.

Subfamily CYPRININ IE.

Dorsal fin much longer than the anal ( length of the latter at most 3/4 of that of the former ), its distance from
the tip of the snout, as a rule less than (. sometimes about equal to) tivice the length of the head. Among the
simple rags at the anterior margin both of the dorsal fin ( 3 or 4) and the anal (2 or 3) the hindmost is gene-
rally hard as bone and spiniferous at the hind margin. Lateral line on the tail occupying the middle of the
depth of the body. Ventral margin terete. Mouth as a rule furnished with barbels. Pharyngeal cartilage as a
rule triangular , with the anterior margin straight or slightly convex. Length of the intestinal canal (■ including
the stomach and oesophagus) as a rule severed times greater than the length of the body.

We have given the first place among these charac-
ters to the one which seems to us to be the most dis-
tinct external token of the direction of development
followed by the subfamily: a long dorsal tin, developed
especially in front, as opposed to a short anal fin. In
the Scandinavian fauna this character is not impaired
by a single exception; but in consequence of another
developmental change, by which the relative length of
the head decreases with age, it sometimes happens that,
in large Crucian Carp for example, the length of the
head is somewhat Jess than half the distance between
the dorsal fin and the tip of the snout, while the Tench
also resembles, as a rule, in this respect the following
subfamily. The systematic significance of the barbels
we have already seen within the families both of the
Sheatfish and the Loach; but here it sometimes happens
that one species of a genus ( Cyprinus ) possesses them,
while others are without them: — and yet these species
interbreed with each other so freely that it seems far
too artificial to separate them into distinct genera. With

its present definition, however, the subfamily includes
all the Scandinavian Cyprinoids that are furnished with
barbels. The same irregularity affects another charac-
ter, the serration of the last undivided and osseous ray
in the dorsal and anal fins: this may be present in
some species of a genus {Barbus), but wanting in others,
while in the Scandinavian fauna it is wanting in two
genera of this subfamily ( Gobio and Tinea), which in
other respects as well (the form of the pharyngeal
cartilage and the length of the intestinal canal) show
signs of a transition to the following subfamily.

The genera that occur within the limits of the
Scandinavian fauna, may be distinguished as follows:

A: Length of the dorsal fin more than twice

that of the anal. Genus Cyprinus.

B: Length of the dorsal tin less than twice
that of the anal.

a: Least depth of the tail less than 2/3

of the length of the head..., Genus Gobio.

b: Least depth of the tail more than -/.

of the length of tire head. Genus Tinea.

Genus CYPRINUS.

Base of the dorsal fin more than tivice that of the anal fin.
ceptionally 16), in the anal fin cd most 7 ( exceptionally 8).
strongly osseous and behind spiniferous.

than 40— thick and hard.

As Gunther" has remarked, rve are fully justified
in retaining the generic name of Cyprinus in its ori-
ginal signification — in the first place as a name for

Branched rays in the dorsal fin at least 17 (ex-
Both these fins with one of the anterior rays
Scales large — their number in the lateral line less
hard.

the Carp — , in spite of the fact that neither Artedi
nor LinNyEUS has always been so exact in his syste-
matic enumeration of the Cyprinoids as to range the

Cat. Brit. Mus., Fish., vol. VII, p. 25, note.

CARPS.

723

Carp first. Bleeker’s objection to this course" — that
the Cyprinus of Artedi and Linnaeus was strictly a
synonym of the Leuciscus of modern writers — has not
gained any adherents among ichthyologists.

The genus, which is indigenous to Asia and Europe,
but has been transported by human agency throughout
the civilised world, is by no means rich in forms —
Guntiier recognises only five species — but these spe-
cies have been transformed by domestication into a
multitude of varieties and often possess monstrously
developed organs. Nilsson established6 for the Crucian
Carp a subgenus, Carassius (without barbels), formed
on the principle resorted to by Cuvier of misusing for
this purpose the Linnaean specific names. But this sub-
division of the genus is of far too little importance
— in spite of the fact that in some authors Carassius
has been recognised as a distinct genus — partly in
consideration of the intermediate position in form of
body occupied by the Goldfish, partly on account of

the ease with which the Crucian Carp and the Carp to-
gether beget progeny capable themselves of reproduction.

The forms belonging to the Scandinavian fauna may
be distinguished by the following external differences:

A: Four barbels, two on each side of the
mouth.

a: Length of the postabdominal part of
the body about equal to the greatest
depth of the body — greater than it
or at least more than 4/5 thereof ___ Cyprinus carpio.

Ik Length of the postabdominal part of
the body less than or equal to 3/4 of
the greatest depth of the body

(Hybrid Carpio Kollarii).

B: Mouth without barbels.

a: 5 or 6 scales in an oblique trans-
verse row between the lateral line
and the anterior part of the base of

the dorsal fin Cyprinus auratus.

b: 7 or 8 scales in an oblique trans-
verse row between the lateral line
and the anterior part of the base of

the dorsal fin Cyprinus carassius.

THE CARP ( SW. KARLEN).
CYPRINUS CARPIO.

Plate XXXI, fig. 1.

Two pairs of barbels , one (small) at each upper corner of the snout, one (larger) just behind each corner of the
mouth. Pharyngeal teeth blunt (molars), more or less cylindrical, with a more or less multistriate grinding,
surface on the top: 1, 1, 3 — 3, 1, 1. Greatest depth, of the body about 1/i of its length: distance between the
foremost point of the insertion of the ventral fins and the beginning of the anal fin greater than the said depth

or at least than 4/5 thereof.

a

Fig. 178. Pharyngeal bones and pharyngeal cartilage of Cyprii
seen from outside (the left); c, pharyngeal cartilage seen f:

carpio ; natural size; a, left pharyngeal seen from above; b. the same
in front and below, the lower (posterior) end pointing upward.

R. hr. 3 (4); D.

3~4-. A F_
17—22 ' ' 5 — 6’ ' 14—17’ ‘ 6

-8(9)’

■6(7)

C. sc + 1 + 17 + 1 4- x\ Lin. lat. (34)36 — 39; Lin. tr. — — hr — 1

(4)5 — 6(7)

Vert. 36—37.

Syn. Kvnqivog et KvTTQiardg, Aristot., Athen., cett. vide Ronde-
j.et, ( Cyprinus ) Univ. Aquat. Hist., part. II, p. 150.

carpe, Belon, Nat., Divers. Poiss., p. 267. Cyprinus,
No. 1, Art. Syn. Pise., p. 3; No. 13, Spec. Pise.,
p. 25.

a Ail. Ichth. Ind. Or. Ne'erl., tom. Ill, p.
h Prodr. Ichth. Scand., p. 32.

73.

724

SCANDINAVIAN FISHES.

Cyprinus Carpio , Lin., Syst. Nat., ed. X, tom. I, p. 320; Bl.,
Naturg. Fiscli. Deutschl., part. I, p. 92, tab. XVI et XVII;
Betz., Fn. Suec. Lin., p. 354; Pall., Zoogr. Ross. Asiat.,
tom. Ill, p. 289; Nn.ss., Proclr. Ichth. Scancl., p. 33;
Schagerstr., Physiogr. Sallsk. Tidskr., p. 295; Cuv., Val.,
Hist. Nat. Poiss., tom. XVI, p. 23; Hckl, Russegg. Reis.,
part. II, p. 1013; Kr., Damn. Fisk., vol. Ill, p. 290;
Nilss., Skand. Fn., Fisk., p. 284; Hckl, Kn., Sussioasserf.
Oesterr. Mon., p. 54; Sieb., Susswasserf. Mitteleur., p. 84;
Canestr., Arcliiv. Zool., Anat., Fisiol., vol. IV (1866), p. 64;
Gthr, Brit. Mus. Cat., Fish., vol. VII, p. 25; Coll., Fork.
Vid. Selsk. Chrnia 1874, Tilliegsh., p. 179; Hessel, U. S.
Comm. Fish., Bep. 1875 — 76, p. 865; Feddeks., Naturh.
Tidskr. Kbhvn, ser. 3, vol. XII, p. 82; Bncke, Fiscli. Fischer.,
Fiscliz. 0., W. Preuss., p. 106; Handb. Fiscliz., Fischer.
(M. v. d. Borne), p. 117; Mor., Hist. Nat. Poiss. Fr.,
tom. Ill, p. 368; Day, Fish. Gt. Brit., Irel., vol. II, p. 158,
tab. CXXIX, fig. 2; Mob., IIcke, Fiscli. Osts., p. 105; Lillj.,
Sv., Norg. Fn., Fisk., vol. Ill, p. 129; Fatio, Fn. Vert.
Suisse, vol. IV, part. I, p. 171.

Rex Cyprinorum 4- Cyprinus nudus ( alepidotus ), Bl., 1. c.,
part. Ill, pp. 131 et 178.

Cyprinus carpio + C. coriaceus + C. specularis + C. rubro-
fitscus + C. nigroauratus + G. viridi-violaceus, Lacep., Hist.
Nat. Poiss., tom. V, pp. 504, 528, 530, 547.

Cyprinus carpio + C. regina + C. elatus, Bonap., Fn. Pal.,
Peso. tab. 108

Cyprinus liungaricus, Hckl, Ann. Wien. Mus. vol. I, part. 2,
p. 222, tab. 19, fig. 1, C. acurninatus, Hckl, Kr., 1, c.,
p. 58.

Cyprinus Nordmanni + C. flavipinnis (ex Kuhl et V. Hass.)

4- C. vittatus, Cuv., Val., 1. c., pp. 66, 71, 72.

Cyprinus bithynicus , Richards., Proc. Zool. Soc. 1856, p. 372
(vide Gthr).

Cyprinus atrovirens + C. flammans + C. sculponeatus + C.
hybiscoides, Bichards., Ichthyol. China, Bep. Brit. Assoc.
1845, p. 287, cett.

Cyprinus hcematopterus 4- C. melanotus 4- C. conirostris, Schleg.

Fn. Japon., Pise., pp. 189 — 191, tab. XCVI et XCVII.
Cyprinus chinensis 4- C. obesus, Basil., Nouv. Mem. Soc.
Nat. Mosc., tom. X, pp. 227 et 228, tab. I, fig. 2, tab. II,
fig. 3.

All these numerous names — and still more, which
we have refrained from giving, as we were unable to
verify the quotations — have been applied in descrip-
tions to the Carp with its many varieties. It has been
treated from time immemorial, we may almost say,
as a domestic animal, and this treatment has affected
its external form. Several of the above synonyms
probably denote hybrid forms, though it is difficult to
give a positive opinion on this head, as the dentition
of the pharyngeal bones has not been specially de-
scribed.

In Germany and Eastern Europe the Carp attains
a length of about l1/i (Heckel and Kner) or l1/ 2
(Benecke) metres and a weight of about 30 — 33 kgm.
(Pallas and Hessel). In Scandinavia it has never
been known to attain a size at all approaching this,

o

and even the account given by Astrom" of three Carp
respectively 23, 24, and 27 Swedish pounds (10 — 1 1 1/2
kgm.) in weight, which were said to have been caught
“about” 1850 in the Hoje River at Lomma, is evidently
based on hearsay, though it cannot be considered to
involve any impossibility. These Carp would at that
weight have been probably 80 — 88 cm. long.

The body is fairly elongated and thick, the greatest
depth, which lies at the beginning of the dorsal fin,
measuring (in specimens 4 — 4V2 dm. long) about 25 —
27 % of the length, and the greatest thickness (just
behind the head) about half the depth or a little more.
Young specimens are in general somewhat deeper6, but
thinner: the thickness increases with age more than the
depth. The dorsal and ventral profiles are fairly re-
gularly arcuate, the former, however, more so, espe-
cially in front. The dorsal edge is somewhat sharpened
just in front of the beginning of the dorsal fin; the
ventral edge, on the other hand, is more terete or even
somewhat flattened. The contour of the tail shows at
the ventral margin a slight break to receive the base
of the anal fin. The least depth of the tail measures
about 12 — 11 % of the length of the body and is as
a rule more than 1/3 greater than the length of the
base of the anal fin or about 1/2 (52 — 47 %) of the
length of the head.

The length of the head varies in adult Carp (3 —
5 dm. long) between about 23 l/2 and 22 1/2 % of that
of the body0; in young specimens it is comparatively
somewhat greater, and it seems generally to be less in
the females than in the males. The head is of a com-
pressed conical form, with sharply rounded snout, and
quite strongly reminds us of a pig’s head. The fore-
head is fair])7 broad and somewhat convex (the breadth
of the interorbital space equal to or somewhat greater
than the length of the snout), but shows between the
eyes a depression, bounded within the convex supra-
orbital regions by two shallow grooves, which converge

a Nagra iakttagelser rorande de vertebrerade djur, som fbrelcomma , i trakten af Lomma, disp. Lund 1859, p. 26.

6 According to Canestrini’s measurements the depth of the body measures 27 — 30 % of its length (including the entire caudal fin),
while this length increases from 128 to 200 mm.

c When the length of the body is measured, as usual in this work, to the end of the middle caudal rays. If we take into account
the entire length of the caudal fin, the above percentages become respectively 21 1/2 and 20.

CARP.

in front towards the snout. This depression is most
distinct in old specimens, in which the externally bare
hind parts of the frontal bones too are more distinctly
marked with radiating striae. The eyes are round (with
the longitudinal diameter slightly greater than the ver-
tical) and are set rather high, by far the greater part
of them lying above a horizontal line from the tip of
the snout to the middle of the caudal fin. In speci-
mens between 3 and 5 dm. long their longitudinal dia-
meter measures about 15 — 14 % of the length of the
head; in younger Carp they are as usual comparatively
larger, their longitudinal diameter, when the body is
2 dm. long, being about 1/5 of the length of the head,
when the body is 3/4 dm. long, about 1/i of the same.
The nasal cavities lie just, in front of the upper an-
terior corners of the orbits. The two nostrils on each
side are fairly large and round, the diameter of the
posterior being about equal to its distance from the
orbit. They are set close to each other, the anterior
obliquely on the inner side of the posterior; and the
margin of the anterior nostril is raised behind in an
obliquely cut, canaliculate form, so as to compose a
lid with which the aperture may be closed. The distance
between the two anterior nostrils is somewhat less than
their distance from the middle of the tip of the snout,
but about equal to half the breadth of the interorbital
space above the centre of the eyes. The length of the
snout varies, however, quite considerably, from 1/3 of
the length of the head in young Carp to about 3D %
thereof in old. The mouth is fully terminal, the cleft,
when the mouth is closed, slightly ascending, and the
gape, when protruded and wide open, of a rounded
quadrangular form. The lips are fleshy; the upper lip
is double, the maxillary bone being also furnished with
a dermal fold, in which the two pairs of barbels have
their origin; the lower lip hangs down along the sides
of the lower jaw in front of the corners of the mouth.
The barbels are flattened at the base and vary con-
siderably in length; the lower (at the corners of the
mouth) may be half as long again, according to Fatio,
as the diameter of the eye, but are usually shorter and
in young Carp much shorter than the diameter of the
eye; the upper (anterior), which are set about half way
along the maxillary bones or a little higher, are only
half or one-third as long as the lower; but we must
not forget that during life these barbels are capable of
extension and contraction. The length of the upper
jaw, measured from the middle of the tip of the snout

725

to the hind extremity of the maxillary bones, is in
adult Carp about 27 or 28 %, and the length of the
lower jaw 31 — 36 % of the length of the head. The
gill-openings extend from the upper posterior corner
of the operculum to the isthmus, about to the perpen-
dicular from the articular knob (upper anterior corner)
of the operculum. The branchiostegal membranes are
below united to the isthmus at a distance from each
other which in old specimens is equal to the longi-
tudinal diameter of the eyes. The branchiostegal rays
are 3 in number, in accordance with the general rule
within the family; but Fatio on one occasion found 4
rays in each membrane. The operculum is obliquely
quadrangular (rhomboidal), with the lower posterior
angle rounded and obtuse. The anterior margin is the
longest, being about equal in length to the postorbital
part of the head, or to about half the entire length of
the head. The lower margin is slightly indented in a very
elongated S-shape or even straight. The surface of the
operculum is striated by granulated ridges with inter-
jacent. grooves, radiating backwards and downwards from
the articular knob. The crescent-shaped suboperculum,
which coasts the inferior margin of the operculum, is
distinctly (about 1/3) longer than the lower jaw. The
triangular interoperculum, pointed at the top, tills the
angle between the two bones just mentioned and the
lower part of the preoperculum, and coasts the inside
of the horizontal (lower) arm of this last bone forward
to the point of union with the articulation of the lower
jaw. The hind angle of the preoperculum is obtuse
and rounded. The cheeks are soft; but the space be-
tween the eye and the uppermost part of the preoper-
culum is filled by the posterior bones of the orbital
ring. The suborbital bones are also thick and firm,
striated like the operculum. The gill-rakers are short
and rather scattered (18 — 20 in the outer row on the
first branchial arch), consisting of pointed triangular
disks with dermal papillte on the inner margin. The
lower pharyngeals (fig. 178), the mucous membrane of
which is fringed at the outer superior margin with
similar, but. smaller gill-rakers, 22 in number, are fur-
nished within this margin as a rule with five teeth each;
but while the teeth a, re being shed, one or two of these
may be wanting or may lie hidden or loose in the
mucous membrane. The largest teeth are 3 in number,
set in an inner row. The first of them is bluntly conical,
with a greater or less degree of lateral compression,
and never shows signs of wear by grinding, but bears at

726

SCANDINAVIAN FISHES.

the top a small annular protuberance ; the middle tooth,
the largest one, is transversely (from behind and in
front) compressed and striated in the same direction,
with four bent furrows on its worn masticatory surface;
the hindmost tooth, as a rule the smallest one, has the
same transversely compressed form and similar mark-
ings, but is furnished with only three furrows. The
other two teeth are set beside each other at right an-
gles to the inner row, in an outward direction from the
middle tooth thereof, and have a more rounded masti-
catory surface, which at least on the inner (larger) one
is marked with stria?. The pharyngeal cartilage is de-
ciduous and in form resembles a more or less equilateral
triangle, with the anterior (upper) side almost straight
or slightly convex.

The dorsal tin begins in old Carp at a distance
from the tip of the snout that measures about 42 % of
the length of the body; but in young Carp its begin-
ning lies farther back. Its posterior part is of fairly
uniform height, but in front it is elevated in a some-
what pointed form, the upper margin being thus more
or less concave. The length of its base measures in old
Carp about 38 — 35 % of that of the body, and its great-
est height (the length of the first branched ray) about
13 — 12 % of the length of the body, the length of the
base of the fin being thus about three times its height;
but in young specimens — up to a length of about 2
dm. — the fin is comparatively much shorter, its height
sometimes measuring as much as half its length".

The anal fin begins in old Carp at about the end
of the second third of the length of the body, and its
base measures about 1/12 or V13 of the same length.
Its height (the length of the first branched ray), on the
other hand, measures about 1/g — ]/v °f the length of
the body and may thus even exceed the greatest height
of the dorsal fin. The form of the anal fin is an
oblique trapezoid, with the outer (lower) margin straight
or slightly convex.

The caudal fin is deeply forked, the length of the
middle rays (about 1j12 of the length of the body) be-
ing scarcely half that of the lobes; but the lobes vary
in form, being sometimes somewhat pointed, sometimes
rounded. Here, as in the generality of the Cyprinoids,
the caudal fin consists of 17 branched rays, one long,

simple ray at each margin (above and below), and at
the base of this ray a varying number of short sup-
porting rays.

The pectoral fins, which with their fleshy base are
set low and obliquely, just behind the lower third of
the gill-openings, are oval and obliquely rounded, the
third branched ray being the longest. The entire length
of the fins is about 16 or 17 % of the length of the
body* * 6, and the distance between the foremost points in
the insertions of these fins and in those of the ventral
fins (the length of the preabdominal region) measures
in old Carp about 21 or 22 % of the length of the body.

The ventral fins are of a more trapezoidal form,
with rounded hind margin. They are also of different
structure, the simple ray at the anterior margin being
furnished at its base with a short, simple, supporting
ray, which, though it is often difficult to distinguish,
appears as a rule in the ventral fins of all the Cypri-
noids. The length of these fins is generally greater in
the males than in the females and varies in old Carp
between about 16 and 14 % of that of the body. The
distance between the foremost points in the insertions
of these fins and in that of the anal tin measures about
27 % of the length of the body.

The covering of scales is subject in the Carp to
the most singular variations. The whole body (except
the head and fins) may be clothed with scales from the
very occiput, where the boundary of the scales lies at
a distance from the tip of the snout which is compa-
ratively greater in young Carp than in old, measures
about 83 — 79 % of the length of the head, and generally,
even in old specimens, is about equal to the depth of
the body at the occiput. Throughout their extent over
the body the scales are rather large (see above, Lin.
lat. and Lin. tr.), of a rounded quadrangular shape,
and so closely imbricated that each scale covers nearly
two-thirds of the adjoining scale behind it. The an-
terior (covered) part of the scale is marked with extre-
mely fine, numerous, concentric strife; but the radiating
grooves are only slightly traced and irregular, and the
anterior margin of the scale is only slightly and un-
evenly indented. On the hind (exposed) part of the
scale, however, both the concentric stria? and the ra-
diating grooves are coarser, and the surface is here

“ According to Canestrini’s measurements the length of the dorsal fin in a Carp 74 mm. long was only 27 % of that of the body

(including the whole of the caudal fin), and its height 13'5 % of the same length.

6 According to Canestrini’s measurements the length of the pectoral fins — which in general shows even relative increase with age —
in Carp 74 — 354 mm. long (including the whole of the caudal fin) is 14 or 15 % of that of the body.

CARP.

727

rough (granulated), furnished with a great number of
small, verrucose protuberances (thickened parts). The
lateral line descends from the temporal region in a
slight, but regular curve down to the middle of the
depth of the body at about the tip of the pectoral fins,
from which point it advances, straight and uninter-
rupted, to the middle of the base of the caudal fin.
The duct in each scale is short, and pierces only about
the middle third of the length of the scale. It has long
been known, however, that, in addition to the ordinary
Carp, there occur varieties (see our figure) which are
distinguished by the absence of scales and the greater
thickness of the skin on a greater or less part of the
body, the scales on the other parts being considerably,
even monstrously enlarged. One of these varieties is
represented in our figure. The largest scales lie in the
lateral line, in a continuous or an interrupted row. One
or two rows of rather large scales generally coast the
dorsal margin of this variety. The belly is clothed with
ordinary scales. As the large scales on the sides of
the body most often possess an iridescent or a vitreous
lustre, this variety has been named spegelkarp (Mirror
Carp). Another variety, which is entirely without scales,
but has a still thicker skin, is known as the Leather
Carp ( laderkarpen ).

There is hardly any external difference between
the sexes, except during the spawning-season, at which
time the males are marked by a dermal eruption, con-
sisting of small, white tubercles, on the cheeks and
gill-covers.

The usual coloration is olive green, on the back
darker, sometimes blackish brown, on the belly lighter,
sometimes yellowish white, on the sides with a coppery
or brassy lustre. It is, however, subject to consider-
able variations, shading on the back into blue or red,
on the sides still oftener into red, a colour especially
characteristic of the so-called Leather Carp. This
ground-colour is diversified by the black edges and
centre of each separate scale. The dorsal and caudal
fins shade most often into gray, the paired fins and the
anal fin into red. In the reddish varieties, however, all
the fins share in the general tone of colour. The iris
is yellow, with a bright golden margin round the pupil.
In captivity albinos sometimes occur.

The natural range of the Carp extends from Japan
and China through Central Asia to Eastern Europe. It

“ Den DansJce Atlas , p. 053.

b See, Yarrell, Brit. Fish., ed. 2, vol. I, p. 350 and Day, 1.

is difficult to fix the limit of the Carp’s natural range
in Central Europe; but it has been introduced in a
state of domestication throughout the rest of Europe,
except the extreme north, and has also been transported
to the East Indian Archipelago, Australia, and North
America. In India it is unknown. Subsequent to its in-
troduction it has spread to the lakes and rivers of these
regions, where it now lives even in a state of freedom.
Kroyer assumes, with great probability, that it was
during the Roman Catholic times, when fish was the
only animal food permitted during Lent, that the Carp
was introduced by the monks into Denmark and the
south of Sweden, and began to be cultivated in ponds
at the convents. We have no positive information,
however, on this point. An old tradition, on the other
hand, quoted from Pontoppidan" by Bloch, Kroyer,
and Nilsson, but also in need of confirmation, ascribes
the introduction to a later date, when Peder Oxe
(L567 — 1575) was Lord Steward of Denmark and owned
large estates in Scania. In England Carp existed, though
their number was small, in 1496, according to Dame
Juliana Berner’s Boke of St. Albans6; and at the pre-
sent day the Carp is very common in that country.

The Carp, like the other Cyprinoids, is by nature a
fresh-water fish, but it can also endure salt water. In
the Caspian Sea, according to Pallas, the Carp lives
in water so salt that hardly any other fish can sustain
life there. In brackish water it occurs, according to
Mobius and Heincke, in the Schlei at Schleswig. In
Sweden it has spread from the ponds to the lakes and
rivers, principally in Scania. It has also been culti-
vated, without doubt, on the estates of Central Sweden,
for it occurs, though extremely rare, in Lake Malar, where
in June, 1888 a fisherman from Sundbyholm (Soder-
manland) caught the female Mirror Carp represented
in our figure (7 3 of the natural size). Neither he nor his
acquaintances had ever seen one of these fishes before.

In a state of nature the Carp haunts places with
a luxuriant vegetation, being by no means averse to a
muddy bottom, but requiring clear water and free access
to the sunlight. Distinction has long been made be-
tween River Carp, Lake Carp , and Pond Carp, which
generally differ not only in their coloration, which
adapts itself to the bottom and the water, but also in
their flavour, which is best in the first-mentioned forms,
worst in the last-mentioned, unless the pond has been

c., p. 103.

Scandinavian Fishes.

92

SCANDINAVIAN FISHES.

supplied with a continual How of fresh water. In
summer, so long as the water is not too warm, Carp
frequent shallow and sunny spots; in winter they flock
together in deep pools.

The adult Carp is sluggish and peaceable in dispo-
sition, though frequently migratory in its habits, espe-
cially during the spawning-season. It is highly sensitive
to heat and cold: during the colder part of the year
it abstains from food. Still it is very tenacious of life,
and may be transported long distances alive. It may
also be transferred from its native haunts to some ar-
tificial receptacle, where it may be kept alive for a
long time, and better seen to and fattened. This is an
old custom, resorted to especially for the purpose of
freeing Pond Carp from their muddy flavour. Packed
in damp moss or ice, and with a bit of bread dipped in
spirits in the mouth, the Carp can live at least twenty-
four hours or more. The above-mentioned Mirror Carp
from Sundbyholm was sent to the Royal Museum alive
in a bucket, and was kept alive in a tub for some days.
One morning it was found to have leapt out of the
tub, and lay on its side apparently dead. It was re-
stored to the water, but floated belly upwards and did
not move a limb. It was then given a dessert-spoonful
of spirits, and began after some minutes faintly to move
its pectoral fins. After a quarter of an hour the dose
was repeated, and within an hour the fish moved about
with ease, as if nothing had happened. During the warm
season, however, the Carp soon dies of suffocation, if
prevented from coming up to the surface to breathe.
Buckland tells® an amusing anecdote of its habit of
breathing air. In an Irish lake fairies were seen danc-
ing on the water during calm, moonlight nights — the
fairies being large Carp that made rings at the surface
by smacking their lips as they breathed.

The food of the Carp is both animal and vegetable,
consisting of seeds, fish-roe, and insects; but it never,
or extremely seldom, bites at a fly. Its favourite
nourishment is composed of decomposing vegetable
substances, worms, Entomostraca, and the larvse of in-
sects. The mud swallowed by it can hardly be included

among its food, but seems necessary to its digestion
and general health.

The spawning-season occurs in May or June, but
may sometimes be protracted till the month of August,
being often interrupted during unfavourable weather.
At this time the Carp is exceedingly restless and bold,
darting wildly about, jumping out of the water, and
leaping high, if necessary, to reach a suitable spawning
place. Here the eggs, which are small (about 1 x/2 mm.
in diameter) and light yellow or yellowish green, are
deposited in shallow water, and attach themselves in
lumps to the water-plants, especially Utricularia, Nym-
phcea , Alisma, and Glyceria b. The males — two or three
to every female — swim at the surface above the fe-
males, as far as the depth of the water permits. The
number of the eggs varies according to the age and
size of the females, but is also different in individuals
of the same size. In a female 3 lbs. (lV5 kgm.) in
weight Bloch" counted 237,000 eggs, and in a female
16V2 lbs. (7x/2 kgm.) in weight, the ovaries of which
weighed 5x/2 lbs. (2x/2 kgm.), Buckland d estimated the
number of the eggs at 2,059,750; but another female,
21 V2 lbs. (9 3 4 */4 kgm.) in weight, contained only 1,310,750
eggs. Sterile Carp are not uncommon, and have always
been highly esteemed for their fat, delicate flesh. An
English fisherman of the name of Tull discovered6 7 a
method of castrating Carp in order to fatten them with
better success.

In a suitable temperature the ova develop so
rapidly that the eyes of the embryo appear on the fifth
or sixth day, and the egg is hatched between the twelfth
and sixteenth days, according to Day, or as early as
the eighth day, according to Benecke. Of the growth
of the fish Canestrini/ tells us that at the age of one
year it weighs 8 gm., at the age of two 32 gm., at
the age of three 500 gm., at the age of four 1 kgm.,
at the age of five 2 — 4 kgm., at the age of six 4 — 5
kgm., at the age of seven 7 — 8 kgm., and at the
age of eight 9—10 kgm. These weights vary of
course according to the abundance of food. The Carp
may reach maturity in the third year.

a Nat. Hist. Brit. Fish., p. 40.

4 The seeds of Glyceria fluitans are also an agreeable food for the Carp.

c L. o., p. 97.
d L. c., p. 38.

‘ See Watson, An Account of Mr. Samuel Tull’s Method of Castrating Fisk., Philos. Trans., vol. XLVIII, art. CVI, p.
(Dec. 1754).

7 Fauna D’ Italia, part. Ill, Pesci , p. 10.

870

CAKP.

729

In Sweden the Carp-fishery is of little importance,
the occurrence of this fish in a natural state and in
any quantity being confined to a few lakes in Scania
— Vombsjon and Ringsjon, according to Nilsson —
with the streams that feed and drain them. The Carp
is taken with the tackle ordinarily employed in fresh-
water fishing, but principally in different kinds of traps.
It is generally too cunning for the net and seine, avoid-
ing the former and swimming under or leaping over
the latter — a ruse which even the ancient Greeks and
Romans knew it to practise. Sometimes it is caught
with rod and line or on nightlines, a. method of fishing
extensively employed in France both as an amusement
and a source of profit. The bait consists of lobworms,
a piece of old Gruyere, or a bit of dough kneaded with
a little honey. The hook must be allowed to sink to
the bottom, and the bait should not be too large, for
the Carp must gulp it down at once — if it begins to
nibble at the bait and feels the hook, it cautiously spits
out both of them. The angler may choose a spot quite
near land, for the Carp resorts freely to the shore for
food, and may be enticed thither without difficulty by
now and then throwing some breadcrumbs into the
water. The best time for angling is the evening.

Large, or at least middle-sized Carp, 3 — 5 kgm.
in weight, are good eating when well cooked, especially
when boiled in wine or stewed with spices. It was
therefore quite natural that, even in olden times, at-
tempts should be made to utilise this fish in the most
convenient and most profitable manner. Hence the
origin of the cultivation of the Carp, an industry which
has made great strides in some countries, especially in
Germany and Austria, and which even in Sweden has
been carried on with success. In his Skanska JResa
LiNNiEUS described the condition of this industry in
Scania in 1749, and his opinion was that “the ground
occupied by a fishpond is far more valuable to its pos-
sessor than the finest tilth” (p. 378). In an academical
thesis (Lund, 1766) Cederlof has left us some econo-
mic remarks on the Scanian Carp-poncls. But this in-
terest flagged, and it was not until the present day
that the cultivation of the Carp was resumed as an
industry. This has been done by the instrumentality
of Mr. Wendt, a German landowner who removed

to Sweden, and in 1879, at Gustafsborg near Perstorp,
a railway station in the District of Christianstad, con-
structed a Carp-breeding establishment. In 1883 he sold
30,000 Swedish pounds (12,752 kgm.) of fish, and in
1884 half as much again (19,128 kgm.), most of this
quantity being exported to Germany". At Langbans-
hytta in Wermland another establishment for the breed-
ing of Carp has lately been opened \

These Carp-ponds are dug, or still better — if their
site admits of this — formed by damming, beside a
river or brook. They should be some feet deep and
at least two in number, so constructed that one or
more of them may be emptied at the same time and
thoroughly drained. In a complete establishment the
ponds must be of two kinds, breeding-ponds (Germ.
Streichteicke, where the Carp may spawn, sick streichen )
and growing-ponds (Germ. Streckteicke , where the Carp
may grow, sicli strecken , from the beginning of the se-
cond year). In some places a third kind of pond may
be seen, the stew (Germ. Fetteich, Setzteich, Abwaclis-
teick, Kammerteick), where the Carp that have reached
the age of three years or more, are kept ready for use
or sale. The size of the ponds depends on circum-
stances; but the breeding-ponds, which may be most
reduced in size, should measure at least 10 — 12
ares (about 1 rood), and to each are 5 females and 3
males may be allowed. These ponds, in which the fish
are to spawn, must also contain a shallow part, where
the water may be heated by the sun, as early in the
year as possible, to a temperature of about +17° or
+ 18° C. (about 63° Fahr.), for at a lower temperature
the Carp will not readily spawn. The growing-ponds
as well as the stews, on the other hand, must contain
a deeper part, at least about 5 ft. in depth, where the
Carp may assemble in winter without danger of the
bottom freezing. The entrance and outlet must of
course be constructed, in the first place, so as to avoid
the possibility of inundation; and precautions should
be taken by means of suitable filters to prevent the
Carp from escaping and to keep out other fishes and
noxious animals. One or two Pike, to cut off the
weaklings and to keep the older Carp in active motion,
do no harm in the ponds, for the survivors thrive and
grow all the better for their company; and Perch and

° See Trybom, Om karpodling i alhnunhet och sarskildt om den vid Gustafsborg i Kristianstads Ian , Landtbruks-Akademiens Hand-
lingar och Tidskrift for ar 1885.

b See Lilleborg, So., Norg. Fisk., vol. Ill, p. 794. When this sheet was in the press, the second number of the ‘tivensk Fiskeri-
TidskrifC appeared, containing a paper by Ii. V. Tiberg on the cultivation of the Carp at Langbanshytta.

730

SCANDINAVIAN FISHES.

Pike-perch are also of service, though not in exactly
the same degree. But large numbers of other fishes,
especially of the Carp’s own family, such as Roach,
Rudd, or Crucian Carp, only do mischief. The most
practical method of keeping the ponds clean, a method
which at the same time renders their bottom better ca-
pable of developing food for the Carp, is to drain them
in turn and leave them dry for a year, during which
period they may be ploughed and sown, in most cases
with good results. They must also be emptied to faci-
litate the transfer of the hsh from one pond to the
other, an operation which should be performed during
the cool season, in autumn or spring, the heat of sum-
mer being at best disagreeable to the Carp on dry land.
It is obvious that the ponds should also be protected
from the inroads of poachers, otters, herons, and other
enemies of the Carp; but ducks do no damage. The
reader who desires fuller information on this head, will
find what he needs either in Norback’s Handledning i
Fiskevard och Fiskafvel (p. 257) or in the numerous
German manuals on the subject, e. g. Max v. d. Borne,
Handbuch der Fischzucht und Fischerei , Berlin 1886,

p. 261.

Among Carp as among other domesticated fishes
we frequently meet with deformed individuals. One of
these examples, not uncommon in its occurrence, with
the snout apparently reduced, is described by Valen-
ciennes'1. The reduction, however, does not really affect
the snout or the jaws, which parts are developed as
usual, or only distorted, the frontal bones, on the other
hand, being anteriorly depressed, and the basisphenoid
and parasphenoid bones reduced.

In the pond the Carp is a victim to all kinds of
diseases. We sometimes hear of Carp being so old that
moss has begun to grow on their bodies, this moss

As we have mentioned above, many cultivators
of the Carp are unfortunate enough to have their
breed of fish spoilt by crossing with the Crucian Carp.

being really a mould fungus, Saprolegnia, of the same
genus as the scourge of the Salmonoids. The best re-
medy for this is a little salt in the pond or a change
of water. By unsuitable food the whole stock of a pond
may be poisoned. Carp may be fed and fattened on
ordinary kitchen-refuse, brewer’s grains, boiled and
mashed potatoes, chopped cabbage, guano, or the dung
of the horse, pig, or sheep. But experience, according
to M. v. d. Borne and others, shows the necessity of
great caution, and the economical results of fattening
have not been satisfactory. Carp are often troubled by
the male frogs, which under the influence of sexual
excitement attach themselves firmly to the head of the
sluggish Carp, and with their forefeet press the eyes
of the fish so hard as to produce blindness.

For all this the Carp is an easily fed and easily
managed domestic animal; but without expensive pre-
paration for table its culinary value is even less than
that of the Pike. In the south of Europe its roe is
made into caviare, consumed principally by the Jews,
who are forbidden by law to eat genuine caviare, the
Sturgeon being destitute of ordinary scales.

The age attained by the Carp is not yet ascer-
tained. Buffon mentioned Carp 150 years old as exist-
ing in the moats at Pontchartrain, and Lebel stated
that the ponds at Lausitz* 6 contained Carp 200 years
of age. It is difficult, however, to test the truth of
these old statements, and the same difficulty applies to
Bloch’s account0 of the capture in 1711, at Bischof-
hausen near Frankfurt-on-the-Oder, of a Carp 6 feet
long, 2V5 feet deep, with scales as large as an 8-
groschen piece, and 70 lbs. in weight. In a state
of freedom, according to Heckel and Knee (1. c., p.
57), the Carp only attains the age of about 12 or 15
years.

The hybrid produced by this cross-breeding has gene-
rally been known among modern ichthyologists by the
name of

a Cuv., Val., Hist. Nat. Poiss., vol. XVI, p. 57.

6 See Lacepede, Hist. Nat. Poiss., tome V, p. 510 and Bloch, Naturg. Deutschl. Fish., part. I, p.
c Cited from Beckmanns Gesch. der Chur., Th. I, p. 574.

95.

HYBRID CARP.

731

CARPI 0 ROLL ARII.

Fig. 179.

Two pairs of barbels, but these more or less reduced, the anterior pair or even one of the posterior barbels {at
the corners of the mouth ) being often rudimentary. Most of the pharyngeal teeth transversely compressed {in front
and behind), with the masticatory surface furnished with one groove , usually: 1, 4 — 4, 1 {sometimes: 3 — 3, some-
times; 1, 1, 4 — 4, 1, 1). Greatest depth of the body about 1/3 of its length; distance from the foremost point
in the insertions of the ventral fins to the beginning of the anal fin less than 4/5 of the said depth.

Fig. 179. Ccirpio Kollarii, natural size, from Rydgard (Scania), taken on the 28th of October, 1858, and its pharyngeal bone
(left) and pharyngeal cartilage magn. 3 diarn. a, b, and c as in the preceding figure.

R. br. 3; D. --- -- ■ A. -f — ; P. - — 1 ; V.

1 9 “ — 21 6 b — 7 15—16“ 8—9

C. x 1 + 17 + 1+a?; Lin. lat. 36- — 37; L. tr. 1; Vert. 36.

(6)7

Syn. Cyprinus Kollarii , Hckl., Ann. Wien Mus., vol. I, p. 223,
tab. 19, fig. 2; Id. ( Carpio ), Russeggers Reisen, part. II,
p. 1014; Cuv., Val. ( Cyprinus ), Hist. Nat. Poiss.,
vol. XVI, p. 76, tab. 458; Hckl, Kn. ( Carpio ), Suss-

“ Sometimes 17, according to Siebold.

•n

732

SCANDINAVIAN FISHES.

wasserf. Oestr. Mon., p. 64; Sieb., Silsswasserf. Mitteleur.,

p. 91.

Cyprinus striatus, Holandre, Fn. d. 1. Moselle, p. 242.

Carpio Sieboldii , Jackel, Fish. Bay., Abh. zool. mineral. Ver.

Regensburg, vol. 9 (1864), II, p. 21.

Carasso- Cyprinus vulgo-Carpio , Fatio, Fn. Vert. Suisse, vol.

IV, part. I, p. 198.

Carpio Kollarii (Sw. Karprudan ) in its Swedish
forms is even externally an intermediate type between
the Carp and the Crucian Carp. The depth of the body
is greater, and the base of the dorsal fin is shorter
than in the Carp, both measuring on an average 1/3
(31 — 34 %) of the length of the body. The changes
of growth and the individual variations both in the
Carp and the Crucian Carp, however, render these
characters of little constancy, though they are generally
sufficient to enable us to recognise a Carpio Kollarii.
The same may be said of the texture of the scales,
which in this hybrid usually show more distinct and
more regular radiating grooves on the anterior (covered)
part of the scale and notches in the anterior margin
thereof. The most important expression of the hybrid
nature of Carpio Kollarii is afforded by the form of
the pharyngeal teeth and the inconstancy of their num-
ber, which is often different even on the right and left
pharyngeals.

The first Swede to describe Carpio Kollarii as an
actual hybrid between the Carp and the Crucian Carp,
was Schagerstrom (in the “Physiografiska Sallskapets
Tidskrift” 1838, p. 295), and Lilljeborg has subsequently

described (in Sveriges oclt Norges Fiskar, vol. Ill, p.
137) at greater length one of the specimens on which
Schagerstrom based his opinion, and which were taken
in a Carp-pond /on the manor of Krapperup in Scania.
This specimen was 3 dm. long. Baron G. C. Ceder-
strom found this hybrid to be plentiful among the
Carp and Crucian Carp in a pond at Rydgard. The
largest of the specimens he presented to the Royal
Museum from this locality, has served as the original
of the figure given above in the natural size.

That Carpio Kollarii is really a hybrid, has been
proved by the experiments of Hessel. “In order to
determine this question,” he writes0, “I myself managed
to bring about such crosses by placing (1) female com-
mon carp with male crucian carp, and (2) female cru-
cian carp with male common carp, in small tanks,
constructed with this end in view; (3) I also put to-
gether female Carpio kollarii with male common carp;
this for the sole purpose of testing the capability of
propagation of the C. kollarii, which had been doubted.
In the two former cases I obtained forms analogous to
Carpio kollarii, sometimes approaching in appearance
the true carp, at others the crucian carp. In the third
case, however, having placed ripe Carpio kollarii to-
gether with Cyprinus carpio, I obtained a product with
difficulty to be distinguished from the genuine Carp.
I took the trouble to feed them for three years, in
order to try their fitness for the table, but their flesh
was exceedingly poor and very bony and could not be
compared by any means to that of the common carp.”

Rep. U. S. Comm. Fish and Fisheries 1875 — 76, p. 868.

CARPS.

733

THE GOLDFISH (s\v. guldfisken).

CYPPJNUS AURATUS.

Plate XXXI, fig. 2.

No barbels. Preabdominal length ( distance between the foremost { uppermost ) points in the insertions of the pec-
toral and ventral fins) less than 3/5 of the length of the base of the dorsal fin. Length of the lower jaw more
than 3/s °f Hie preabdominal length. 5 or 6 scales in an oblique transverse row between the later al line and the
anterior part of the base of the dorsal fin. Pharyngeal teeth set in one row , transversely (in front and behind )

compressed , and with one groove in the masticatory surface: 4 — 4.

Fig. 180.

Pharyngeal bone (left) and pharyngeal cartilage of Cyprinus auratus , magn. 3 diam.

a, b, and c as in the preceding figure.

R. hr. 3; D. — .4. 2 — ; P. - V. — :

18®— 20 5—7 15 6 8 c

C. x + 1 + 17 4- 1 + Lin. lat. 29d — 30; Lin. tr. - 1

7

Vert. 30—31.

Syn. Cyprinus pinna ani duplici, caudce trifurca, Lin. Vet. Akad.

Handl. 1740, p. 403, tab. I, figg. 1 — 8; Fn. Suec ., ed. I,
p. 125, tab. II, fig. ad. num. 331.

Cyprinus auratus , Lin., Syst. Nat., ed. X, tom. I, p. 322;
Raster, Opusc. subseciva, tom. II, lib. II, p. 78, tab. IX;
Bl., Fisch. Deut.schl., part. Ill, p. 132, tab. XCIII et XCIV
(+ Cypr. macrophthalmus, Ausl. Fiscli., part. IX, p. 51,
tab. CCCCX); Lacep. {Cypr. auratus + C. argenteus 4- C.
telescopus + C. macrophthalmus + C. quadrilobatus ) Ilist.
Nat. Poiss., tom. V, p. 553; Yarr, Brit. Fish. ed. I, vol.
I, p. 315; Cuv., Val. {Cypr. lineatus — obs. tamen D: 4/o3 >
Lin. lat. 33; Lin. tr. 11 — + C. thoracatus + C. Langs-
dorfii + C. auratus ) Hist. Nat. Poiss., vol. XVI, p. 96,
cett. ; Richards. {Cypr. lineatus + C. carassioides (ex Gray)
+ C. Burgeri + C. gibelioides (ex Cant.) + C. Cuvieri +
C. Langsdorfii + C. thoracatus + C. abbreviatus + C. au-
ratus) Rep. Brit. Assoc. 1845, p. 291, cett.; Basil. {Caras-
sius Pekinensis + Car. coeruleus + Car. discolor + Cyprinus
auratus + Cypr. macrophthalmus + Cypr. quadrilobatus)
Nouv. Mem. Soc. Natur. Mosc., tom. X (1855), p. 229, cett.,
tab. Ill, fig. 3, tab. V, figg. 1 — 5, tab. IX, fig. 2; Blanch.
( Cyprinopsis , ex Fitz.) Poiss. cl. eaux clouces Fr., p. 343;
Gthr {Carassivs) Brit. Mus. Cat., Fish., vol. VII, p 32;
Fedders., Naturh. Tidskr. Kbhvn, ser. 3, vol. XII, p. 83;

Fatio, Fn. Vert. Suisse, vol. IV, part. I, p. 205; Day, Fish.
Gt. Brit., Irel. , vol. II, p. 166,. tab. CXXX, fig. 2; Lillj.
{Cyprinus) Sv., Norg. Fn., Fisk., vol. Ill, p. 158.

Carassius Langsdorffii + Car. Burgeri + Car. Cuvieri + Car.
grandoculis, Schleg. (Sieb.) Fn. Japan. Poiss. p. 192, cett.
tab. XCVI1I.

The Goldfish is everywhere so well knotvn that any
special description of it seems unnecessary. In its nor-
mal type it is an intermediate form between the Carp
and Crucian Carp, with a more shallow and more terete
body and longer dorsal fin than the latter. During
youth, hoAvever, the Crucian Carp and the Goldfish may
be almost exactly similar in these respects, especially as
the Goldfish undergoes exceedingly great variations in a
domesticated state. The rule is that the latter generally
has comparatively larger fins and shorter preabdominal
region than either the Carp or the Crucian Carp, the
young Goldfish being thus most easily recognised —
where the coloration does not decide the question — by
the fact that the tips of the pectoral fins extend behind
the perpendicular from the insertions of the ventral fins.

The Goldfish, the hin-ju of the Chinese, has its
original home “in a small lake, Tschekiangb near the

a Sometimes

b

11

e In modern

15, according to Fatio and Canestrini.

14, „ ,, Fatio.

9, ,, ,, Fatio.

25, ,, ,, Fatio and Canestrini.

maps Tschekiang is the name given to

one of the eastern provinces of China, on the coast, between 27° — 31°

N. lat.

734

SCANDINAVIAN FISHES.

town of Tschangwahyen in the province of Hiantschewfu,
at the foot of the mountain TsienkingV’ From this
locality it was first transported to the northern pro-
vinces of China and to Japan, long before it became a
luxury in Europe at the middle of the last century.
The assumption has indeed been made that it had been
introduced at an earlier date into Europe by the Portu-
guese, after they had discovered the route to India
round the Cape of Good Hope; and at the present day
it is said to be a common fish in the streams of Por-
tugal. But it was not until the eighteenth century
that the cultivation of the Goldfish was begun in Eu-
rope. The French East Indian Company then presented
some Goldfish to Madame Pompadour, and about 1730
the species was naturalised in England. From that coun-
try it subsequently spread over the whole of Europe;
and at present the breeding of Goldfish is practised on

Fig. 181. The Telescope-fish, a variety of the Goldfish.

After Gunther.

the largest scale in Germany. The neighbourhood of
Havre in France also supplies the market with con-
siderable quantities of Goldfish.

In its original state the Goldfish is olive green,
darker — sometimes blackish brown — on the back and
lighter — sometimes whitish — on the belly, with a
golden (brassy) or silvery lustre, and has the form
shown, with only slight deviations from the natural
type, in our figure (Plate XXXI, fig. 2). Cultivation
and selection6 are the causes of the well-known altera-
tions both in colour and form c. The colour shades
1) into blood-red or orange, ending in the lustrous
gold of the typical Goldfish, 2) into paler yellow or

white, a transformation which gives the Silverfish its
name, or 3) into brown, blue, or nearly pure black.
These tints either become predominant both on the
body and the fins, or in fhe light varieties, leave
patches and traces of the original colour. The changes
of form tend not only to a more terete shape of body,
but also to the most grotesque modifications of the fins.
The dorsal fin is reduced more and more, until it
finally disappears, or it may also be divided into an
anterior and a posterior part. The caudal and anal
fins are doubled ; and when these changes are accom-
panied by the protrusion of the eyes, they culminate
in the variety described by Linnjeus in 1740, the
Telescope-fish (fig. 181). The internal organs also
undergo monstrous alterations. The development of
the air-bladder may suffer such distortion as to leave
the fish a helpless cripple.

In this manner the Goldfish has been cultivated
and deformed for the amusement of people of rank
(‘in Magnatum oblectamentum’, Linnaeus), above all at
the imperial court of China, where officials have been
especially appointed to take charge of the Goldfish, and
where, as Lacepede remarks, the women may well re-
quire this diversion to break the monotony of their
idle existence. Among these ladies it has been a shift-
ing fashion to keep one variety or another, and an
interesting observation of nature to watch the amatory
passages of the Goldfish, as the male caresses the gravid
female, rubbing his body against hers. The peaceful
and sociable life of these fish and the ease with which
they may be bred, without the need of any trouble-
some attendance, have rendered them agreeable and
cheap pets. By ringing a bell — the Chinese always
have one hanging beside the ponds — each time the
fish are fed, they may be trained even in large pools
to obey this summons and to come up to the shore,
when the bell is sounded, to show themselves. By
means of warmth and abundant food Goldfish may
be induced, in suitable aquaria or in fish-ponds, to
breed three or four times in the course of the summer.
The Goldfish hatched in spring are 3 — 7 cm. long
by autumn, the largest being now ready for sale, as
they have generally acquired by this time the proper

a According to the Jesuit missionary Le Comte, who wrote an account of his travels at the end of the seventeenth century. See
Baster, 1. c., p. 80. At the same period Iyampfer, one of Olaus Rudbeck’s followers, published a book of travels and a history of Japan,
where he mentions the Goldfish, the lcing-jo of the Japanese.

h Land and Water, May 3rd 1879 and C. Wagner, Wasser-Cidtur, Bremen 1881.
c Cf. Baster, 1. c.. tab. IX.

CRUCIAN CARP.

735

coloration. In its earliest youth, the duration of which
may, however, be shortened by cultivation, the Goldfish
wears its original colour. Traces thereof appear in our
figure on the lips and the pectoral and ventral fins.

In many parts of Southern and Central Europe
the Goldfish has become completely naturalised. In
Sweden there can hardly be any locality where the
natural temperature is high enough fully, to satisfy the
climatic requirements of the Goldfish; but in Denmark
it is kept in ponds at several places, and even in Sw*e-
den it can live and multiply in the open, as shown
by an experiment made at Helsingborg. In 1888 five
handsome Goldfish were put into a fountain in the Park

of that town, and left to find their own support among
the algm growing on the bottom of the basin, and
among the insects they might capture at the surface
of the water. Here they survived a severe winter, and
seemed to thrive excellently, having gained both in
weight and size by the following summer. They also
considerable increased in number, the parent stock being
now surrounded by a numerous progeny".

The flesh of the Goldfish was tasted and com-
mended long ago by Raster (1765). It is said to be
excellent, whether boiled or fried, and superior to that
of the Carp. The Goldfish of commerce yields most pro-
fit, however, by its sale as stock for ornamental waters.

THE CRUCIAN CARP (sw. rudan).

CYPRINUS CARASSIUS.

Plate XXX, figs. 2 and 3.

Mouth without barbels. Preabdominal length more than 3/5 of the base of the dorsal fin. Length of the lower
jaw less than 3/5 of the preabdominal length. 7 or 8 scales in an oblique transverse row between the lateral line
and the anterior part of the base of the dorsal fin. Pharyngeal teeth set in one row , for the most part trans-
versely compressed , with simple or one-grooved masticatory surface: 4 — 4.

Fig. 182.

Pharyngeal bones and pharyngeal cartilage of Cyprinus carassius. Nat. size.

a, h , and c as in the preceding figure.

3 4 3 i

R. hr. 3 ; D. - — ; A. — — — ; P. — —

16—19 6—7 13—16

; V.

7 — 9 ’

C. x + 1 + 17 + 1 + x: Lin. lat. (31)33 — 35; Lin. tr.

1 ; Vert. 32.

Syn. Karas (Karass) et Giblichen, Gesn. De Aquat., Paralip., p. 16;

Cyprinus brevis , Schonev., Ichthyol. Slesv. Hols., p. 33:
Cyprinus pinna dorsi ossiculorum viginti, linea laterali recta,
Art., Ichthyol., Gen. Pise., p. 4; Syn. Pise., p. 5; Descr.
Spec. Pise., p. 29; Lin. Fn. Suec., ed. I, p. 122 (pinna am
err.); Cyprinus Hamburger dictus, Gronov., Act. Upsal. 1741,
p. 75; Cyprinus, No. 4, Klein, Hist. Pise. Natur. Miss. V,
p. 59, tab. XI, figg. 1 et 2.

Cyprinus Carassius, Lin., Syst. Nat., ed. X, tom. I, p. 321;
Penn., Brit. Zool. (ed. 1776), vol. Ill, p. 319; Bl ., Fisch.
Deutschl., part. I, p. 69, tab. XI; Retz., Fn. Suec. Lin., p.
355; Swartz, Sv. Zool., vol. I, tab. No. 10; Ekstr., Vet.-
Akad. Handl. 1830, p. 192; Pall., Zoogr. Ross. Asiat., tom.

a See the Svenska Dagblad for the 22nd of August, 1889.

Ill, p. 297; Nilss ., Prodr. Ichth. Scand., p. 32; Ekstr.,
Vet.-Akad. Handl. 1838, p. 213; Id., Skand. Fisk., ed. 1,
p. 140, v. We., tab. 31 et 32; Yarr., Brit. Fish., ed. 2, vol.
I, p. 355; Cuv., Val., Hist. ATat. Poiss ., vol. XVI, p. 82,
tab. 459; Nilss., Skand. Fn., Fisk., p. 290; Blanch. ( Cy -
prinopsis ) Poiss. d. eaucc donees Fr., p. 336; Lindstr. Gotl.
L. Hush. Sallsk. Arsber. 1866, p. 16 (sep.); Mela Vert.
Fenn., p. 317; tab. X; Lillj., ( Cyprinus ) Sv., Norg. Fn.,
Fisk., vol. Ill, p. 147.

Cyprinus Gibelio, Bl., 1. c., p. 71, tab. XII; Ekstr. 1830,
1. c., p. 196; Nilss., Prodr., p. 33: Yarr., 1. c., p. 358;
Cuv., Val., 1. c., p. 90; Hckl., Kn. (Carassius') Siisswasserf.
Oestr. Mon., p. 70; Coll., Forh. Vid. Selsk. Chrnia 1874;
Tilliegsh., p. 180; Blanch. ( Cyprinopsis ) 1. c., p. 340; Norh.
( Carassius ) Handl. Fiskev., Fiskafv., p. 417, tab. XLVIII,
fig. 130.

Carassius vulgaris , Nordm., Voy. Russ. Mer. (Demidoff), tom.
Ill, p. 479; Kr., Damn. Fisk., vol. Ill, p. 294; Hckl. Kn.,

Scandinavian Fishes.

93

736

SCANDINAVIAN FISHES.

1. c., p. 67; Dybowski, Vers. Monogr. Oypr. Livl., p. 41;
Sieb. Siisswasserf. Mitteleur ., p. 98; Mgkn, Finl. Fiskfn.
(disp. Helsingf. 1863), p. 37; Canestr., Arch. Zoo!., Anat.,
Fisio]., vol. IV, fasc. 1 (1866), p. 66; Gthr, Cat. Brit.
Mus., Fish., vol. VII, p. 29; Feeders., Nat. Tidskr. Kbhvn,
ser. 3, vol. XII, p. 82; Bncke, Fisch., Fischer., Fischz. 0.,
W. Preuss., p. 109; Mor., Hist. Nat. Poiss. Fr., tom. Ill,
p. 374; Mob., Hcke, Fisch. Osts., p. 106; Day, Fish. Gt.
Brit., Irel., vol. II, p. 164; Bncke, Hanclb. Fischz., Fischer.
(M. v. d. Borne), p. 119.

Oyprinus moles, Agass., Mem, Soc. Sc. Nat. Neuchat., tom. I
(1836), p. 37.

Oyprinus humilis + ( ?) C. buceplialus, IIckl, Ann. Wien., Mus.
vol. II, pp. 156, 157, tali. 9, fig. 4.

Oyprinus Linncei, Bp., Cat. Met. Pesc. Enr., p. 27; Malm
( Carassius ) Gbgs, Boh. Fn., p. 556.

Oyprinus oblongus, Hckl, Kn., Susswasserf. Oestr. Mon., p. 73.

A: The Lake Crucian Carp ( Cyprinus carassius, auctt.)
— Plate XXX, fig-. 2.

The size of the Crucian Carp is very variable, being
influenced by the water in which the fish lives, and by
the supply of food. Most specimens are small, but
sometimes the species attains a length of 4 dm. or
more. The largest specimen Ekstrom saw measured
35 cm. to the base of the caudal fin, i. e. 42 cm. to
the end of the caudal lobes. Lill.teborg mentions se-
veral specimens 45 cm. long from Upland, and he once
saw a still larger Crucian Carp, which was taken in
Lake Hedervik (Upland) and weighed nearly 3V2 kgm.

The body is rather thick, but compressed and very
deep, the greatest depth", which coincides with the
perpendicular from the beginning of the dorsal fin to
the bases of the ventral fins, measuring half the length
of the body to the base of the caudal fin or even to
a- point half-way along the said fin. The greatest thick-
ness, which lies almost in the same perpendicular, mea-
sures more than 1/3 (about V5) of the greatest depth.
'Fhe back is broad and convex at the occiput, gradually
decreasing in breadth towards the dorsal fin, at the
base of which it is strongly compressed and thin. From
the end of the dorsal fin to the caudal fin the back
resumes its broad, convex shape. Forming a more or
less obtuse angle at the beginning of the dorsal fin,
the entire dorsal profile runs in a high arch to the end
of the said fin, where a depression begins and extends
to the caudal fin. The belly is flat in front of the
ventral fins, between these fins and the anal fin con-
vex, with a low carina along the median line. The

curve of the ventral profile, less marked than that of
the dorsal, is regular to the beginning of the anal fin,
from which point it rises more abruptly to the peduncle
of the tail. This finless part of the tail is deep6 and
very short, the distance between the anal and caudal
fins measuring half the depth of the tail at the ter-
mination of the former fin.

The head, which is not very large, occupies in
adult specimens about 22 % of the length of the body,
or rather more than 1/i of the length to the base of
the caudal fin, while in young specimens its relative
length is somewhat greater. The breadth measured
across the gill-covers is about equal in adult specimens
to the distance from the tip of the snout to the hind
margin of the preoperculum, in young specimens to
that of the orbit. The forehead is broad and convex,
its breadth at the middle of the eyes measuring in
adult specimens about 38 — 41 % of the length of the
head. The profile is straight from the snout to the
occiput, where a slight break generally interrupts its
continuity. The snout is blunt. Its length in adult
specimens is about 1/3 of that of the head. It projects
slightly in front of the mouth, which is small, pro-
trusile, and turned slightly upwards. The lips of young
specimens are fairly thick, especially the underlip; but
in old specimens they shrink, and sometimes leave the
sharp inferior edge of the intermaxillary bones to form
the very margin of the upper jaw. The upper jaw
projects somewhat, though only slightly, beyond the
lower; but the length of the latter is generally some-
what greater than that of either the snout or the oper-
culum. The eyes are circular and middle-sized, being
comparatively larger as usual during youth, and vary-
ing in longitudinal diameter between about 70 and 40
% of the breadth of the interorbital space. They are
set so high that an horizontal line drawn from the
middle of the margin of the upper jaw touches the
lower edge of the iris or of the pupil. The nostrils
resemble those of the Carp; but the two anterior nostrils
lie at a somewhat greater distance from each other than
from the tip of the snout. The operculum is flat,
’without any marked convexity, and rendered rough to
the touch by rows of tubercles radiating towards the
inferior margin. Its upper margin is united by the
skin to the side of the body. In this species also the

a About 41 — 43 % of the length of the body to the end of the middle caudal margin.
0 Least depth of the tail about 13 / of the length of the body.

CRUCIAN CARP.

737

branchiostegal membranes coalesce with the sides of the
isthmus, at a distance from each other which is equal
in old specimens to the longitudinal diameter of the
eye. The size of the gill-openings is consequently not
very considerable, . though greater than in the Carp,
the head being much deeper. The branchiostegal mem-
brane forms a broad, membranous rim, usually broader
than in the Carp, along the margins of the operculum
and suboperculum. The gill-rakers are of the same
soft structure as in the Carp, but more numerous, the
outer row on the front of the first branchial arch con-
taining about 26 — 30. The lower pharyngeals are

furnished at the outer margin with about 19 si-
milar but shorter spines. At the inner margin they
have a short row of four teeth (fig. 182), the first of
which pretty closely resembles the corresponding tooth
in the Carp, being terete, or even somewhat compressed
at the sides, but with hooked tip and obliquely cut
crown. The three posterior teeth, on the other hand,
are strongly compressed in a transversal direction (in
front and behind), and their crowns are bent close to
each other. The pharyngeal cartilage has about the
same triangular shape as in the Carp.

The body is covered with large, imbricated, and
striated scales, resembling in essential respects those of
the Carp, but with the radiating grooves on the co-
vered (anterior) part of each scale more distinct and
opening in sharp sinuses at the anterior margin of the
scale. As a rule six (4 — 8) grooves appear at this
margin. Another groove usually runs straight or ob-
liquely across the scale, and from the middle of this
groove (the indistinct nucleus of the scale) two or three
similar grooves extend obliquely backwards along the
free (posterior) part of the scale, where the concentric
stria; show undulating breaks. The hind margin of the
scale thus displays numerous and fine, but irregular
notches. The lateral line is straight or slightly sloping,
and lies somewhat nearer to the ventral edge than to
the dorsal. It is covered by 33 (31 — 35) scales. It
terminates sometimes at the middle of the bodv, more
frequently above the vent; and is seldom complete all
the way to the caudal fin. The perforating ducts of
the lateral line are still shorter than in the Carp. In
most cases we find 7 rows of scales above the lateral
line and 6 below it. Sometimes there are 8 rows above
and 7 below, in which case the top and bottom rows
consist of smaller scales. The vent lies somewhat be-
hind the beginning of the hindmost third of the bod}'.

The dorsal fin is long, but shorter than in the
Carp, though its length undergoes even relative increase
with age, varying between about 26 and 32 % of that
of the body. Its height, on the other hand, is generally
greater than in the Carp, being apparently subject to
greater individual variation (about 14 — 18 % of the
length of the body). It begins as far from the tip of
the snout as from the base of the caudal fin, or a litfle
nearer the latter, exactly above the bases of the ventral
fins, or a little farther forward, and ends in most cases
above the middle or the termination of the anal fin.
Its height is almost uniform, or slightly greater in the
anterior half; and its margin is arcuate in adult spe-
cimens, straighter in the young. The rays vary in
number between 20 and 23. The first four rays are
simple, the first of all being an almost imperceptible
spine, most often hidden in the skin, the second some-
what longer, the third more than twice as long as the
second, and the fourth dentated behind on both sides,
soft at the tip, and nearly as long as the first branched
ray. The last ray is cloven to the base, and may
easily be counted twice over.

The anal fin is short but fairly high, with rounded
margin. Its length is about 10 or 11 %, its height
about 12 or 13 % of the length of the body. It con-
tains 9 or 10 rays, the first three simple. The first is
a rather small spine, the second twice as long as the
first, the third thick and hard, with soft and jointed
tip, dentated behind on both sides, and somewhat shorter
than the first branched ray. The last branched ray is
cloven to the base.

The pectoral fins are obliquely rounded at the tip,
and contain 1 simple and (in most cases) 14 branched
rays. Their length may rise from 14 to 18 % of that
of the body. The ventral fins are both broader and
longer, being more obliquely rounded at the tip. Their
length varies between 16 and 19 % of that of the body.
Here, as in the preceding forms, the first ray is short
and set very close to the second, which is thick and
hard, with soft tip, and nearly as long as the first
branched rays. Both pectoral and ventral fins are
generally longer in the males than in the females.

The caudal fin is broad, when folded somewhaf
forked, almost truncate when expanded. The middle rays
occupy 13 — 18 % of the length of the body, and in adult
specimens are 2/3 as long as the longest lateral rays.

The colouring of the Lake Crucian Carp is very
variable, and is closely connected with the nature of

738

SCANDINAVIAN FISHES.

the water inhabited by the dish. It is purest and most
handsome in lakes with clear water and weedy bottom.
In such cases the back is olive green, the sides are
lighter and suffused with brassy yellow, growing still
lighter towards the belly, which is whitish yellow. The
top of the head is of a darker olive green than the
back; its sides are coloured with a handsome brassy
yellow. In muddy lakes with turbid water the colora-
tion is darker, the back and the top of the head being
dark green, the suffused colour on the sides hardly
visible, and the belly dark yellow. The iris is yellow
in specimens that live in clear water ; but the darker
the colour of the body, the greater the density of the
dark dots with which the iris is punctated, and which
in these specimens give it a brownish or pure brown
appearance. The colour of the dins is no less variable.

It conforms to that of the body, and thus becomes
darker or lighter. The dorsal and caudal fins are
generally grayish, and in colour closely resemble the
back. The pectoral, ventral, and anal fins are more
reddish brown. During youth, “up to a length of at least
42 mm.“, the Crucian Carp is marked on the peduncle
of the tail with a whitish band and, in front of this,
with a black band, sharply defined posteriorly” (Malm).

The common Crucian Carp may undergo the same
change of colour as its nearest congener, the Goldfish.
Pallas states that the most handsome specimens of this
golden form occur in the desert lakes along the course
of the River Ural and in its salt tributary, the So Le-
na j a. v. Siebold observed a similar, but stunted form
(of the following variety) in small, stagnant pieces of
water near Braunsberg and Konigsberg in East Prussia.
B: The Bond Crucian Carp ( Cyprinus gibelio, Bl.,
auctt.) — Plate XXX, fig. 3.

In Sweden the Pond Crucian Carp is usually of
insignificant size. Its ordinary length is about 7 — 12
cm., while the largest specimen Ekstrom saw measured
nearly 2 dm. In 1835 Fries took a specimen 23 cm.
long at Morlanda (Bohuslan); and Malm mentions the
capture of a female 34 cm. long in the pond at the
park of the Gothenburg Horticultural Society.

The body of this variety, as compared with that
of the preceding one, is more elongated, less compressed
in front, and shallower. The greatest depth, which lies
somewhat nearer to the head than in the Lake Crucian ,

Carp, is less than half (about 2/5) of the length of the
body to the base of the caudal fin, or about 1/3 of the
length to the end of the said fin. The greatest thick-
ness is more than 1/3 of the greatest depth.

The shape of the back is the same as in the com-
mon Crucian Carp. The dorsal profile runs in a re-
gular and not very high arch, without any angle at
the beginning of the dorsal fin, and is in general less
strongly depressed between the end of the dorsal fin
and the beginning of the caudal fin than in the pre-
ceding variety. The elevation at the very tip of the
tail (the urostyle), on the other hand, is sometimes
more marked in the Pond Crucian Carp. The belly
from the isthmus to the ventral fins is convex, seldom
flat as in the preceding variety, and between the ventral
and anal fins compressed, almost carinated. The curve
of the ventral profile is nearly equal to or somewhat
sharper than that of the dorsal, whereas in the common
Crucian Carp the former is more gradual than the
latter. From the anal to the caudal fin the depression
is more distinct. The tail is deep and short, though
longer than in the preceding variety, the distance be-
tween the anal fin and the caudal fin being there al-
most exactly half the depth of the body at the end of
the former fin, but here 2/3 thereof.

The head is apparently larger — which is always
the case when the fish acquires a more elongated form
— but its relative length is really the same as in the
Lake Crucian Carp, varying between about 27 % (in
young specimens) and about 22 % (in old) of the length
of the body. Its breadth, however, measured across
the gill-covers, is greater in proportion to the breadth
of the trunk here than in the preceding variety. Seen
in profile, the head also appears shorter and more ob-
tuse than in the latter. The forehead is somewhat
broader and less convex. The snout is blunter, and
the cleft of the mouth ascends more sharply, the lower
jaw closing the mouth like a lid, and the angle formed
by the lower jaw at its articulation being thus rendered
more distinct. The pharyngeal teeth in number and
form exactly resemble those of the Lake Crucian Carp.
The opercula are distinguished by their being more or
less convex, whereas in the preceding variety they are
almost flat. The branchial arches, the branchiostegal
membranes, and the eyes, both in form and position,

a Even at a length of 81 ram. this marking is still present in the specimens preserved in spirits in the Royal Museum. That it may
be even longer persistent in the more elongated varieties of this species, appears from figs. 5 and 6 in Siebold’s Susswasserftsche Mitteleuropas.

CRUCIAN CARP.

739

resemble those of the Lake Crucian Carp; and this ap-
plies also to the nostrils, though they are sometimes
set a little higher.

The scales are like those of the preceding variety
in form, number, and distribution. The lateral line
shows some difference, being generally somewhat more
curved, and lying nearer the back. The position of the
vent is the same.

The dorsal fin, which in the number of the rays
resembles that of the Lake Crucian Carp, is distin-
guished from the latter by the difficulty which often in-
volves the detection of the first ray, and which not un-
frequently causes this ray to be overlooked. In form the
difference is greater, the dorsal fin of the Pond Crucian
Carp being always higher in front and less arcuate at
the margin. The anal fin may lie recognised by its
somewhat greater height and less rounded anterior cor-
ner. The caudal fin, even when strongly expanded, and
even in old and large Pond Crucian Carp, is concave at
the hind margin. The pectoral and ventral fins of the
two varieties are alike in all essential respects. The above

differences in the form of the fins in general range the
Pond Crucian Carp on a level with young individuals.

The coloration of this variety is generally darker
than that of the Lake Crucian Carp. The back is of a
dark olive green, and the top of the head still darker.
The dark tint of the back grows somewhat lighter down
the sides, the inner coat of brassy yellow being more
distinct here than on the back, where it is often in-
visible. The belly is dark yellow. The dorsal fin is
of the same colour as the back; the other fins are gray,
more or less deeply tinged with red. The iris is brassy
yellow, but often rendered brown by the dense dots
with which it is strewn. The pupil, however, always
has a fine, brassy yellow rim.

In the structure and arrangement of the internal
organs there is only one difference: in the air-bladder
of the Pond Crucian Carp the round anterior part is
comparatively larger, and the posterior part not so
sharply curved, shorter, and narrower, a circumstance
which seems to depend on the stronger contraction of
the hind part of the abdominal cavity.

The Crucian Carp, which in Scania and Bohuslan is
called Karussa (probably from the Danish Karudse ), oc-
curs almost everywhere in Sweden, even in the District
of NorrbottenC In Norway and Finland, according to
Collett and Mela, the northern range of the Crucian
Carp is about equally great, extending to at least 66°
N. latC The individuals that occur in these northern
regions do not, however, attain any considerable size.
The species seems also to diminish in size towards the
west, for though even in the District of Gefieborg it
attains a weight of 3 3/4 lbs. (17 hectogrammes), Collett
mentions a specimen 260 mm. long as the largest Cru-
cian Carp from the neighbourhood of Christiania pre-
served in the University Museum of that city; and this
specimen probably weighed no more than 7 or 8 hecto-
grammes. In the central and southern provinces of
Scandinavia the species is extremely common, and it is
here too that it attains the greatest size. In Wermland
it is rarer and smaller. The Crucian Carp is also spread
over the whole of Europe0 and Central Asia, together
with Siberia, eastwards to Amur and China.

a Cf. Underd. Bet. Forsl. Ny Fiskeristadga 1883, Bilaga III, ]

6 Gkimm ( Fishing and Hunting on Russian Waters 1883, p
“about 65° N. lat.”

c Apostolides ( La Peche en Gr'ece , 1883), however, does not

The favourite haunts of the Crucian Carp are small
lakes with shallow water and weedy and oozy bottom.
When it is found in larger pieces of water, as in the
western island-belt of the Baltic or Lakes Wener, Wetter,
and Hjelmar, its occurrence is confined to shallow, weedy,
and muddy inlets. It also thrives well in small ponds,
peat-haggs, and other collections of stagnant water, and
is the only one of our indigenous fishes, except the
Three-spined Stickleback, that is plentiful and multiplies
freely in such localities, for the Carp cannot be regarded
as an indigenous Scandinavian species. Hence the Cru-
cian Carp, like the Carp, is kept in ponds, almost as a
domestic animal, to supply the wants of the table. It
can even put up with polluted water. Pallas also tells
us that the Crucian Carp, which does not even disdain
the salt water of the steppes, is the first fish to make
its appearance in new-formed lakes and fens, of which
fact lie saw instances in the basin of the Isset, a tri-
butary of the Tobol (Siberia).

In such small pools, where there is never a suf-
ficiency of food for any considerable number of fishes,

. 158.

. 13) sets the northern limit of the Crucian Carp’s range in Russia at
include the Crucian Carp among the fishes of Greece.

740

SCANDINAVIAN FISHES.

the Crucian Carp gradually degenerates, and after some
generations assumes the form described above under the
name of the Pond Crucian Carp. This form never oc-
curs in lakes or such localities as were first mentioned
as the favourite haunts of the species. There only the
Lake Crucian Carp is met with; but if specimens of this
variety be put in a small fish-pond, in the course of
some years the pond will be found to contain Crucian
Carp of an intermediate form, difficult to refer to either
variety, and at a still later period only Pond Crucian Carp.

These two forms are consequently not distinct spe-
cies, as was long assumed on Bloch’s authority. It is
probable, however, that the Crucian Carp does not de-
generate with equal rapidity everywhere, but that the
course of degeneration is arrested at an earlier or later
stage, according to the nature of the water and the
supply of food. Small pools are, at all events, often
inhabited by the said intermediate forms, which seem
to have lived and multiplied there for a long time
without further transformation.

That the Pond Crucian Carp resumes its original
form, is shown by the fact that if a few small speci-
mens of this variety lie placed in a- large pond, where
the supply of food is abundant, they soon acquire the
intermediate form. But when they have been allowed
to multiply to any great extent, so that food is scarce,
and the pond overstocked, they revert to the normal
form of the Pond Crucian Carp. Furthermore, we have
actual evidence to show that specimens of the Pond
Crucian Carp, which have been transferred to lakes,
have at length been restored to the original form of
the Lake variety. In a small lake in Sodermanland
Eksteom found large Lake Crucian Carp, the progeny
of Pond Crucian Carp which, according to the state-
ments of trustworthy persons still living at the time,
had been taken 40 or 50 years previously from a neigh-
bouring pond still inhabited by the latter variety.

Eksteom elucidated this question in the Proceedings
of the Swedish Academy of Science for the year 1838;
and at the present time his opinion on the subject is
almost universally accepted.

The Crucian Carp is very sluggish in temperament,
never undertaking any long excursions, and always
staying near its birthplace, unless compelled by fortui-
tous circumstances to change its abode. This innate
distaste for active motion deprives it of any great ti-
midity and renders it oblivious of danger, as it hides
in the ooze at the bottom, where the water is deep, or

among the weeds nearer the shore. When it ascends
towards the surface, which seldom happens except when
the heat of the sun entices it forth, it is however shy,
and takes to flight at any noise. It then makes straight
for the bottom, and there seeks a safe refuge, from
which it is not easy to unearth. Its movements in the
water are often active and speedy, though they lack
endurance.

The spawning-season occurs in spring and summer,
some days earlier or later according to the situation of
the water where the fish has its home. In Central
Sweden the Crucian Carp spawns in June, about mid-
summer, when the weather is warm and tine. The fish
then ascend to shallow water near shore. Here they
spawn among the weeds, assembling in dense shoals
and circling rapidly round each other, now and then
raising their snouts above the surface, where each fish
leaves a bubble that bursts with a faint sound. A
murmuring hubbub, like the noise of boiling water, is
thus produced. The roe is deposited at the bottom on
the weeds, to which it adheres, and is soon hatched
when the weather is favourable. The fry do not grow
very rapidly, but soon reach maturity. This faculty
of early reproduction, combined with the comparatively
large size of the ovaries and the smallness of the eggs,
their diameter, when deposited, being only slightly more
than 1 mm., as well as their great number (100,000
— 300,000 in each female), renders the Crucian Carp
extremely prolific. The duration of the spawning-
season is protracted, in this species as in others, by
the older fish spawning first, the younger ones later in
the year. But the erroneous supposition that the Cru-
cian Carp spawns several times a year, has arisen from
the circumstance that during the months of July and
August, when the weather is calm and fine, these fishes
shoal in exactly the same manner as when they are
spawning.

The Crucian Carp is a glutton, and lives on insects,
worms, water-plants, and mud. The flesh ranks as a
delicacy, but its reputation depends principally on the
skill of the cook. Large Crucian Carp, taken in clear
water, are best and by no means bad eating; but small
specimens from muddy lakes always have a more or less
tainted flavour, and their flesh is also bony.

The Crucian Carp is extremely tenacious of life.
In lakes and ponds where the bottom freezes, it can
survive the winter, and after being kept frozen for a
long time, may be restored to life by cautiously thawing

CRUCIAN CARP.

741

the ice. How great sufferings it is capable of enduring,
may be gathered from Ekstrom’s account of a specimen
described in his “ Morhofiskar" . “This specimen was pro-
cured from the island-belt, where it had been kept
several days in a cauf. About five o’clock in the mor-
ning it was taken out of the cauf and carried to the
rectory. It was then laid on a gutting-board and left
there during the whole time occupied in describing and
drawing it. About six o’clock p. m., when the fish had
been out of the water quite thirteen hours, it was cut
in two close to the beginning of the dorsal tin. The
heart, liver, etc. were removed, and the surface of
section was drawn. When all this was finished, and
the pieces were to lie thrown away, the forepart of the
fish still showed signs of life, the gill-covers being
opened and closed, and the mouth protruded and drawn
back. I left the pieces where they lay, in order to
observe how soon all manifestations of life would cease.

Not until nine o’clock in the evening had all signs of
vitality disappeared. The fish had thus lived sixteen
hours without water, three hours cut in two and with-
out heart.”

Of the age attained by the Crucian Carp E inn a-: us
tells us ( Skanska JResan, p. 256) that a Crucian Carp
had lived “certainly more than 70 years” in a spring
at Ma in the parish of Svenskop. During all this time
it had not grown to a greater length than 6 in. (15
cm.), and the colour of the back was very dark. “The
meagre water of the spring had probably denied it a
sufficiency of food.”

The Crucian Carp is taken chiefly in traps (see
above, p. 33, fig. 7), during the spawning-season in
gill-nets, and often in trammel-nets''. In the last case
the water should be thick, for otherwise the fish seldom
suffers itself to lie driven into the net, but buries it-
self in the mud. (Ekstrom, Smitt.)

a Fr. tramail (trots mailles'), Svv. skottnat, skotnot, or pul snot. In this fishery, which is generally employed for the Cyprinoids, and
which is unfortunately very destructive if practised during the spawniug-season, the necessary tackle comprises, besides the coble, the net
itself (fig. 183), the ‘shoes’ (Sw. kabbarne , fig. 184), a pole, and a ‘beater’ (Sw. puls, fork, or terfvel , fig. 185). The depth of the net is
between 12 and 16 dm., the length 14 — 16 metres if the net is to be managed by one man, but much greater when there are two fisher-
men. Besides the net itself (the inner net — fig. 183, G), which resembles an ordinary gill-net, and is made of very fine twine, the size

of the meshes depending on the kind of the fish for which the net is to be used, the trammel should consist of an outer net (Sw. grimma )
with very large meshes, and made of coarse and strong twine. The outer net should be equal in length to the inner net, if it is to extend
along only one side of the trammel, or twice as long, if, as is usually the case, it is to cover both sides (fig. 183, E and F). When

74*2

SCANDINAVIAN FISHES.

Genus GOBIO.

Base of flie dorsal fin less than twice as Jong as that of the anal. Branched rays in the dorsal fin at most 8,
in the anal at most 7. Neither of these fins with any spiniferous ray. Scales middle-sized — 40 or a few more
in the lateral line — and rather thin. Distance between the anal fin and the vent about equal in length

to the base

In this genus and the following one we pass fr.om
the Macroenteric Cyprinoids and also draw nearer to
the Leuciscine group, the dorsal tin being reduced both
in the number of the rays and the length of the base.
The external form of the body is also approximated
to the Leuciscine type. Still its affinity to the genus
Barbus , which abounds especially in India, connects
Gobio with the true Carps. The genus Gobio was

of the fin.

established by Cuvier" as a subgenus of Cyprinus /
but Fleming6 was the first to adopt it as the name of
a distinct genus. Only two species, both European,
have hitherto been recognised within this genus. One
of them, the smaller, but more elongated of the two
( Gobio uranoscopus ), belongs exclusively to Austria and
Bavaria. The other is

tlie fish comes in contact with the inner net, it pushes a part thereof in front of it through one of the meshes of the outer net, and is
thus enclosed in a kind of pocket, from which it cannot retreat. The size of the meshes in the outer net varies according to the depth of
the trammel, which should be five meshes deep. In the inner net, which is to form the pockets, the netting should be deeper, a trammel
12 dm. deep requiring an inner net 161/, dm. in depth. The art of constructing a trammel-net is difficult enough; but space permits us
only to refer the reader to the figure. The head rope (Sw. flarntelnen ) should, if possible, be furnished with round floats (Sw. flam ) of pine-
bark, attached at a distance of 17 — 20 cm. from each other. The foot rope (Sw. stentelnen ) should be weighted with plummets of lead.
At each end of the head rope a so-called ‘shoe’ is fastened. The shoe is made of wood, in the form of a pointed and hollow cone, and
from its base there projects a wooden disk, pierced at the end with a hole, by means of which the shoe is attached to the head line.

In addition to the net described above the fisherman should have a coble or punt, furnished at the prow with a ring of iron, rope,
or twisted rushes, about 15 cm. in diameter. The pole , with which the net is shot, is made of spruce, and should be 7 or 8 metres
long, not too thick to be easily grasped by the hand, and not so thin as to bend beneath the weight of the net. The tackle should be
completed by a so-called ‘beater’, with shaft 4 or 5 metres long. The shaft should consist of a thin stake, sharpened like a sword at one
end to enable the fisherman to drive it with ease into the bottom, and furnished at the other end with a lump of wood which varies in shape,
but generally resembles a hemisphere hollowed into the form of a funnel. When this end of the ‘beater’ is thrust into the water, the air
contained in the hollow is forced below the surface, and increases the noise of the blow.

Armed with these implements, the fisherman betakes himself some fine summer day to a shore overgrown with grass or reeds. When
he has found a suitable spot, he passes the pointed shaft of the beater through the ring at the prow of the coble, and drives it into the
bottom hard enough to moor his boat safely. Then he lets down the net, which should have been properly arranged beforehand, into the
water at the bows, takes the pole, inserts its end into the shoe, and thus shoves out the net in an oblique direction towards the shore. When
he has spread half the net to the full length of the pole, he suddenly plucks the latter out of the shoe, and the net stays in position. He
now inserts the tip of the pole in the same manner into the shoe at the other end of the trammel, and sets this part of the net in a si-
milar way, so that, when both halves of the net are shot, it lies in the form of a snow-plough (V), with the entrance of the angle turned
towards the shore, and its point towards the bows of the coble. The fisherman next draws up the ‘beater’, adjusts the middle part of the
net, the part which the length of the pole has not enabled him to set before, and with the aid of the ‘beater’ pushes the coble round one
end of the net towards the shore. Having got as close in shore as possible, and keeping the net in front of him, he turns the other end
of the ‘beater’ downwards, and plunges it into the water in the direction of the net, thus driving the fish before him, and gradually pushing
the coble forward, until he is able to touch the net with the ‘beater’. With the same implement he then lifts the foot rope of the net
above the surface, takes the foot rope and head rope in his hand, and draws up the net into the boat. After arranging the trammel by
taking one of the ropes in each hand and laying the net carefully in the coble, he proceeds to another spot, where he repeats the operation.

When the trammel-net is so large that two men are required to manage it, it is set like a seine in a semicircle, and when they are
ready, one of the fishermen rows the coble from land towards the trammel, while the other stands in the boat and with the ‘beater drives
the fish into the net.

a R'egn. Anim., ed. 1, tom. II, p. 193.

6 Brit. Anim., p. 186.

CARP-FISHES.

743

THE GUDGEON (sw. sandkryparen or slattingen).

GOBIO FLUVIATILIS.

Plate XXXI, fig. 3.

Least depth of the tail about 1/3 (30 — 35 %a) of the length of the head. Dorsal and anal fins (sometimes the
other fins also) with spotted rays. Pharyngeal teeth set in two rows , the largest ones with compressed , simple ,

obliquely set masticatory surface: the tip hamate : 2(3), 5 — 5(4), 2(3)''.

a b

Fig. 186. Pharyngeal bones and pharyngeal cartilage of Gobio fluviatilis. Magn. 3 diain. a, b, and c as in the preceding figure.

2—3 2—3

R. hr. 3; D. ; A. - V. -

7—8 6—7 7-

C. a + 1 + 17+1 + sc; Lin. lat. 41 c — 45 ; Lin. tr. - 1 ;

4

; P- — — ;
’ 13—16

Vert. 37f— 41.

Syn. Gobio , Auson., Mosell., vers. 132; Gobio fluviatilis , Rondel.,
De Pise., part. II, p. 206. Fundulus, Chari.., Onomast.
Zoic , p. 157; Schonev., Ichthyol. Slesv. Hols., p. 35.
Cyprinus quincuncialis maculosus, maxilla superiore longiore,
eirrhis duobus ad os, Art., Ichthyol., Gen., p. 4; Syn., p. 11;
Spec., p. 13. Gudgeon , Penn., Brit. Zool. (ed. 1776) tom.
Ill, p. 316.

Cyprinus Gobio, Lin., Syst. Nat., ed. X, tom. I, p. 320; Bl.,
Fisch. Deutschl.., part. I, p. 57, tab. VIII, fig. 2; Retz ., Fn.
Suec. Lin., p. 355; Pall., Zoogr. Ross. Asiat., tom. Ill, p.
295; Agass., Isis, vol. XXI (1828), p. 1049, tab. XII, fig.
2; Nilss., Prodr. Ichthyol. Scand., p. 33; Schagerstr.,
Physiogr. Sallsk. Tidskr. 1837, p. 295; Gthr ( Leuciscus ),
Jahresh. Ver. Vat. Natnrk. Wiirttemb., IX (1853), p. 268.
Gobio fluviatilis, Flmng, Brit. Anhn., p. 186; Cuv., Val., Hist.
Nat. Poiss., vol. XVI, p. 300, tab. 481 (+ Gob. obtusi-
rostris, p. 311); Bonap. ( + Gob. venatus ) Iconogr. Fna Ital.,
Pesc., tab. 110, figg. 5 et 6 ; Kr., Danm. Fisk., vol. Ill,
p. 334; Nilss., Skand. Fn., Fisk., p. 300; Dybowski, Cy-
prinoid. Livl., p. 72; Sieb., Siisswasserf. Mitteleur., p. 112;
Mgrn, Finl. Fiskfn., p. 40; Canestr., Arch. Zool., Anat.,
Fisiol., vol. IV, fasc. 1 (1866), p. 80; Gthr, Cat. Brit.
Mus., Fish., vol. VII, p. 172; Buckl., Nat. Hist. Brit. Fish.,
p. 100; Bncke, Fisch., Fischer., Fischz. O., W. Preuss.,
p. 115; Mob., Hist. Nat. Poiss. Fr., tom. Ill, p. 386;
Fatio, Fn. Vert. Suisse, vol. IV, p. 280; Mela, Vert. Fenn.,
p. 320, tab. X; Day, Fish. Gt. Brit., Irel., vol. II, p. 172;
tab. CXXXT, fig. 2; M5b., Hcke, Fisch. Osts., p. 107; Her-
zenst., Warp., Not. Fischf. Amurb., p. 28; Lillj., Sv. ,
Norg. Fn., Fisk., vol. Ill, p. 161.

a Sometimes 39, according to Kroyer.

,, 4, 6 — 5, 3, according to Fatio.

,, 36, according to Moreau.

Gobio vulgaris, Hckl., Kn., Siisswasserf. Oestr. Mon. p. 90.

Gobio Benacensis, Pollini; Gobio lutescens, De Fil.; Gobio
Pollinii, De Betta; vide Fatio, 1. c., p. 294.

The Gudgeon, according to Bloch, attains a length
of 8 in. (21 cm.), and Pennant was told of a specimen
that weighed half-a-poimd; but in Scandinavia it always
belongs to the small fishes, being at most about 15 cm.
long. The body is elongated and fairly thick, with
broad, convex back, flat belly, and more or less per-
pendicular sides. The greatest depth of the body, which
occurs just in front of the beginning of the dorsal tin,
measures about 16 — 1 8 1/2 % of its length, and the
greatest thickness, across the opercula, is about 2/s of
the greatest depth. The least, depth of the tail is about
8 (sometimes 8V2) or 7 % of the length of the body.
The dorsal profile thus forms a very elongated curve,
which generally shows quite a distinct depression at
the occiput. The ventral profile is straighter in front
and ascends, strictly speaking, only behind, at the be-
ginning of the anal fin.

The form of the head, bluntly pointed in front,
otherwise passes evenly into that of the trunk, but is
diversified in the nasal region, just in front of the eyes,
by a swelling, which, according to Lilljeborg, is larger
in the males than in the females — a statement of which
we have failed to find corroboration in our specimens

94

Scandinavian Fishes.

744

SCANDINAVIAN FISHES.

— and which in its most advanced development has
given rise to the establishment of a distinct species,
Gobio obtusirostris. This swelling — even when it is
not very prominent — and also the projection beyond
the mouth of the blunt tip of the snout most strongly
characterize the head of the Gudgeon. The length of
the head varies between about 1/i and a little more
than * 1/5 (26 — 22 %) of that of the body, being as usual
comparatively greater during youth. The eyes are ex-
tremely mobile, round, set fairly high — their upper
margin being almost in a plane with the slightly con-
vex forehead — and rather large, their longitudinal
diameter varying between about 7 and 478 % of the
length of the body or about 27 and 21 % of the length
of the head. In young specimens this longitudinal dia-
meter is about equal to the breadth of the interorbital
space, in old less, sometimes only 7/io thereof. The posi-
tion of the eyes is also such that the length of the snout

— with considerable individual variations" — measures
on an average 2/5 of that of the head. The nostrils lie
on the rounded margin between the sides and back of
the snout, much nearer to the eyes than to the tip of the
snout. Together they form on each side a round cavity,
divided externally by the oblique partition Avail betAveen
them, which is raised into a lid. The mouth is small
and lies, as Ave have mentioned, entirely on the under
side of the head, with the maxillary bones extending
back to about a line Avith the anterior nostrils and
Avith the articulation of the loAver jaAv in a line Avith
the anterior margin of the orbit. The lips are fleshy,
and the dermal fold that, lies on each maxillary bone
is elongated behind into a barbel of moderate length,
Avhich extends when laid back hardly any distance, if
at all, behind the perpendicular from the anterior mar-
gin of the eye6. The palatal curtain Avithin the upper
jaw is well-developed. The tongue is fleshy, but hardly
at all free from the bottom of the mouth. The gill-
rakers are short, scattered, blunt, and soft, developed
on the first branchial arch only in the inner (posterior)
row, which contains about 8. Among the pharyngeal
teeth (fig. 186) only the three posterior in the inner
roAv have a distinct, obliquely cut masticatory surface,
Avhile the others — both the two anterior, thicker teeth
in the inner row and the tAvo or three teeth in the
outer i*oav — are of a straighter cylindrical form, though

obliquely hooked at the tip. The pharyngeal cartilage
is soft, scarcely cartilaginous, and triangular, with the
upper (anterior) side slightly curved upAvards (convex).
The pseudobranchim are distinct on the inside of the
upper part of the hyomandibular bone, though small,
with about 6 filamentous lamellae. The operculum is
trapezoidal, its upper and loAver sides being parallel,
but the former only about 2/3 as long as the latter and
throughout united to the dorsal side. The branchiostegal
membrane is extraordinarily narroAV along the hind
margin of the operculum, but all the broader along the
suboperculum and in its loAver part, where it is ex-
tended by three, fairly broad, sabre-shaped rays. The
height of the gill-openings is about equal to the post-
orbital length of the head; their upper angle lies hardly
on a level Avith the superior margin of the eye, and
their loAver angle lies in about a line Avith the hind
margin of the preoperculum, Avhere the branchiostegal
membranes on the ventral side coalesce Avith the
isthmus.

The dorsal fin is trapezoidal, Avith the upper mar-
gin straight or slightly concave and the high anterior
margin composed of the first three (simple) rays. The
first of these rays is extremely short, sometimes scar-
cely possible of external detection, Avhile the second is
nearly half as long as the third. The fin begins at a
distance from the tip of the snout,' Avhich measures on
an average 42 % (varying between 41 and 43 %) of
the length of the body, and its length (base) is 11 or
12 % (in the males sometimes nearly 13 %) of the
same. Its height (the length of the third ray) is always
greater than its length and varies between about 16
and 21 % of the length of the body.

The anal fin is of the same form and structure as
the dorsal. Its distance from the tip of the snout is
on an average 62 % (varying betAveen 60 and 63 %),
its length ll/2 % (varying betAveen 7 and 8 7 2 %), and
its height 1372 % (varying betAveen 12 and 16 %) of
the length of the body.

The caudal fin is deeply forked at the hind mar-
gin, most so in old specimens, and thus forms an ob-
tuse angle Avith soineAvhat convex edges to the lobes.
The length of the middle rays, Avhich occupy about
11 — 9 % of the length of the body, is as a rule in
young specimens equal to or a little more than half,

° Varying between 34 and 45 / of the length of the head.

6 Canestrini, however, mentions a specimen, a male 741/0 mm. long, in which the length of the barbels was somewhat more than

GUDGEON.

745

in old specimens a little less than half that of the
longest outer rays.

The paired fins are similar to each other in general
form, when the ventrals are not fully expanded. Both
pairs are of an oblique oval shape; but the pectoral
fins are more pointed arid longer. The length of the
latter varies between about 20 and 15 %, that of the
ventral fins between about 15 and 13 % of the length
of the body. In their position too, these paired fins
resemble each other, the pectoral fins when expanded
being drawn down to the flat ventral margin, where
they project horizontally outwards like the ventral fins.
The position of the latter is such that the distance from
the tip of the snout to the foremost (outermost) point
of their insertion is about 43 % of the length of the
body, and from the said point in the insertion of these
fins to the corresponding point in that of the pectorals
about 23 % of the same.

The scales are of moderate size, but rather thin,
and their form varies considerably, being rounded and
quadrangular, semi-elliptical (rounded behind), or round-
ed and triangular. In the anterior (covered) part, which
is the shorter, but deeper, we find only concentric, fine
strife; the posterior part, on the other hand, is marked
with a number (as many as 24) of radiating strife, the ends
of which at the hind margin of the scale form fine and
fairly regular notches in this margin. The lateral line is
fairly straight, in front slightly curved in a downward di-
rection, and keeps to the middle of the sides of the body.

In coloration the Gudgeon is one of the most sin-
gular of our Cyprinoids, not indeed for any remarkable

beauty of coloui though in this respect it is but

little inferior to our most handsome species, the Min-
now — but for its possession of more varied hues than
most of the Scandinavian forms of this family. The
back is grayish green, darker (shading into brown) or
lighter according to the colour and light of the envi-
ronments of the fish, and transparent when the light
falls directly upon it. The belly is milky white. The
colour of the back extends forward on the top of the
head, the snout, and the outside of the barbels. On the
upper part of the sides of the snout, just below the
nostrils, appears a dark, fairly broad, longitudinal band,
which is continued across the upper part of the eye
more or less distinctly on the temple. Below the eye

too, lies a more or less distinct band, formed by more
or less densely agglomerated collections of dark pig-
ment. Similar irregular spots also occur on the oper-
culum and preoperculum; but the sides of the head are
otherwise coated with a brassy and silvery lustre finely
punctated with brown. The lower jaw and the branch-
iostegal membrane are of the same colour as the belly.
The iris is below of a silvery lustre, but finely punc-
tated with brown. The silvery lustre of the sides of
the body shades more or less distinctly into violet, and
this bluish colour collects at certain points into a more
or less regular row of undefined spots above and along
the lateral line, now of a lighter blue, now shading
even into blackish brown. The scales of the lateral
line are marked, at least in the anterior part there-
of, by a dense agglomeration of dark brown pigment
on each side of the canal itself; and more scattered
dots of similar pigment at the margins of the scales
form a network of brown over the whole body. All
the fins are light and transparent, of a more or less
pronounced yellowish green. This last colour is most
prominent on the rays, which are also marked — -in the
dorsal and caudal fin always, in the other fins some-
times, but with far less density — with brownish, elon-
gated spots, together forming more or less regular trans-
verse bands and most distinct on the caudal fin.

The most trustworthy external distinction between
the sexes during the spawning-season is the dermal
eruption, consisting of small, fine, verrucose tubercles
on the head and the forepart of the back, which is
general among the male Cyprinoids. A dark blue spot
on the gill-cover, more prominent than in the females,
is also a usual characteristic of this sex during the said
period. Furthermore, according to Fatio, the longest
rays in the pectoral fins of the male are thickened during
the spawning-season; but we have failed to find any e A i-
de ns e of this. The most general difference between the
sexes lies, however, in the longer and higher fins of the
males; but the changes of growth — young specimens
of both sexes being similar in this respect — impair the
validity of the sexual characters drawn from this rela-
tion. We have, however, found one of these characters
to hold good in all our specimens: in all the males
the length of the lower jaw is less, in the females
more, than 55 % of the length of the ventral fins".

“ The position of the vent, which in other cases gives the most easily expressed sexual character, and which has been stated in the
Gudgeon to lie in the females behind, in the males in front of, the middle of the body, varies so considerably that in a male 107 mm. long
for example we found the distance from the vent to the tip of the snout to be 58 mm.

746

SCANDINAVIAN FISHES.

The internal organs agree with the description given
above of the structure of the Brachy enteric Cyprinoids.
The stomach extends back to about a line with the in-
sertions of the ventral tins, from which point the in-
testine first runs forward, to the left of and above the
stomach, until at the very front of the abdominal cavity
it bends suddenly back to return straight to the vent.
The entire length of the digestive canal is only about
3/5 of the length of the body. The dorsal part of the
musculature contains on each side an ellipsoidal organ
of hitherto unexplained signification, proximally and
dorsally attached to the fourth rib (the first of the nor-
mal ribs). This organ is white and fairly large, being
nearly 3 mm. long in a Gudgeon 110 mm. in length.
It consists of transversely striped muscle fibres, exter-
nally longitudinal, internally transversal, and is attached
by connective tissue to the said rib. As for its mor-
phological significance, Ave can find no better comparison
than with the above-mentioned (p. 704) introitus cap-
sules vesica in the Cobitoids.

The Gudgeon has a wide geographical range to the
east and west, from Amur and China" to England and
Ireland. To the north and south its range extends in
Europe from Southern Finland (Lat. 62° or 63° N.), the
extreme south of Sweden, and Liim Fjord, to Central
Italy. In Greece and Spain it is unknown. Its Swedish
range consists principally of Scania, where it is fairly
common in several localities. When Retzius in 1800
introduced it into the fauna, of Sweden, it already pos-
sessed at Lake Finja a special Swedish name, Slatting
{slat — smooth). The species is also known to occur in
Scania in Lake Vester and the Ronne River to the west,
where according to Schagerstrom it is called Oral an-

o

ning, and in the streams of Helge and Arup to the east.
From the River Morrum in Blekinge Baron G. C. Ce-
derstrom in 1856 forwarded several specimens to the
Royal Museum. The Zoological Museum of Upsala,
according to Lilljeborg, has received specimens through
Dr. Scheutz from the neighbourhood of Wexio. Accord-
ing to Nilsson the Gudgeon also occurs in the River
Nissa at Halmstad and is there called stensugare (Stone-
sucker)* 6. Krdyer did not know it from the Danish is-
lands; but in recent times it has been found in Zealand
(Feddersen). In Jutland it is common south of Liim
Fjord. In Germany and the Baltic Provinces of Russia

it is also common, as well as in Central and Western
Europe. It is one of the inhabitants of the Cavern of
Adelsberg. In the water-courses of Switzerland it ascends,
according to Fatio, to a height of about 800 metres
above the level of the sea; south of the Alps it is of
rarer occurrence.

The habits of the Gudgeon are fairly well expres-
sed by its Swedish name of Sandkrypare (Sand-creeper).
As it lies, usually in companies, close to the bottom,
with the paired fins horizontally expanded, on a bed of
sand, gravel, stone, or even of mud, it still keeps a
careful watch, and by fits and starts displays great ac-
tivity. It takes a, hook freely. It is fairly tenacious of
life, but like all the Cyprinoids, sensitive to changes in
the weather and incapable of enduring bad respiration.
The living specimens in our aquaria kept close to the
bottom and remained still in a packed mass, as long as
the water was fresh; but as soon as it had stood an
hour or so the crowd dispersed, and its members ascended
severally to the upper layers of the water. In spite of
this the Gudgeon can live in muddy or even polluted
streams, and readily haunts the openings of sewers,
provided they are in running water.

Valenciennes" experimented on this fish in order
to ascertain the effect produced on it by atmospheric
rarefaction. The air-bladder became completely empty,
its gas passed into the intestine, the belly swelled up,
and the fish floated on its back, but Avas alive and
seemed to steer its course in the Avater. After 24 hours’
continued operation of the air-pump he restored the
fishes to a normal atmospheric pressure; and after the
lapse of 6 hours they Avere quite recovered, Avith the
air-bladder full almost exclusively of nitrogen.

As Ave have remarked above, the comparatively
short intestinal canal of the Gudgeon indicates that its
food is chiefly animal. It lives principally on insects
and their larvae, crustaceans, Avorms, and the roe and
fry of various fishes; but it is fond of rooting up its
food in sand and mud, and also devours decomposing
substances, both animal and vegetable.

The spaAvning-season of the Gudgeon occurs in
Sweden at midsummer or earlier — in more southern
countries even at the beginning of April. Before this
time it has ascended from its winter-quarters to shal-
loAver spots in rivers or brooks. In the River Hofdala,

a Herzenstein and Warpachowsky, 1. c.

6 Renewed attempts to procure the Gudgeon from this locality have, however, failed.
c Cuv., Val., 1. c., p. 14.

GUDGEON.

747

according to Baron Cedeesteom, the Gudgeon is often-
est taken in early summer in traps ( rgssjor , see above,
p. 33, fig. 7), which are set in the middle of the river
against the stream. The Gudgeon ascend the river in
the evening to spawn", but in the morning return to
Lake Finja. On the 25th of May, 1858 Cedeesteom
observed the Gudgeon spawning in this locality; he was
told that the spawning had already lasted a week here
and along the shores of Lake Finja. Rusconi describes
the spawning of the Gudgeon in Lombardy as follows6:
“During my stay at Desio, on a most lovely day in
July, I was walking early in the morning along the
shore of the little lake of Villa Traversi. Suddenly a
noise reached my ear. I thought at first that some
one was beating the water with sticks or with the flat
of an oar. On glancing along the shore 1 soon detected
the spot from which the sound proceeded, as well as
the cause of this disturbance: it was caused by spawn-
ing fishes. Eager to obtain a closer view of this sight,
I stealthily made my way towards them, and under
cover of the bushes that fringed the shore, 1 got near
enough to observe them with ease and without betray-
ing my own presence. They lay at the mouth of a
small brook, the water of which was cool and clear, but
so scanty that the pebbles at the bottom were almost
dry. They Avere Gudgeons. They approached the mouth
of the brook. With rapid strokes they came swiftly on
and advanced about a. metre up the brook, not leaping,
but in a manner gliding over the pebbles. After this
first spurt they stopped, bent the trunk and tail alter-
nately to the right and left, and in this Avay rubbed
the ventral side against the bottom. With the excep-

tion of the belly and the loAver part of the head their
whole body iioav lay out of the water. They retained
this position for seven or eight seconds. Then they
dealt a sharp bloAv with the tail on the bottom, splashing
the water in all directions, turned round, and darted
back to the lake, soon to repeat the same operation”.

The eggs are comparatively large Avhen deposited,
about 1 1/2 mm. in diameter, but also rather few in
number, at most some thousands in each female. They
are transparent, with a dash of blue or yellow. Valen-
ciennes supposed that the Gudgeon spawns several times
a year. The only evidence of this seems to be that the
female does not deposit her roe all at once, but bit by
bit. That the spaAvning-season is of long duration,
hoAvever, is shoAvn by the fact that Fatio in SAvitzerland,
and Lilljeborg here in the North, found females fall of
roe, but not yet ready to spaAvn, late in July0’.

To the angler the Gudgeon affords excellent sport.
It readily takes a bait of flies or worms; and tenacious
of life as it is, it may be used Avith advantage as live
bait for larger fish. Its greatest value lies, however,
in the flesh, Avhich is of delicious flavour and easy of
digestion. Buckland Avrites: “When out gudgeon-fishing
on the Thames, be sure and take a frying-pan, as gud-
geons taken out of the Avater and immediately fried are
delicious. Clean, Avipe, and Hour, then Avell fry in
boiling fat, or, better, in oil, till they are crisp and of
a light brown colour. Such a fish dinner is ahvays a
great feature in a pic-nic on a fine day”. A someAvhat
similar method is employed in Scania in the prepara-
tion of fiskaltaga (fish-cake) from Gudgeons and other
small fishes.

Genus TINCA.

Base of the dorsal fin less titan twice that of the anal. Branched rags in the dorsal fin at most 9, in the anal
at most 8. Neither of these fins furnished with, a spiniferous rag. Scales small and thin — their number in
the lateral line at least about 90. Distance between the anal fin and the vent onlg V2 or V3

of the base of this fin.

By the deeper form of the body this genus is more comes nearer the Leuciscines, among Avhich the MinnoAv

closely approximated to the true Carps; but it also with its small scales reminds us most of Tinea. Only

a According to Fatio the Gudgeon spawns in the daytime.
b See Bbehm, Thierleben , Aufl. 2, Abth. 3, Bd. 2, p. 275.

c Valenciennes also states that the spawning lasts from April to the end of July or the middle of August. The Gudgeons we re-

ceived this year (1891) at the beginning of June from the Finja, had just commenced spawning.

748

SCANDINAVIAN FISHES.

genus is the same as that of the preceding one. The
name is derived originally from Ausonius (in the fifth
century) and Rondelet (in the sixteenth century). It
was adopted as a specific name by Linnaeus, as the
name of a subgenus of Cyprinus by Cuvier6, and raised
by Fleming c to the rank of a generic name.

THE TENCH (sw. LINDAREN OF. SUTAREN).

TINCA VULGARIS.

Plate XLI, fig. 4.

Barbel at each corner of the mouth small. All the fins rounded. Mouth situated at the tip of the snout, be-
ginning of the dorsal fin just in front of the middle of the body. Muciferous pores in the cephalic system of
the lateral line especially distinct in the supraorbital , suborbital, and preoperculo-mandibular branches. Pharyngeal
teeth, set in a single row, with narrow, one-grooved masticatory surface: 4(5) — 5(4).

one species of this genus is known, for the Siberian
species, Cyprinus perenurus" , which Pallas established,
and which has otherwise been referred to this genus,
is based merely upon a defective description in the
manuscripts left by Steller and has been explained
by Warpachowski as a Phoxinus. The history of the

Fig. 187. Pharyngeal teeth and pharyngeal cartilage of Tinea vulgaris. Natural size.

a, b, and c as in the preceding figure.

R. hr. 3 ; D.

3(4) 3(4).

8(9)’ ‘ 7(8)’

P. ; V.

15 — 17

2

(8)9 :

30—33

C. x + 1 + 17+ 1 + x; L. lat. (90)100 — 114(120); L. tr. — — 1;

Vert. 37—41.

Syn. Tinea, Auson., Mos., vers. 125; Rondel., De Pise., part. II,
p. 157; Bel., Nat., Div. Poiss., p. 325; Schonev., Ichthyol.
Slesv. Holst., p. 76. Cyprinus mucosus totus nigrescens,
extremitate caudse cequali, Art. Ichth., Gen. Pise., p. 4; Syn.
Pise., p. 5 ; Descr. Spec. Pise., p. 27. Cyprinus pinna ani
ossiculis undecim, cauda integra, Lin., Fn. Suec., ed. I,

p. 122.

Cyprinus Tinea, Lin., Syst. Nat., ed. X, tom. I, p. 321 ;
Bl., Fisch. Deutschl., part. I, p. 83, tab. XIV ( + Per
Goldschlei, p. 90, tab. XV); Retz., Fn. Suec. Lin., p. 354;
Pall., Zoogr. Ross. Asiat., tom. Ill, p. 296; Ekstr., Vet.-
Akad. Handl. 1830, p. 200; Nilss., Prodr. Ichth. Scand.,
p. 34; Ekstr., v. Wr., Skand. Fisk., ed. 1, p. 205, tab.
52; Gthr ( Leuciscus ) Jahr. Ver. Vat. Naturk. Wiirtemb., IX
(1853), p. 274; Reut., Sundm. {Tinea) Finl. Fisk., tab. VIII.

Tinea vulgaris Flmng, 1. c.; Cov., Val., Hist. Nat. Poiss.,
vol. XVI, p. 322, tab. 484; Yap.r., Brit. Fish., ed. I, vol. I,
p. 328; Kr., Damn. Fiske, vol. Ill, p. 351; Nilss., Skand.
Fn., Fisk., p. 297; Hckl, Kn., Siisswasserf. Oestr. Mon.,
p. 75; Dybowsky, Cypr. Livl., p. 66; Sieb., Siisswasserf.
Mitteleur., p. 106; Mgrn, Finl. Fiskfn. (disp. Helsingf.
1863), p. 39; Canestr., Arch. Zool. Anat., Fisiol., vol. It',
fasc. I (1866), p. 69; Blanch., Poiss. d. eaux donees Fr.,
p. 317; Lindstr., Gotl. L. Hush. Sallsk. Arsber. 1866, p.
17 (sep.): Gthr, Cat. Brit. Mas ., Fish., vol. VII, p. 264;
Coll., Forh. Vid. Selsk. Chrnia 1874, Tillaegsh., p. 183;
Mor., Hist. Nat. Poiss. Fr., tom. Ill, p. 383; Bncke, Fisch.,
Fischer., Fischz. O., TV. Preitss., p. Ill; Mela, Vert. Fenn.,
p. 318, tab. X; Day, Fish. Gt. Brit., Irel ., vol. II, p. 188,
tab. CXXXIV, fig. 2; Mob.. Hcke, Fisch. Osts ., p. 114;
Norb.. Handl. Fiskev., Fiskafv., p. 430, fig. 138; Lillj.,
Sv., Norg. Fn., Fisk., vol. Ill, p. 169.

''Tinea Chrysitis (+ var. aurata), Agass., Mem. Soc. Sc. Nat.
Neuch.. tom. I, p. 37; Bonap. (+ T. italica), Iconogr.
Fna Ital., tom. Ill ( Pesci ), tab. 93.

Tinea Linnei , Malm, Gbgs, Boh. Fn., p. 564.

a Zoogr. Ross. Asiat., tom. Ill, p. 299.
b R'egn Anim., ed. 1, tom. II, p. 193.
c Brit. Anim., p. 186.

TENCH.

749

The average length of the Tench in Sweden is
about 2 or 3 dm. The largest specimen Ekstrom ever
saw was 47 cm. long — already an extraordinary size
— but Nilsson had a specimen nearly 55 cm. in length.
According to Blanchard the fish attains at the age
of one year a weight of about 125 gm., at the age
of three years about 1 — 1 1/2 kgm., at the age of six
or seven years 3 — 4 kgm. This last weight is gener-
ally given, on Bloch’s authority, as the maximum
weight of the Tench, and according to the reports sent
in to the Swedish Fisheries Committee of 1881 the
species attains in the District of Jonkoping a weight
of 8 Swedish pounds (3% kgm.), but this statement
needs confirmation". According to Fatio Tench 4 dm.
long weigh about 1 7 3 kgm., and Tench 5 dm. long

27a kgm-

The body is thick, but laterally compressed and
fairly deep, the greatest depth, at the beginning of the
dorsal tin, being more than 7 4 of the length6. The
depth is comparatively greatest, however, in the hind
part of the body, for the Tench is the only one of our
Cyprinoids in which the minimum depth of the tail, at
least in adult specimens, is about 14 %c (13 1/2 — 15 1/2 %)
of the length of the body or 3/s (58 — 65 %) of the length
of the head**. The dorsal profile ascends only slightly,
but in a regular curve. The back is convex through-
out its whole length, flatter behind than in front of
the dorsal fin. The belly, on the other hand, is flatter
in front of than behind the ventral fins, but in the
males is sometimes concave in the median line between
the ventral fins and the anal fin.

The head, the length of which is about 22 — 24 %
of that of the body, is thick and somewhat compressed.
Its breadth is about equal to its vertical depth at the
anterior margin of the orbit. The forehead is broad
and evenly convex, and lies in a line with the snout
and occiput. Its breadth is always somewhat (about
7,0) greater than the length of the blunt snout, which
in adult specimens occupies about 38 % of the length
of the head. The mouth is small, but turned sharply
upward; the lips are thick. The length of the barbel
at the corner of the mouth is usually less than half

the diameter of the eyes. The length of the upper
jaw is about 1/i (24 — 27 %) and that of the lower jaw
about 7s (30 — 34 %) of the length of the head. The
lower jaw is always somewhat shorter than the suture
between the suboperculum and the operculum, which
suture is as a rule equal in length to the snout. The
eyes are small, but comparatively larger as usual in
young specimens: during the growth of the fish from
15 to 40 cm. their longitudinal diameter (slightly greater
than the vertical) varies between about 15 and 13 %
of the length of the head or 40 and 34 % of that of
the snout. They are set about half-way between the
occiput and the tip of the snout, and so high that the
line from the middle of the margin of the upper jaw
to the middle of the caudal fin touches the inferior
margin of the iris. The nasal cavities, which lie twice
as far from the tip of the snout as from the eyes, are
furnished, here as in the preceding genera, each with
two closely adjoining nostrils. The anterior nostril is
the smaller and has a projecting dermal flap at the
hind margin. The gill-openings are fairly large, their
height being about equal to the least depth of the tail.
The opercula are smooth, rounded at the margin, and
furnished with a broad rim, which extends from the
upper angle of the gill -opening to the point where the
branchiostegal membrane is attached to the isthmus, at
a, distance of half the diameter of the orbits from the
branchiostegal membrane of the other side. The three
rays in each branchiostegal membrane are bent, broad,
and strong. The gill-rakers are short and scattered,
numbering about 13 in the outer row on the first
branchial arch, 16 on the outer margin of the pha-
ryngeals. The five, or sometimes only four (more usu-
ally five on one side, four on the other) pharyngeal
teeth (fig. 187) are more or less (the foremost tooth
least) transversely (behind and in front) compressed,
with simple masticatory surfaces, depressed in the trans-
verse median line, and with the upper inner corner
more or less (most in the hindmost tooth) hooked. The
pharyngeal cartilage is reddish brown like horn, of a.
pointed oval shape (triangular with rounded lateral
corners and arcuate anterior margin), and almost hooked

a Yarrell tells us (from Daniel’s Rural Sports') of a Tench 33 in. long and 11 lbs. 9 4 oz. in weight, which had long lived
closely confined among some roots in a pond choked up with mud.

b In adult specimens about 27 — 31 % of the length of the body. In young specimens about 7 cm. long this percentage is about
21, according to Canestrini.

c In young specimens, however, according to KR0YER, about 11 */2 %.

d In exceptional cases, however, these percentages may be found in the Crucian Carp and the White Bream.

750

SCANDINAVIAN FISHES.

at the inferior extremity. It may be easily detached
from its place.

The body is covered with very thin, oblong, and
imbricated scales, which are deeply inserted in the skin
and thus appear externally to be very small. One of
the larger scales above the lateral line in a Tench 4
dm. long is 9 mm. in length and 5 mm. in breadth,
but only the hindmost 2 mm. project out of the follicle.
The nucleus lies far forward in the covered part. A
great number of radiating grooves run from the nucleus
in all directions to the margin of the scale (the most
numerous and densest to the hind margin); and the
tine, numerous, concentric striae encircle the nucleus,
running parallel to the margin of the scale. The scales
and follicles are clothed with a fairly thick, soft, and
slimy epidermis. The lateral line bends down in front,
and then runs straight along the body, about equally
distant from the back and belly. It contains about
100 scales. Above the line there lie about 30 scales,
below it about 20, in an oblique row from the be-
ginning of the dorsal fin.

The height of the dorsal tin is greater than the
length of its base: the former measures in adult spe-
cimens about 18 % (17 — 19 %) and the latter about
13x/2 % (12 — 15 %) of the length of the body. The
distance between this tin and the tip of the snout is
about 48 % (47 — 49 %) of the length of the body, or
about twice as long as the dorsal margin of the ped-
uncle of the tail (between the dorsal and caudal tins).
Its first four rays are simple; but the first ray is al-
ways so short that it does not even project above the
skin; the second ray is also short and in old specimens
difficult of detection, the third about half as long as
the fourth, which is only slightly shorter than the se-
cond or third among the branched rays. These last
two rays are the longest in the tin. Sometimes, how-
ever, even the fourth ray may be branched; and simple
or branched, but narrow rays may lie inserted here
and there between the branched rays, which are other-
wise as a rule 8 in number.

The anal fin is of exactly the same structure as
the dorsal and of essentially the same form; but its
base is still shorter in proportion to its height. The
latter measures about 14 % (13 — 15 Vs %) of the length
of the body, the former about 8 % (7 — 9 %) of the
same. The beginning of the tin lies at a distance from
the tip of the snout of about 63 % (60 — 64 %) of the
length of the body. The vent is usually separated

from the beginning of the anal fin by a distance about
equal to the diameter of the eyes (a little more or less).

The caudal tin is broad and slightly concave at
the end, almost truncate when expanded. The length
of its middle rays, measured from the point where the
scales end, is about 14 % (157s — 13 x/4 %), that of its
longest lateral rays about 16 % (18 — 1 5 x/2 %), of the
length of the body.

The paired tins are broad and obliquely oval. The
most distinct external differences between the sexes find
expression in the dimensions of these tins. In young
Tench and in the males the ventral fins are longer than
the pectoral tins, in adult females shorter. The length
of the pectoral tins is about 1 6 x/2 % (16 — 18 %) of that
of the body: the length of the ventral tins is in the
females less than 17 % (15 — 16 %), in the males more
than 17 % (about 18 %) of the same. It is also a rule
that in the females the ventral fins, when laid back, do
not extend to the vent, while in the males their tips
reach beyond it. In the males too, the second ray in
these tins is very thick and broad. The distance be-
tween the ventral tins and the tip of the snout is about
42 % (39 x/ 2 — 44 %) of the length of the body, their
position being generally farther back in the females.

The coloration of the Tench adapts itself to the
water in which the fish lives. In clear water the whole
body is yellowish green on a golden ground, with tine,
golden dots at the tip of each scale. The top of the
head is darker than the back, the latter in its turn
darker than the sides, which fade below into the whitish
yellow colour of the belly. This last colour also ex-
tends to the lower parts of the head and of the gill-
covers. All the fins are light green. The iris is cop-
pery red. — When the fish has been so long out of the
water that life is extinct, the colour is entirely changed.
The body is now dark green. The back, and in parti-
cular the top of the head, are blackish green. The tine,
golden dots have vanished. The belly is yellow. The
tins are dark purplish red or nearly black. These last
organs are also the first parts of the body to change
colour. — When the Tench lives on a muddy bottom,
it presents the appearance shown in our figure. When
it is found in the forest tarns, where the water is ge-
nerally very dark, the entire upper part of the body is
black, as though dipped in ink, only the belly being
somewhat lighter. A not uncommon variety is the Golden
Tench, of a spotted or plain orange colour. This form
is often kept in fish-ponds both in Germany and England.

TENCH.

751

The internal organs essentially resemble those of
the preceding genus. The liver follows the intestinal
canal backwards in two long, pointed lobes, the right
lobe being the longest and extending' back to the vent.
The intestinal canal is short and thick, only slightly
longer or even shorter than the fish, with only two
coils and without dilatation or special stomach. The
air-bladder is large and, as usual, contracted at the
first third of its length. The anterior, smaller part is
cylindrical with rounded ends, the posterior part pointed
behind. The ovaries and testicles are paired and oc-
cupy the ordinary position.

The Tench is more of a European fish than the
preceding species, but otherwise has essentially the same
geographical range. It occurs throughout the whole of
Europe, northwards in places to the 62nd degree of
latitude. It is also known in Asia Minor, and in the
extreme south of Europe it is common. How far its
range extends in Siberia, is unknown; but Pallas says
that it is common around the Yenisei. In the west of
Europe it does not seem to go so far north as in the east
of the same continent. Thompson" regarded the Tench
as of foreign origin in Great Britain and Ireland. Day
says that it is commoner in the east of England than
in the west. Kroyer had no knowledge of its presence
in the north of Jutland; but according to Feddersen
it has subsequently been found in Lake Ravnstrup, a
little north of the east end of Liim Fjord. In Norway,
according to Collett, the Tench occurs at only two
spots, situated in the coast regions of the extreme
south, namely Kragero and the park at the Nas Iron-
works. To the latter locality it has been transplanted
from Denmark. In the southern tracts of Sweden it is
fairly common: but the northern limit of its range lies
in about 60° N. lat., in Werrnland and Westmanland,
according to the reports sent in to the Fisheries Com-
mittee of 1881, though it is said to have been planted
in the District of Gefleborg. In Finland, according to
Mela, its range extends to 61° 40' N. lat. According
to Reuter it occurs even at Archangel; but according
to Grimm 1 its range to the north is bounded in Russia
by the 62nd degree of latitude. According to Fatio
it ascends in Switzerland to lakes situated as much as
1,600 m. above the level of the sea.

In Sweden the Tench is most commonly found in
small lakes, ponds, and fens, with a muddy and weed)'
bottom. It is worthy of remark that this fish also oc-
curs in the central part of the western island-belt of
the Baltic, where it invariably chooses its haunts, how-
ever, in shallow and weedy inlets in the innermost
part of the archipelago. Sluggish and indolent by na-
ture, it loves quiet and is destitute of the activity dis-
played by the majority of the following genera of this
family. Except during the spawning-season it lives al-
most constantly at the bottom, most often embedded in
mud among the weeds. It is especially prone to this
latter habit in winter, at which time it generally lies
still in a kind of dormancy; but, according to Siebold,
it is sometimes met with in this position even in the
hottest summer. Now and then, though seldom, it may
be seen in summer, when the water is calm, at the
surface. Being very tenacious of life, it may be trans-
ported considerable distances without dying, and is thus
easily planted in ponds. It was one of the fishes earli-
est selected for this purpose.

The spawning-season generally occurs in Sweden
at the beginning of June or somewhat later, according
to the state of the weather. The oldest females spawn
first, the younger ones later in the season. The spawn-
ing takes place in shallow and weedy inlets, ponds, and
small lakes, without any boisterous demonstrations. The
fine, yellowish eggs are deposited on the weeds; they
are generally hatched in a week’s time. The fry grow
rather slowly c, though the rate seems to vary according
to the spawning-place and the state of the water. The
number of the spawning females is usually less than
that of the males. Yarbell estimates the proportion
of the sexes at two males to one female, or not less
than three to two. This disparity in number might
have a detrimental effect on the propagation of the
species, if the fecundity of the Tench did not com-
pensate the scarcity of females. In a female 1 s/4 kgm.
in weight Bloch estimated the number of the ova at
297,000. Harmer, according to Day, found as many
as 383,253 ova in one female.

The food of the Tench consists of mud, worms,
and insects. It is seldom caught in the seine, generally
in fish-traps (Jcatsor, see fig. 204, p. 816, belotv, and

“ Nat. Hist. Irel., vol. IV, p. 136.
b Fish., Hunt. Russ. Wat., p. 14.

c In the month of October Lilljeborg took a young Tench 70 mm. long.

95

Scandinavian Fishes.

752

SCANDINAVIAN FISHES.

ryssjor, see above, p. 33, fig. 7). It bites freely at a
bait of the common earth-worm. The qualities of the
tlesh as an article of food are extolled by some and
decried by others". This depends in most cases upon
the manner of dressing it for table, and a Tench may
lie made into a good dish. The flesh is white, free
from bones, and firm, but is considered indigestible and
therefore unwholesome. The muddy flavour which it
generally possesses, disappears if the fish be scalded in
hot water before cooking, to remove the thick coat of
of slime which covers the body. This slime has been
supposed to deter predatory fishes from attacking the

Tench. But this is not the case: both the Pike and
Salmon eat Tench with avidity, and small Tench are
excellent live-bait, especially in trolling for Pike.

The Tench, which is known in Sweden as Lin-
daren, Sutaren (the Danish Slider), and Skomakaretf ,
was regarded by the popular belief of ancient times as
the physician of the other fishes; and in human medi-
cine it was employed in several ways. It was used as
a remedy for ague and a preventive of malaria, by its
aid jaundice could be removed from the constitution^
and its liver was a cure for toothache.

’(Ekstrom, Smitt.)

Subfamily LEUCISCINtE.

Base of the dorsal Jin about equal in length to that of the anal ( the latter varying between 70 and 130 % of
the former , but less than 16 % of the length of the body). Distance between the dorsal fin and the tip of the
snout more than twice the length of the head. No spinous ray in the dorsal or the anal find. Lateral line ( where
present) situated at the middle of the depth of the tail or lower. Abdominal margin in front of the vent uninter-
ruptedly covered with scales in the median line. Mouth without barbels. Length of the lower jaw more than 47 %
of the base of the anal fin. Pharyngeal cartilage as a rule oblong (oval), rounded or pointed in front. Length

of the intestinal canal as a rule less than that of the body.

As we have mentioned above, we have chosen as
the principal character of the subdivisions of the Cyp-
rinoid family, the correlative size of the dorsal and
anal fins. Within the Scandinavian fauna this distinc-
tion is adequate; but the natural connexion between
this subfamily and the following one produces hybrid
forms, which even in our fauna are exceptions to the
rule — a Bream form (the so-called Abramidopsis Leuck-
artii) with Leuciscine characters and Leuciscine forms
( Bliccopsis ) with Abramidine characters.

The Scandinavian forms belonging to the subfamily
Leuciscince are distributed among the following genera:
A: Length of the base of the anal fin less
than 19 % of the distance between this
fin and the tip of the snout.
a: Distance between the dorsal fin and
the tip of the snout more than 82 %
of that between the anal fin and the
same point. Scales small _ Genus Phoxinus.

b: Distance between the dorsal fin and
the tip of the snout less than 82 %
of that between the anal fin and the
same point Scales of moderate size Genus Leuciscus.

B: Length of the base of the anal fin more
than 19 % of the distance between this
fin and the tip of the snout.
a: Distance between the dorsal fin and
the tip of the snout less than 86 %
of that between the anal fin and the
same point. Lateral line complete.

«: Beginning of the dorsal fin situated
behind the middle of the body.

Length of the lower jaw less than

2/. of that of the head Genus Scardinius.

(3: Beginning of the dorsal fin situated
in front of the middle of the body.

Length of the lower jaw more

than 2/3 of that of the head Genus Aspiits.

b: Distance between the dorsal fin and
the tip of the snout more than 86 % of
that between the anal fin and the
same point. Lateral line incomplete Genus Leucaspius.

a An allusion to the contempt with which the Tench was regarded survives in the well-known lines of Ausonius:

“Quis non et virides vulgi solatia tineas
Norit’’ . . .

“Who knows not the green Tench, the mob's delight?’'
b Sutare and Skomakare both mean shoemaker, a name perhaps derived from the colour of the fish. Tr.
c Icteri magnes , Linn^us.
d In Scandinavian forms.

CARP-FISHES.

753

Genus PHOXINUS.

Scales very small , at least about 80 in a row along the sides of the body and about 30 in a transverse row on
each side of the trunk. Lateral line usually incomplete. Jaws equally projecting. Lobes of the caudal fin blunt
(rounded). Base of the anal fin less than 19 % of the distance between this fin and the tip of the snout.

The genus of the Minnows, which was introduced
into the system by Agassiz", is well defined in the
Scandinavian fauna, not only by its small scales, but
also by the elongated and terete form of the body.
From Siberia, however, Wakpachowski has described
some forms6 more closely approximated by the deep
and laterally compressed form of the body to the other
Leuciscince. In Southern Europe too, the limit between
this genus and the other genera of the subfamily is
difficult to maintain, forms of the genus Leuciscus c oc-
curring there, which have as many as 80 scales in the
lateral line, though, to judge by the descriptions, they
differ from Phoxinus in having at most 26 scales in a
transverse row. In North America the genus Phoxinus ,
as defined by Jordan and Gilbert^, also contains spe-
cies with larger scales. A chromatic character fairly
constant in the Minnow, a more or less prominent,
dark stripe along the sides of the body, and another
fairly characteristic peculiarity, the tumid form of the

snout, reappear in the Leuciscus ( Telestes ) muticellus
of Southern Europe, a form which also composes a re-
markable connecting-link between the genera of this
subfamily, and which has sometimes borne the system-
atic name of the Minnow. Valenciennes c and Gunther '
have therefore found it advisable to unite the genus
Phoxinus with Leuciscus. But it cannot be denied that
the Minnow is the most singular of all our Cyprinince ,
or that it is well deserving of a distinct rank as a
connecting-link between the dark and small-scaled Tench
of the preceding subfamily and the following White-
fishes with their larger scales.

The name of Phoxinus is derived from Aristotle,
who merely remarks, however, that the fish is a fresh-
water form, capable of reproduction at its very birth
and always full of roe. Following Belon and Rondelet,
ichthyologists have assumed that Aristotle here re-
ferred to the Minnow.

a Mem. Soc. Sc. Nat. Neuch., tome I (1835), p. 37.
b Bull. Acad. St. Petersb., tom. XXXI (1887), p. 533.

c Leuciscus rnicrolepis. Cf. Canestrini, Arch. Zool., Anat., Fisiol., vol. IV (1866), p. 109.
d Bull. U. S. Nat. Mus., No. 16, p. 242.
e Cuv., Val., Hist. Nat. Poiss., vol. XVII, p. 363.

/ Cat. Brit. Mus., Fish., vol. VII, p. 207.

754

SCANDINAVIAN FISHES.

THE MINNOW (sw. elritzan).

PHOXINUS APHYA.

Plate XXXIII, fig. 3, c? and $.

Body elongated and terete , its greatest depth being less than 1f5 (17 — 19 V2 %) of its length a, greatest thickness
at least half the greatest depth , and least depth ( least depth of the tail) less than half the greatest depth or than
V3 of the length of the tail from a line with the end of the base of the anal fin to ■ the middle of the base of
the caudal fin ■ Length of the head about 21 — 23 V2 % of that of the body. Pharyngeal teeth pointed, hooked at
the tip, with the masticatory surface sharpened , but smooth or even hardly distinguishable, and set in a double or

single roiv: 2, 5(4) — 4(5), 2 or 5 — 4.

a b

Fig. 188. Pharyngeal bones ( a and b, as in the preceding figure) and covering of the pharyngeal process (c) in Phoxinus aphyci.

Magn. 3 diam.

R. br. 3; D.

(2)3

A.

(2)3

V.

(1)2

P.

7(8)’ “ (6)7’ " (6)7(8)’

C. x 4- 1 4- 1 7 4- 1 4- xi L. lat. (85)90 — 110; P. tr.
Vert. 38 — 40.

1

13 — 16 6’
15—20
14 — 16’

>Syn. Oi cpofivoi, Aristot., De Anim., lib. VI, cap. XIII et XIV.

Phoxinus , Veron, Belon, Nat., Div. Poiss., p. 324. Pisci-
culus varius, Rond., De Pise., part. II, p. 205. Phoxinus
Icevis , Schonev., Ichthyol. Slesv. Hols. p. 57. Cyprinus tri-
dactylus varius oblongus teretiusculus, pinna ani ossiculorum
octo, Art., Ichthyol., Syn. Pise., p. 12 4- Cyprinus biunci'
alis, iridibus rubris, pinna ani ossiculorum novem, Id., ibid,
(excl. syn. Schonev.) et Descr. Spec. Pise., p. 30. Lin., Fn.
Suec., ed. I, p. 125, No. 331; Id., Alkufwa , It. Westroy.,
p. 232.

Cyprinus Phoxinus, Lin., Syst. Nat., ed. X, tom. I, p. 322;
Bl. Fisch. Deutschl., part. I, p. 60, tab. VIII, fig. 5; Retz..
Fn. Suec. Lin., p. 356; Pall. Zoogr. Ross. Asiat., tom.
Ill, p. 330; Flmng ( Leuciscus ), Brit. Anim., p. 188; Ekstr.,
{Cyprinus), Vet.-Akad. Handl. 1830, p. 166; Nilss., Prodr.
Ichthyol. Scand., p. 29; Id. ( Leuciscus ), Skand. Fn., Fisk.,
p. 319; Sundev. {Cyprinus), Stkhlms L. Hush. Sails. Handl.,
H. 6 (1855), pp. 82 et 175; Thomps. {Leuciscus), Nat.
Hist. Irel., vol. IV, p. 138; Lindstr., Goth L. Hush. Sallsk.
Arsber. 1866, p. 18 (sep.); Gthr, 1. c. ; Olsson, Of vers.
Vet.-Akad. Forh. 1876, No. 3, p. 131; 1882, No. 10, p.

48; Day, Fish. Gt. Brit., Irel., vol. II, p. 185, tab. CXXXIV
fig. 1; Mob., Hcke, Fisch. Osts., p. 113.

Cyprinus Aphya, Lin., Syst., 1. c., p. 323; Bl., 1. c., part. Ill,
p. 143, tab. XCVII, fig. 2, quae figura tamen pessima potius
forsan ad Leuc. grislaginem referenda; Retz., 1. c.; Nilss.,
Prodr., 1. c. ; Kr. {Phoxinus), Damn. Fisk., vol. Ill, p. 524;
Coll., Forh. Vid. Selsk. Christ. 1874, Tilla?gsh., p. 183;
1879, No. 1, p. 96; N. Mag. Naturv. Christ., Bd. 29 (1884),
p. Ill; Malm, Gbgs, Boh. Fn., p. 564; Fedders., Naturb.
Tidskr. Kbhvn, ser. 3, vol. XII, p. 91; Mela, Vert. Fenn.,
p. 321, tab. X; Lillj., Sv., Norg. Fisk., vol. Ill, p. 179.
Cyprinus rivularis, Pall., Russ. Reis., part. II, p. 717.
Phoxinus Icevis , Agass., 1. c. ; Hckl, Kn., Susswasserf. Oestr.
Mon., p. 210; Dybowski, Cypr. Livl ., p. 105; Sieb., Siiss-
wasserf. Mitteleur., p. 222; Mgrn, Finl. Fiskfn. (disp.
Helsingf. 1863), p. 48; Bncke, Fisch., Fischer ., Fischz. 0.,
IF. Preuss., p. 141; Mor., Hist. Nat. Poiss., tom. ID, p.
392; Fatio, Fn. Vert. Suisse, vol. IV, p. 638.

The Minnow is the smallest Scandinavian Cyprinoid,
with the exception of the Owsianka (Leucaspius de-
lineal us), and is generally no larger than the latter.
Its ordinary length in Sweden is about 6- — 9 cm. Our
largest specimen, from Lake Korn in Northern Bohus-
lan, measures 105 mm. to the end of the caudal lobes

“ Except in gravid females.

b Fewest in the males, in which, according to Fatio, the number is further reduced during the spawning-season.

MINNOW.

755

or 99 mm. to the tips of the middle caudal rays. From
Ireland, however, though the species is supposed to
have been planted there, Thompson mentions a speci-
men 6 in. (152 mm.) in length. The females attain
a greater size than the males. In shoals of spawning
Minnows Heckel and Kner found the length of the
females to vary between 92 and 105 mm., that of the
males between 52 and 79 (exceptionally 92) mm. The
largest specimens in their possession were 131 mm.
long and came from Hungary.

The body is elongated and of fairly uniform thick-
ness, the anterior part of the trunk showing so little
compression that the greatest thickness is at least aj3
of the depth. Posteriorly the lines of the body con-
verge, as usual, even from the sides, but the tail is
still comparatively terete. The depth of the head at
the occiput is slightly greater than its breadth. The
back is broad and depressed in the median line into
a groove, which may be exchanged in front of the
dorsal tin for a low carina. The belly is even and
terete, or, at the isthmus, flat. The dorsal profile shows
a more or less distinct break at the occiput, but is
otherwise regular, like the ventral profile, which differs
from that of the other Whitefishes in being free from
any perceptible break, even at the vent or along the
anal fin.

The most characteristic point in the form of the
head consists of a swelling (thickening of the ethmoid
bone) on the snout between the nostrils. The occiput
and forehead are slightly convex or (especially the
latter) nearly flat; the gill-covers and cheeks almost
perpendicular. The eyes are middle-sized and set so
high that their superior margin lies almost in the same
plane as the forehead. Their longitudinal diameter
measures in young specimens (x/2 dm. long) about 30
%, in average- sized specimens about 28 %>, and in spe-
cimens 1 dm. long about 23 %, of the length of the
head; the vertical diameter is 9/io — Vs °f the longi-
tudinal. They lie almost entirely in the anterior half
of the head, the postorbital length of the latter being-
only slightly less than the length of the snout and the
eye together. The breadth of the interorbital space is
always greater than the diameter of the eyes and va-
ries between 40 and 30 % of the length of the head.
The horizontal profile of the snout is obtusely (slightly)
rounded, and its tip generally projects a little beyond
that of the lower jaw. The cleft of the mouth, with
its fleshy lips, is small and turned slightly upwards.

The length of the upper jaw from the middle of the
tip of the snout is, as a rule, equal to that of the
snout, or about 30 % of that of the head. The length
of the lower jaw is about the same as the breadth of
the interorbital space, or on an average 1/.i of the length
of the head. The nostrils are round and rather large,
but set close together, the posterior nostril being usu-
ally covered to a great extent by the dermal flap that
rises from the narrow bridge between them. They lie
much nearer to the eyes than to the tip of the snout.
The gill-openings are comparatively large, extending
above along about half the upper margin of the gill-
cover, and separated at the isthmus by a distance some-
times scarcely half the diameter of the eyes. The
branchiostegal rays as well as the rim outside them are
broad. The gill-rakers are small; in the outer row on
the first branchial arch they are about 8 in number,
scattered, verrucose, and hardly distinguishable. The
pharyngeal teeth show considerable variation in number.
In most cases they are set in two rows, the large row
on the inside of the lower arm of the bones generally
containing 5 teeth on the left side and 4 on the rig-ht,
and the smaller row, outside the former one, consist-
ing of 2 small, almost cylindrical teeth. Sometimes,
however, not a trace of this smaller row can be found.
We have never found the pharyngeal cartilage present:
according to Fatio it is of an obtuse heart-shape, with
the posterior part attached like a knob to the anterior.

Our figure (fig. 188, c) shows, on the other hand,
how the mucous membrane of the pharynx covers the
downward process of the occipital bone at the spot
where the pharyngeal cartilage is otherwise developed.
This figure may also assist in the explanation of the
form possessed by the pharyngeal cartilage in the ma-
jority of the following Leuciscines, at least in the larger
ones. The surface of this cartilage is heart-shaped.
On each side of it the roof of the pharynx shows deep
hollows, converging in a forward and upward direction
(downward in the figure, which is drawn from a spe-
cimen laid on its back, with the head turned towards
the artist). But at the middle of the base of this
heart-shape the mucous membrane is continued (up-
ward in the figure) in a rounded (convex) coat over
the lower (posterior) part of the said occipital process,
until it leaves this process and passes into the upper
wall of the oesophagus. This convex part forms the
foundation of the callosity which, especially in the
larger Whitefishes (see, for example, the Chub, fig.

756

SCANDINAVIAN FISHES.

191, c; p. 769 below), composes a continuation of the
pharyngeal cartilage in an inverted canaliculate form,
more or less sharply detined and more or less raised
above the remaining surface of the cartilage.

All the fins are of a somewhat rounded shape, re-
minding us in some degree of the Tench. The dorsal
tin begins at a distance from the tip of the snout which
on an average measures very nearly half the length of
the body, but varies between 48 and 52 % thereof.
Its base measures on an average Vio °f the length of
the body, varying between 9 and 11 % of the same.
Its height (the length of the longest, the first branched
ray) is always greater than the length of its base and
measures in the females about 13 — 15 %, in the males
nearly as much as 17 %>, of the length of the body.
As a rule the first ray is imperceptible above the skin.
The same remarks apply to the anal fin, which is of
the same obliquely quadrilateral form, with more or
less distinctly rounded corners, and as a rule of the
same size. The distance between this fin and the tip
of the snout is greatest in the females and varies be-
tween about 55 and 61 % of the length of the body.
The caudal fin is not very deeply forked. Its middle
rays occupy in average-sized specimens about 9 — 11 %
of the length of the body and measure more than half
(53 — 64 %) of the length of its longest rays.

In this species too, the paired fins afford the most
trustworthy distinctions of form between the sexes.
Both pairs are rounded and obliquely oval. The pec-
toral fins are set, as in the Gudgeon, so low that when
expanded they lie almost in the same plane as the
ventrals. In both sexes the length of the pectoral fins
is as a rule about 13 — 15 % of the length of the body,
but in the males it may rise during the spawning-
season to 18 % thereof. In the males the upper (an-
terior) rays (5 — 7) are thick and have more markedly
distinguished articulations than in the females. The
pectoral fins of the Minnow thus supply us with si-
milar sexual characters to those afforded by the vent-
ral fins of the Tench. Here, on the other hand, the
ventral fins of the two sexes differ both in their po-
sition — somewhat further back in the females — and
their length, which in the males is more, in the fe-
males less than 13 % of the length of the body. The
distance between these fins and the tip of the snout
measures during youth and in the males about 40 — 42
% , in the females 42 — 46 % of the length of the body,
and the distance between them and the anterior angle

of the pectoral fins (the preabdominal length) during
youth and in the males 19 — 22 %, in the females 22 — 26
% of the same length. In consequence hereof the tips
of the pectoral fins, when laid back, may sometimes
extend in the males nearly to the bases of the ventral
fins, while the tips of the latter fins in the same posi-
tion reach in the females scarcely to the vent, in the
males beyond this point or even to the beginning of
the anal fin. The vent, which protrudes considerably
and is fairly wide at its junction with the urogenital
aperture, lies perceptibly, though not far, in front of
the beginning of the anal fin. Here, as in the Cobitoids
(see above), we find no special scaly appendage at the
outer angle of the ventral fin.

The covering of scales shows quite a considerable
external resemblance to that of the Tench; but the
scales are of exactly opposite form, deeper than long,
of a more or less broad elliptical shape, with the longi-
tudinal axis turned in the transverse direction of the
body. Instead of being inserted in the skin for the
greater part of their length, as in the Tench, they are
chiefly free and hardly imbricated in the longitudinal
direction of the body, though in a transverse row on
the back each scale may slightly overlap the one below
it. The texture is rather coarse, but densely striated
with concentric ridges, which are interrupted through-
out the circumference of the scale by radiating grooves
extending to a greater or less distance inward from the
margin. The striae are thus divided into patches of
oblong triangular shape, crossed by transverse streaks
and with their apex directed towards the central nucleus
of the scale. The belly is naked in front and at the
isthmus, but is furnished on each side in front of the
pectoral fin with a triangular patch of about 8 rows
of scales. The lateral line pierces its scales throughout
the greater part of their breadth (in the longitudinal
direction of the body), but is very often partially or
entirely wanting on the sides of the tail. It forms a
downward curve from the temporal region, so sharp
that it reaches the middle of the sides in front of the
perpendicular from the tip of the pectoral fin, when
laid back, and runs back from this point in a straight
line to the middle of the base of the caudal fin.

The coloration in conjunction with the well-pro-
portioned form of the body renders the Minnow one of
our most handsome fishes. The colours show considerable
variation, however, according to the locality, the season,
the sex and the mood of the fish. No long description

MINNOWS.

757

can so clearly delineate the coloration of the Minnow
as v. Wright’s two figures: a female and a male in
the spawning-dress. The back is olive brown or nearly
blackish, purer green between the dark transverse bands
that run down towards the sides. The middle of the
sides generally shows a pattern of equally dark shade,
a, row of blackish brown spots, partly or completely
coalescing into a stripe, which runs from the snout,
interrupted by the eyes, to the lower part of the caudal
fin, where it ends in an obliquely set spot, as a rule
the most constant and persistent trace of this stripe,
even in specimens preserved in spirits. The ventral
side is light, milky white or silvery. Very often the
colour of the sides is dashed with gold, especially on
the gill-cover and above the dark stripe; and the vent-
ral side assumes a reddish tint, punctated more or less
densely with blackish brown dots, the last distribution
of colour being most prominent on the forepart of the
body in the males. All the fins are transparent, shad-
ing into yellow, gray, or green. In the spawning-dress
all the colours are brighter, most so, as usual, in the |
males: the sides become emerald green, the belly red-
dish, the golden lustre comes forth on the suboper-
culum, and on the upper part of the operculum, as far
as it is free from the side of the body and forms the
upper margin of the gill-opening, there appears a lust-
rous, white spot, especially conspicuous in the males
on the black ground composed of the occiput, the pre-
operculum, a part of the operculum, and the remainder
of the branch iostegal membrane. In the males ready
to spawn this lustrous white also extends to the bases
of the paired fins and the inner anterior corner of the
anal fin. The corners of the mouth are carmine red.
The iris glitters with a lustre of silver and gold. In
light environments the fish has a lighter dress, and the
body grows somewhat transparent. The fish is also
lighter by day than at night, has a richer dress when
well fed than 'when starved, and changes colour rapidly
enough under the influence of the passions. “The
coloration varies considerably in different individuals,”
writes Ekstrom; “it also changes speedily on the death
of the fish. To observe the numerous and bright hues
with which this fish is adorned, it should be seen in
the water or a,t the moment of its capture. Though
kept alive in a vessel of water, it changes colour soon

enough.” In England a gold-coloured variety of the
Minnow has been found".

The Minnow has many Swedish names: Elritza
(Germ. Elritze), in Halland and Bohuslan Ailing or
Allepytta (Dan. Elbut or Elbutte ) and Alkula, in Dais-
land Alknfva , in Oster Gothland Alkutta, in Wester
Gothland Hundgadda and Gli, in Westerbotten Glirr ,
in Dalecarlia Qvidd and Iggling (Linnaeus), in Jemtland
Blindsill (Olsson), in Lapland Solsensodg (Linnaeus),
among the islands of the Baltic Hamntorsk in the Di-
strict of Stockholm (Sundevall) and Lortbuk in Soder-
manland (Ekstrom), and in Gothland Laxbadd (Lind-
strom). Artedi also gives the names of Mudd and
Skitspigg , Linnaeus that of Budd (Germ. Butt). This
multitude of names is alone sufficient to prove the com-
mon occurrence of the Minnow throughout Sweden and
in the island-belts of the Baltic and the Gulf of Both-
nia. The species is also common in Norway both to
the extreme north (south to about 69° N. lat.) and the
extreme south (north to about 63° N. lat.), but not in
the intermediate districts (Collett, cf. the occurrence
of the Perch in Norway, see above, p. 28, note d). Its
geographical range embraces the whole of Europe —
with the exception, as far as is known at present, of
the Iberian Peninsula — and Northern Asia, east to the
Amur. In Switzerland, according to Fatio, it ascends
the cold Alpine streams and lakes to a height of nearly
2,500 metres above the level of the sea. In Norway, ac-
cording to Collett, it lives high up in the birch region
of the fells, at an altitude of about 900 metres.

The appearance6 and habits of the Minnow have
been compared, not without reason, to those of Salmon-
fry in the stage termed parrs. They are also often
found in company, the Minnow being generally an in-
habitant of .clear streams and brooks with sandy or
gravelly bottom. But the Minnow lives, as we have
mentioned, not only in running fresh water and in
lakes, but also in the sea, where it often appears in
shoals off the piers among the island-belt. Ekstrom
describes its habits in these latter haunts as follows:
“It lives off rocky promontories and stony shores in
deep water, especially where there is a current, associ-
ates exclusively with its own kin, and seems to avoid
places frequented by other species. Where it occurs,
it is always found in large numbers. During the greater

a Manlky, see Day, 1. c.

* “Facies Truttse sen Salmonis”.

Lin., Fn. Suec.

758

SCANDINAVIAN FISHES.

part of the day it lies still, almost motionless, at the
bottom, where it seeks its food. Now and then, how-
ever, it rises to the surface, quick of movement and
voracious for its size. With greedy eagerness it catches
the insects that fall into the water. It dies almost in-
stantaneously when taken out of its native element.”
In fresh water it is less sensitive to the influence of
the air and is sometimes seized with the same wan-
dering spirit as the Salmon, leaping over obstacles with
vigour relatively the same and no less eagerness. It
sometimes forces its way along the tiniest, half-dry
brooks and pools, if only it feels running water — and
may subsequently be found in puddles, where it is
scarcely possible to explain its presence. In fresh water
it is also more sociable in its relations to other fishes:
in the Nissa River (Halland) I have often caught Min-
nows in company with Salmon-fry.

Its food is chiefly composed of insects, small crus-
taceans and mollusks, and worms. In the island-belt
it often keeps watch at the landing-places, where fish
is gutted and rinsed, and seizes small fragments of
the offal. Fish-roe and small fry, even of its own
species, also form a part of its diet. It does not dis-
dain a bait of wheat dough, though it bites less rea-
dily at this; and in an aquarium it may be fed on
bread-crumbs. Its voracity entices it forward at the
least cause and gives it an inquisitiveness of which
the fisherman may take advantage. Fatio tells us
that he caught quantities of Minnows in a landing-
net, by holding it still in a stream. As soon as the
leader and the van of the shoal had entered to see
what they might discover in the net, the others fol-
lowed without hesitation.

The spawning-season seems to vary according to
the early or late arrival of the summer warmth. In
the tracts bordering on the Rhine the Minnow is taken
in large numbers, together with a multitude of other
small fishes (Salmon-fry among others), all known by
the common name of Rumpdien , during the months of
May and June, at which time the Minnow ascends the
brooks to spawn on a, stony or gravelly bottom. Ek-

strom states that in the island-belt of Sodermanland
it spawns at the end of June or beginning of July.
According to Fatio the spawning may begin in the
Swiss valleys at the middle of April, but in the higher
Alpine regions is sometimes delayed till August. The
eggs are not very numerous, at most about 1 ,000 in
the same female, but comparatively large, being 1 — 1 x/4
mm. in diameter. They are hatched, sooner or later
according to the temperature of the water, in from 6"
to 15 h days. “It is a mistake,” writes Sauvadon0, “to
suppose that the eggs of the Minnow are strewn about
and carried away by the current among the gravel and
stones, as soon as they have been deposited. During
the spawning the fish are so numerous and packed so
close together that their bodies are enough to neutralise
the gentle current that otherwise passes over the
spawning-place. Furthermore, these small, glutinous
ova adhere to each other and fill all the interstices be-
tween the stones, on which they may often be found
in layers 1 — 5 cm. thick and 5 — 20 cm. long. All
these eggs stick fast together and form a layer firm
enough to withstand a current ten times stronger than
that Avhich runs at the spawning-place. On the 30th
of May I have myself gathered at least 3 kgm. of
Minnow-eggs.” In the first week of August Yarrell
found young Minnows three-quarters of an inch (19
mm.) long. The Minnow reaches maturity, according
to Fatio, at a length of 35 — 40 mm.

At the approach of autumn the Minnow retires to
deep water, there to pass the winter in company with
other Cyprinoids. As food the Minnow is not to be
despised, especially if the gall-bladder, which otherwise
gives it a bitter taste, be removed. In Sweden, how-
ever, it is seldom eaten; here and there in the country
it is fried and made into the so-called fish-cake. Fur-
ther south, where it is still more plentiful, it is a more
appreciated dish. In England, according to Day, it is
eaten freely, being prepared for table like Whitebait'7.
In Sweden it is most important as bait for Salmon
and Perch; but as it is not very tenacious of life, it
must not be left long on the hook.

a According to Davy, see Day, 1. c.

6 According to Fatio.

c Bull. Soc. Zool. D’Acclimat. Paris, serie 2, tome IV (1867), p. 721.

d Buckland says that on the 16th of September, 1394 Bishop William of Wykeham, the founder of Winchester College, gave a
dinner to the King and Queen and 210 other guests, the menu including a number of fish courses, among others 7 gallons of Minnows.

CARP-FISHES.

759

Genus LEUCISCUS.

Scales as a rale middle-sized", numbering in all the Scandinavian species about 40 — 60 in the lateral line, which
is complete. Jaws projecting equally far, or nearly so. Lobes of the caudal fin pointed. Base of the anal fin
less than 19 % of the distance between this fin and the tip of the snout. Distance between the dorsal fin and
the tip of the snout less than 86 % of that between the anal fin and the same point.

This genus was already known by its modern
name* 6, which means Wliitefish, in the time of Galen
(the second century A. D.). At many places, both in
Sweden and elsewhere, Whitefish is a trivial name
common to the Cyprinoids (especially Leuciscus and
Alburnus ) and Coregonoids. Belon referred the name
of Leuciscus to the Dace, Rondelet both to the Roach
and the Dace. Klein, as we have mentioned above,
gave the genus a definition which corresponded in es-
sential respects to that of the modern subfamily Leu-
ciscince. Linnaeus did not recognise this genus — he
employed Leuciscus as one of his specific names for
the Dace — but CuvieiT adopted it as a subgenus; and
it was subsequently raised by Fleming'7 to the rank of
an independent genus. The number of the species is
so great, about a hundred being known, that attempts
have with reason been made to subdivide the genus.
But the ground on which most authors, following Hec-
kel’s example, have proposed to base this subdivision
— the number and arrangement of the pharyngeal teeth,
their similarity or dissimilarity on the two pharyngeals

and their distribution in one or two rows — has proved
untenable, as we have already been enabled to conclude
from the Minnow, where the smaller row may some-
times disappear without leaving a trace of its presence.
The Scandinavian fauna contains only four recognised
species, which may be distinguished as follows:

A: Anal fin concave.

a: Least depth of the tail less than
43 % of the length of the peduncle
of the tail between the perpendi-
cular from the end of the base of
the anal fin and the middle point
in the line of demarcation between

the scales and the caudal fin Leuciscus grislagine.

b: Least depth of the tail more than
43 % of the length of the peduncle
of the tail measured as above.
a a : Base of the dorsal tin less than

\ 4 of the distance between this
fin and the tip of the snout... Leuciscus idus.
bb: Base of the dorsal fin more than
l/4 of the distance between this
fin and the tip of the snout... Leuciscus rutilus.

B: Anal fin convex... Leuciscus cephalus.

a As for the exceptions, see the observations on the preceding genus.

6 From the Greek hevx.6g, white.

c Regne Animal , ed. I, tom. II, p. 194.
d Brit. Anim., p. 187.

Scandinavian Fishes.

96

SCANDINAVIAN FISHES.

7 60

THE DACE ( sw. stammen).

LEUCISCUS GRISLAGINE.

Plate XXXII, fig. 2.

Scales in the lateral line about 52 ( 50“ — 54). Branched rays in the dorsal fin 7. Least depth of the tail less
than the length of the base of the dorsal fin, or than 43 (41) % of the length of the peduncle of the tail between
the perpendicular from the end of the anal fin and. the middle point in the termination of the scales at the base
of the caudal fin. Outer ( lower posterior ) margin of the anal fin concave. Pharyngeal teeth curved at the tip
or straight , with narrow and smoth or hardly perceptible masticatory surface: 2(3), 5 — 5, 2(3).

Fig. 189. Pharyngeal bones and pharyngeal cartilage of Leuciscus grislagine, 3 times the natural size. a, b , and c as in fig. 178.

3 3 1 2

R. hr. 3; D. — ; A. .P ; V. — ; C.x + 1 + 17

1 8b 17 — 18 8

g

+ 14- x; L. lat. 50rt — 54; L. tr. — 1 (supra pinn. ventr.) ;

4

Vert. 41 — 44 c.

Syn. Leuciscus, Belon, Nat., Din. Poiss., p. 313. Leucisci 2:da
Spec., Rondel., Pise., part. II, p. 192. Cyprinus novem
digitorum, rutilo longior et angustior, pinna ani radiorum
decern, Art., IchthyoL, Syn. Pise., p. 9 (Gallis Vandoise ).
Cyprinus oblongus, figura rutili, pinna ani ossiculorum decern
Art., ibid., Gen. Pise., p. 5; Syn. Pise., p. 5; Descr. Spec.,
p. 12 (Suecis Stcim). Cyprinus pedalis gracilis oblongus
crassiusculus, dorso crasso, pinna ani ossiculorum novem, Art.,
ibid., Syn., p. 10 (Tigurinis Hasel). Lin., Fn. Suec., ed. I,
p. 123 (Angermannis Stum ).

Cyprinus Grislagine, Lin., Syst. Nat., ed. X, tom. I, p. 323;
Retz., Fn. Suec. Lin., p. 357 ; Ekstr.(?), Vet.-Akad. Handl.
1830, p. 157, tab. IV; Nilss., Prodr. IchthyoL Scand., p. 27;
Agass. ( Leuciscus grislagine + L. argenteus + L. rostratus
. + L. rodens + L. majalis ), Mem. Soc. Sc. Nat. Neuch.,
tom. I (1836), p. 38; Ekstr., v. Wright {Cyprinus), Skand.
Fisk., ed. I, p. 69, tab. 14; Kr. ( Leuciscus ), Danm. Fislce,
vol. Ill, p. 472; Sundev. (Cyprinus), Stockh. L. Hush.

Handl., H. VI, 1855, p. 81; Nilss. (Leuciscus), Skand
Fn., Fisk., p. 303; Widegr. (Cyprinus), Fislcfn., Fisker.
Norrb. Tj., Landtbr. Akad. Handl. 1861, p. 12 (sep.); Coll.
( Leuciscus ), Forh. Vid. Selsk. Chrnia 1874, Tillsegsh., p. 181;
Malm, Gbgs, Boh. Fn., p. 562; Mela, Vert. Fenn., p. 326,
tab. X; Id. et Sundm., Finl. Fisk., tab. XXVI; Lillj., Sv.,
Norg. Fisk., vol. Ill, p. 198.

Cyprinus Leuciscus, Lin., Syst. Nat., 1. c. ; Bl., Fisch. Deutschl.,
part. Ill, p. 141, tab. XCVII, fig. 1; Pall., Zoogr. Ross.
Asiat., tom. Ill, p. 318; Hckl, Kn. (Squalius), Sussivasserf.
Oestr. Mon., p. 191 (+ Squ. lepusculus, p. 186, + S.

chalybceus, p. 188, + S. rodens (ex Agass.), p. 189, + S.
rostratus (ex Agass.), p. 192); Dyb., Cypr. Livl., p. 126;
Sieb., Sussivasserf. Mitteleur., p. 203; Mgrn, Finl. Fislcfn.,
p. 47; Blanch, Poiss. d. eaux douces Fr., p. 401 (+ Squ.
bearnensis, p. 400, + S. burdigalensis (ex. Val.), p. 403);
Feddersen, Naturh. Tidskr. Kbhvn, ser. 3, vol. XII, p. 88;
Bncke, Fisch., Fischer., Fiscliz. O., W. Preuss., p. 139;
Mor., Hist. Nat. Poiss. Fr., tom. Ill, p. 425; Fatio, Fn.
Vert. Suisse, vol. IV, p. 582.

Cyprinus Dobula, Lin., Syst. Nat., 1. c.; Bl. 1. c., part. I,
p. 42, tab. V; Yarr. (Leuciscus) Brit. Fish., ed. 1, vol. I,
p. 346 (+ Leuc. vulgaris, p. 353, + Leuc. Lancastriensis
(ex Shaw), p. 355).

“ Sometimes 47, according to Sjebold.

6 9

99 ° 1 99 99 99

c According to Fatio; 40 — 45 according to Jeitteles.

DACE.

761

Leuciscus vulgaris , Flmng, Brit. Anim ., p. 187; Cuv., Val.,
Hist. Nat. Poiss., vol. XVII, p. 202 (+ Leuc. rostratus
(ex. Agass.) + L. rodens (ex. Agass.) + L. lancastriensis,
(ex. Sijaw), p. 216 + ]j. burdigalensis , p. 218, nec L. gris-
lagine, p. 220); Gthr, Cat. Brit. Mus., Fish., vol. VII, p.
226; Day, Fish. Gt. Brit.. Irel., vol. II, p. 180, tab.
CXXXIII, tig. 1; Mob., IIcke, Fiscli. Osts., p. 109.

The Dace is one of the smaller species of the ge-
nus. Its ordinary length is between 15 and 20 cm.,
and in Sweden probably does not exceed 25 cm.
Further south it seems to attain a somewhat greater
size; Fatio found in the Rhine a gravid female of this
species, 275 mm. long and 300 grammes in weight.
Day states that in the Thames the Dace attains a length
of about 3 dm.

Of all the Scandinavian species within the Leucis-
cine subfamily the Dace is the most elongated in pro-
portion to the depth of the body, and is easily recog-
nised by this character. The body is deepest a little
in front of the dorsal fin, and the greatest depth mea-
sures in adult specimens, 18 — 21 cm. long, about 21
— 24 % of the length of the body. In younger speci-
mens the greatest depth is even relatively less, as else-
where in the family. The least depth of the body (of
the peduncle of the tail) measures in adult specimens
about 8 V2 — 7 Vs % of its length. The back is terete,
though less so just in front of the dorsal tin, and the
sides are moderately compressed. The back rises in a
gradual curve from the head to a. point a little in front
of the dorsal tin, though it is straighter than in the
Scandinavian congeners of the Dace, and the curve does
not increase in sharpness at the occiput. From this
point the dorsal profile slopes gently, almost in a straight
line, to the base of the caudal fin. The ventral profile
forms a similar or even more gradual curve from the
chin to the vent, which protrudes slightly, and then
runs straight to the caudal fin.

The head is of moderate size (in adult specimens
about 21 — 19 % of the length of the body), with broad
and rather flat forehead, sloping 'with a slight convexity
towards the broad and very obtuse snout, which pro-
jects a little in front of the jaws. The mouth is small
and turns slightly upwards, as in the rest of th e -Leu-
ciscince , the upper jaw is slightly more prominent than
the lower, and the corners of the mouth extend a little
behind the perpendicular from the posterior nostril.

The length of the upper jaw from the middle of the
tip of the snout measures in adult specimens rather
more than 1/i (about 25 V2 — 28 %) of that of the head.
The length of the lower jaw is equal to or somewhat
greater than that of the snout, or about V3 (32 — 35 %)
of that of the head. The eyes are fairly large, their
longitudinal diameter being only slightly less than 1 4
(about 23 — 24 V2 %) of the length of the head. They
are generally set just in front of the middle of the
head" and so high that the line drawn from the middle
of the caudal fin to the margin of the upper jaw
touches the inferior margin of the eye. The least
breadth of the interorbital space is about 1/3 (32 x/2 — ^
34 %, exceptionally 37 %) of the length of the head, or
1V2 — lVs times the longitudinal diameter of the eyes.
The nostrils are rather large, and lie in a deep cavity,
nearer to the eye than to the tip of the snout. The
gill-rakers are small and scattered, but pointed, num-
bering 8 in the outer row on the first branchial arch
and 7 on the pharyngeal bones. Each pharyngeal is
armed as a rule with 7 teeth (fig. 189), set in two
rows. The inner roAv contains 5 long and fairly straight
teeth, curved up Avar ds at the tip, but usually not much
hooked6, and Avithout incisions or spines on the indi-
stinct masticatory surface. The teeth in the outer roAv
are generally 2, sometimes 3, in number, short, straight,
and conical. The pharyngeal cartilage is soft, thin at
the edges, heart-shaped, and deciduous. The gill-
openings do not extend at all above the upper margin
of the gill-cover, and beloAv they coalesce Avith the
isthmus at a distance from each other of about half
the diameter of the eyes.

The dorsal fin, the height of Avhich is greater than
its length at the base, lias an obliquely truncate mar-
gin Avith somewhat rounded corners. Behind it is half
as high as in front, a rule Avhich applies to most of the
Leucisenue. It begins at a distance from the tip of the
snout Avhich in adult specimens measures on an average
half the length of the body, varying, hoAvever, between
46 V2 and 54 % thereof. Its base occupies 710 (varying
in our specimens between 9 1/2 and 10V2 %) of the length
of the body; and its height (the length of the longest
ray), Avhich is as a rule equal to the length of the
pectoral fins, varies in our specimens between 15 and
1 6 1/3 % of the length of the body. The first three rays

a

depending

b

The postorbital length of the head varies in onr specimens between 4 8 1 / 2 and 54 % of its entire length, the
on the breadth of the opercular rim.

In specimens from Lake VVener, however, we find the hooked tips well developed in the two hindmost teeth

variations essentially

of this row.

762

SCANDINAVIAN FISHES.

are simple, the first being hardly perceptible externally a,
the third the longest ray in the whole tin, or equal in
length to the fourth. The last seven rays are repeatedly
branched at the tip, and the last ray is cloven almost
to the base.

The anal tin is of nearly the same shape as the
dorsal, and of exactly the same structure, except that
it has an additional branched ray. Its corners, espe-
cially the posterior, are more pointed, and the margin
is slightly concave. The tin begins at a distance from
the tip of the snout that measures on an average some-
what less than 2/a (varying in our specimens between
63 and 66 V2 %) of the length of the body, and its base
occupies on an average about 1j11 (varying in our spe-
cimens between 8‘9 and 9'8 %) of the same length. Its
height is on an average about Vs (varying in our spe-
cimens between 12 and 13 %) of the length of the body.

The pectoral tins are obliquely pointed, and con-
sist of 1 simple and 17 or 18 branched rays, of which
the last (lowest) two or three are small and difficult to
distinguish. The ventral tins, on the other hand, are
broader and more rounded, and are furnished on their
front (outer) side, as usual in this family, with two
simple rays, so closely united that the first is not al-
ways distinguishable. Their length measures in young
specimens about 14 %, in old about 121/, * of that of
the body. The distance between the foremost (outer-
most) point in their insertions and the corresponding
point in the insertions of the pectoral fins is on an
average about 1/i (varying in our specimens between
23V2 and 28 %) of the length of the body. The di-
stance from the former point to the tip of the snout
varies between about 44 and 48 % of the length of the
body, and to the beginning of the anal fin between
about 20 and 2 1 1 2 % of the same length. The bases
of these fins are furnished as usual with a pointed,
lanceolate appendage.

The caudal fin is forked, with lobes of equal length.
Its middle rays occupy about 8— 7 % of the length of
the body, and are less than 1/'2, sometimes only 1/3 , as
long as the longest rays.

The scales which cover the body, are of moderate
size, but in comparison with those of the Roach father

small. They are imbricated, and lie in distinct series.
They are of a rounded quadrilateral form, but the two
anterior corners are generally well marked and separ-
ated by sinuses from the middle of the anterior margin,
which is convex and crenulated by three or four small
undulating sinuses, into which the grooves6 radiating
forward from the central nucleus open. The posterior
(free) part of the scale contains from two to nine such
grooves, more or less distinct. The lateral line descends
at first, and then runs (from about the middle of the
abdominal region) parallel to the ventral line and nearer
to the belly than to the back. The number of scales
in the lateral line (about 52 in Dace from the north)
is the greatest possessed by any Scandinavian Leuciscus
except the Ide. The said number would thus give us
the most easily applicable character for the Dace, if it
were not stated from more southern localities that the
species may have only 44 scales in the lateral line0.

The coloration shows great resemblance to that of
the kindred species; but the colours of the Dace are
paler, less diversified, and not so bright. The top of
the head and the back are dark olive brown, the sides
silvery gray and lustrous, with a strong tinge of pale
ochreous yellow during the spawning-season. The iris
is silvery with a dash of yellow and a, narrow ring of
deeper yellow next the pupil; it is finely punctated with
green pigment, and has a dark, curved band at the top.
The sides of the head are silvery with a strong tinge
of yellow, and shade into various colours. The dorsal
fin is plain, pale grayish brown, the caudal fin bright
olive green, and all the lower fins pale with a dash of
yellow, especially at the base. The rays of the last-
mentioned fins are flame-coloured at the middle, and
the bases of the ventral fins are of the same hue. In
young specimens the lower fins are light and quite
colourless.

In the internal organs, on examining them within
the abdominal cavity, we found no essential difference
from other species.

The geographical range of the Dace extends over
the whole of Europe north of the Alps, with the ex-
ception of Scotland and Ireland, and over Western
Siberia, where the species has been found, according

a Artedt overlooked the first, short ray, but counted the last, deeply cloven ray twice over, thus arriving at the correct number of
rays in the dorsal fin.

b The number of these grooves is generally about 11; but on the caudal scales they may be much more numerous, as many as 28,
though only a few, about 9, extend to the nucleus of the scale.

c Jeitteles, Fische der March bei Olmtitz, II Abth., pp. 15 and 16.

DACE.

763

to Mela, in the River Obi0. In Scandinavia it is evi-
dently a northern and eastern fish, being' rare in Den-
mark. In the north of Sweden it is common in all
the large rivers and lakes connected with the Baltic,
from the Muonio to the Dal elf inclusive. In these
regions it is found especially in the large lakes and
also in the rivers, penetrating into Lapland as far as
the rivers afford it a passage. In the Tornea Elf it
is unknown above Kengis Force; but Widegren says
that it ascends from this river into the Muonio, in 68°
N. lat. In Lycksele Lappmark it is taken in quantities.
Its most common provincial name in the north is the
one given above. In Dalecarlia it is known as the
Straffing, and in the neighbourhood of Gefle as the
Veling. It is also common in the large rivers and
lakes of Wermland and in Lake Wener, where it is
called Staffing. South of the River Gotha we have
personally no information of its occurrence in Sweden.
Of the other Swedish names for this species we will
mention Skall-id, Skalljer , and Acfrag. In the island-
belt of Stockholm Sundevall found the Dace to be
very rare, though, according to Ekstrom, it occurs,
but is not very common, among the islands oft’ the
coast of Sbdermanland*. In Norway, according to Col-
lett, the Dace is common in the south. It is dispersed
throughout Finland, and in Russian Karelen was taken
by Malmgren in the River Kem, near the White Sea.
In Denmark, where it was not known with certainty
by Kroyer, it occurs, according to Eeddersen, in the
Nips, a river of south-western Jutland.

The Dace, like the Ide and Chub, is partial to
running water, and in spring ascends from the lakes
and the Baltic island-belts into the rivers to breed. It
is more easily overpowered by the current, however,
than the others, and therefore prefers quiet streams0.
It passes the winter in the depths of large lakes, or in

deep water among the islands on the Baltic coast.
Early in the spring, soon after the ice has broken up,
the Dace assemble in large shoals, begin to ascend the
rivers, and commence spawning in May on a sandy
bottom, as a rule eight or ten days after their arrival.
On the completion of the spawning, they usually re-
main at most a, fortnight longer, and then desert the
spawning-place. In the Klar Elf, however, the Dace, it
is stated, remains all the year round. It is one of the
less prolific Cyprinoids, the female containing a com-
paratively small number of rather large eggs. Fatio
counted about 17,400 ova, about 2 mm. in diameter,
in a female 27 cm. long.

The Dace is a timid fish, and takes to flight at
the least noise. Its movements in the Avater are very
rapid and active, and it often leaps over the net. It
is tenacious of life, young Dace being therefore excellent
live bait, especially as their skin is bright and lustrous.
The flesh is white, not very bony, and of good flavour.
Still, OAving to its insignificant size, the Dace is in little
request, and it is only in localities Avhere it is plenti-
ful, and Avhere for the time being other fish is scarce,
that this species is used as food.

The Dace is taken in the seine during the Avhole
summer, Avhere this method of fishing is employed for
other species; but only solitary specimens are caught
in this manner. It is principally taken 1) in gill-nets,
which are set in quiet bends and long Avindings of the
streams, such places being favourite haunts of the Dace
on bright and warm summer days, 2) with trammel-
nets, Avhich are used at the same time of year in shalloAV
and Aveedy inlets of the lakes, 3) in traps (see above,
pp. 32 and 33), which are set during the spawning-
season at places that the fish must pass, the narroAvest
channels and those easiest to close being selected for
this purpose. (Ekstrom, Smitt.)

a Pallas (1. c., p. 314) applies to liis Gyprinus lacustris the Finnish name of Kortsa, which, according to Mela, belongs to the
Dace. Thus, according to Pallas, the Dace is common throughout Western Siberia and especially plentiful in Lake Baikal. The Cyprinus
Grislagine of Pallas is, according to Grimm, a variety of Roach.

6 Unless Ekstr6m’s description of the Dace in the Vet.-Alcad. Handl. for 1830 referred principally to the Ide.
c Lilljeborg was told, however, that “the Dace can surmount rapids with a fairly strong current”.

764

SCANDINAVIAN FISHES.

THE IDE (sw. iden).

LEUCISCUS IDUS.

Plate XXXII, fig. 1 and Plate XXXV, fig. 1.

Scales in the lateral line about 57 (55 — < !Oa ). Branched rays in the dorsal fin 8b. Least depth of the peduncle
of the tail more than 43 % of its median length , hut as a rule somewhat less than the length of the base of the
dorsal fin. Outer (lower posterior) margin of the anal fin concave. Pharyngeal teeth hooked at the tip or straight ,

with terete and smooth , indistinct masticatory surface: 3 , 5C — 5, 3.

Fig. 190.

Pharyngeal bones and pharyngeal cartilage of Leuciscus idus , natural size, a, b, and c as in the preceding figure;
d, tlie last (hindmost) tooth in the inner (larger) row, seen from behind.

JR. hr. 3; I).

8 b

C. x + 1 + 17 + 1 + x;
Vert. 46 — 47-".

A.

Lin.

; P.

10 — IP

lat. 55—60"

— 1 — ; V. — —
17 — 18e 8—

r 8—9^

L. transv.

4—5

9’
1 ;

/ Syn . Capito fluviatilis i lie, quem Jesen appelant Germani quidam:
nos differentia caussa coeruleum cognominemus licet, Gesn.,
Hist. Anim., lib. IV, Paralip., p. 9 + Capito fluviatilis subruber,
quem Germani orfum appellant, ibid., p. 10. Cyprinus iride
sublutea; pinnis ventralibus anique rubris, Akt. Ichthyol.,
Gen., Pise., p. 5; Syn. Pise., p. 14; Descr. Spec., Pise., p.
6 + Cyprinus Orfus dictus, ibid., Syn. Pise., p. 6 4- Cypr.
cubitalis, pinna ani ossiculorum quatuordecirn, ibid., p. 7.
Cyprinus pinna ani ossiculis tredecim rubra, Lin. Fn. Suec.,
ed. I, p. 121.

Cyprinus Idbarus, Lin., Syst. Nat., ed. X, tom. I, p. 324;
Retz. Fn. Suec. Lin., p. 356 = Cyprinus microlepidotus,
Ekstr. Fiscli. Sclieer. Morka , p. 18, tab. II.

Cyprinus Idus, Lin., Syst.., 1. c.; Retz., 1. c., p. 358; Ekstr.,
Vet.-Akad. Handl. 1830, p. 146, tab. Ill; Nilss., Prodr.
Iclitliyol. Scand., p. 27 ; Ekstr., v. Wright, Skand. Fisk.,
ed. I, p. 59, tab. XI; Cuv., Val. ( Leuciscus ), Hist. Nat.
Poiss ., vol. XVII, p. 228; Kr., Damn. Fisk., vol. III. pag.

447; Lillj. (Cyprinus), Vet.-Akad. Handl. 1850, p. 305; Nilss.
(Leuciscus), Skand. Fn., Fisk., p. 306; Sundev. (Cyprinus),
Stockh. L. Hush. Siillsk. Handl., H. 6 (1855), pp. 81 et
170; Vet.-Akad. Handl. 1855, p. 15, tab. Ill, fig. 7 — 11;
Lloyd, Scandinavian Adventures, vol. I, p. 55 cum fig. (Ide);
Lindstr. (Leuciscus), Gotl. Fisk., Goth L. Hush. Siillsk.
Arsber. 1866, p. 17 (sep.); Nystr., laktt. fn. Jemtl. Vatt.
(disp. Ups. 1863), p. 4; Gthr, Cat. Brit. Mus., Fish., vol.
VII, p. 229; Coll., Fork. Vid. Selsk. Chrnia 1874, Tillaegsh.,
p. 181; ibid. 1879, No. 1, p. 96; Olss., Ofvers. Vet.-Akad.
Forh. 1876, No. 3, p. 130; Malm, Gbgs, Boh. Fn., p. 562;
M5b., Hcke, Fiscli. Osts., p. 108; Mela, Vert. Fenn., p.
328, tab. X; Reut. et Sijndm. Finl. Fisk., tab. XIII; Lillj.,
Sv., Norg. Fisk., vol. Ill, p. 207.

Cyprinus Orfus, Lin. Syst., 1. c.; var. hujus speciei, vide
Hci<l, Kn., 1. c., p. 150 + Idus miniatus, p. 151.

Cyprinus Jeses , Lin., Syst., 1. c., p. 325 ; Bl., Fische Deutschl.,
part. I, p. 45, tab. VI; Cuv., Val. (Leuciscus), 1. c., p. 160;
Mor. (Idus), Hist. Nat. Poiss. Fr., tom. Ill, p. 417.

Idus melanotus, Hckl, Kn., Siissivasserf. Oestr. Mon., p. 147;
Dyb., Cypr. Liu., p. 141; Sieb., Siissivasserf. Mitteleur., p.
176; Mgrn, Finl. Fiskfn., p. 45; Bncke, Fisch., Fischer.,
Fischz. O., W. Preuss., p. 133; Grimm., Fish., Hunt. Russ.
Wat., p. 14.

a In exceptional cases 54 or 61.
b In exceptional cases 7 or 9.

c According to MObitjs and Heincke sometimes 2, 5 or 4, 5, the latter numbers being also given by Ekstrom.
d In exceptional cases 9 or 12.
e In exceptional cases (young) 16.

/ Sometimes 10, according to Heckel and Knee.

g Sometimes 45, according to KrGyer, or 48, according to Lilljeborg.

I]

The Ide is one of the larger species within this
genus. According to the reports sent in to the Swedish
Fisheries Committee of 1881, it attains in Sweden a
length of about 9 dm. and a weight of 6 4/5 kgra. These
statements, however, undoubtedly refer to exceptions,
unless they are exaggerated, or perhaps depend on a,
confusion of this species with the Asp ( Aspius rap ax).
The ordinary length of the Ide in an adult state is
about 3 or 4 dm. At a length of 5 dm. it weighs
at most from 2 to 2 ’/a kgm.

The body is moderately compressed or even rather
bulky. The greatest depth, which lies just in front of
the beginning of the dorsal hn, usually measures in
young specimens rather more than 1/5, in old 25 — 30 %,
of the length of the body, and the greatest breadth or
thickness is not quite half the depth. The latter measure-
ment varies, however, according to sex, age, or season,
in young specimens being sometimes only about 2/6 of
the depth, and rising in old specimens to about 45 —
51 % thereof. The least depth of the body is in young
specimens about 10 % (9V2 °/o\ in old about 11 %, of
its length. The back, which is fairly broad and convex,
and which rises more or less abruptly, according to the
fatness of the fish, from the occiput, forms a slight
upward curve. The belly from the isthmus to the vent
is fairly straight, except during the spawning-season,
when it forms in the female a downward curve, be-
ginning at the chin. Between the pectoral and ventral
lins the belly is flat, but subsequently, between the
latter and the vent, becomes convex and somewhat
carinated.

The length of the head usually occupies in mode-
rate-sized specimens a little more than J/5 (22V2 — 21 1/2 %)
of that of the body. In old specimens this proportion
is almost exactly 1f., and in young ones the head is
as usual larger in comparison with the rest of the body.
The front part of the head is broad and convex, forming
a gradual curve from the snout to a point somewhat
in front of the nostrils, where it is slightly depressed".
The breadth of the interorbital space increases, even
relatively, with age from a little more6 than 37 % to
nearly 46 % of the length of the head, the latter in
specimens 4 or 5 dm. long. As a rule, however, this
breadth is less than 3/4 of the base of the dorsal fin,
and we have never found it to exceed 85 % thereof.

765

The sides of the head are. somewhat compressed and
flattened. The snout is blunt, broad, and only slightly
projecting, the margin of the upper jaw (middle of the
intermaxillaries) being the most prominent point. The
mouth is small and turned slightly upwards. In full-
grown specimens the corners of the mouth do not ex-
tend behind the perpendicular from the middle point
between the eye and the nostrils, while in young spe-
cimens they reach comparatively a little further back,
or to a line with the anterior margin of the eye. The
length both of the snout and the upper jaw increases,
even comparatively, with age, most, however, in the
former case; the length of the snout increasing in spe-
cimens 8 — 42 cm. long from about 27 to 34 % of the
length of the head, and that of the upper jaw from
about 27 to 31 % of the same. A retrogression to the
characters of youth may sometimes be found, however,
in old specimens. When the mouth is opened, the
upper jaw is slightly protruded. The length of the
lower jaw measures about 36—40 % of that of the head.
The longitudinal diameter of the eyes decreases compa-
ratively, during growth, in the above-mentioned speci-
mens, from about 30 to 15 % of the length of the head,
i. e. from a little less than the breadth of the inter-
orbital space to only V3 thereof. At the same time
the postorbital length of the head increases from some-
what less than 1/2 to about 56 % of its entire length.
The nostrils are set, as usual in the genus, close to
each other and nearer to the eye than to the tip of the
snout, the narrow partition-wall between them being
elevated into a kind of lobe. The anterior nostril is
round, the posterior crescent-shaped and larger. The
distance between the two anterior nostrils is about half
the breadth of the interorbital space. As usual in the
genus, the head is scaleless, and is covered on the sides
with a thin, transparent skin, through which the oper-
cular and facial bones are distinctly visible. The occiput
and forehead, on the other hand, are clothed with a
thicker, firmer, and richly muciferous skin, which has
a smooth and even surface. The branchiostegal mem-
brane extends behind the gill-cover in a broad rim;
its three rays are flat, broad, and curved; and below,
as is usually the case in the genus, it is united to the
isthmus, in a line with the hind margin of the pre-
operculum. Idle pseudobranchiae are well-developed.

a With regard to this depression see above, p. 716, rote a.
b In exceptional cases a little less.

766

SCANDINAVIAN FISHES.

The gill-rakers in the outer row on the first branchial
arch are 10 in number, flat at the tip, and divided into
two or three digitate branches. The pharyngeal teeth
(fig. 190) are set, as mentioned above, in two rows, 5,
as a rule, in the inner row, and 3 smaller teeth in the
outer one. During the shedding of the teeth, however,
these numbers are altered, and we find, on the one
hand, new and more or less undeveloped teeth (at first
cucullate), lying loose in the gums or beginning
to fix themselves, and on the other hand, old teeth
more or less completely detached from their alveoli.
The masticatory surface of the largest (middle and
posterior) teeth -in the inner row does not become quite
distinct until the fish has attained some considerable
size. The front tooth in this row (as well as the first,
the two first, or even all three teeth in the outer and
smaller row) is straighter and conical or cylindrical,
the remaining teeth being hooked at the tip.

The dorsal fin begins at a distance from the tip
of the snout that in young specimens measures about
48 or 49 % of the length of the body, in old 50 or
51 % thereof. Its origin generally lies above the middle
of the insertions of the ventral fins; but on examining
a number of specimens we find a variation even in this
respect. The height of this fin is greater than its length,
the former measuring on an average 18 — 16 % of the
length of the body, and the length of its base 11 or
12 % of the same. The margin is obliquely truncate,
with more or less rounded corners, at least in front;
and the fin leans slightly backwards. The beginning
of the anal fin lies at a distance from the tip of the
snout that in young specimens and the males varies
between 60 and 65 %>, in the females between about
67 and 68 % of the length of the body, though this
measurement is also subject to individual variations.
The height of the fin varies between 13 and 15 % of
the length of the body, the length of its base between
10 and 12 % of the same. When it is folded, the mar-
gin is deeply concave; when expanded, almost straight,
though even then the concavity is quite perceptible.
The length of the rays in proportion to each other is
usually such that the tips of the third and fourth rays,
which are the longest, extend to the tip of the last ray,
when the fin is completely folded. The caudal fin is
forked, with pointed corners, the lower lobe being
hardly any longer than the upper. The length of the
middle caudal rays occupies in young specimens about
12 — 10 % of that of the body, in old about 8 — 7 %

thereof, and measures in the former about 1 /2, in the
latter about 2/ 5, of that of the longest rays in the fin.

The pectoral fins are of oblong shape, with rounded
tip when laid back; their length varies between about
1 7 a/2 and 15 % of that of the body. The ventral fins
are broader and more rounded. The distance between
the latter fins and the tip of the snout measures in old
specimens about 45 or 46 %, in young specimens about
43 % , of the length of the body, the distance between
the foremost point in their insertions and the corre-
sponding point in the insertions of the pectoral fins
(the preabdominal length) in old specimens about 23 or
24 %, in young about 21 or 22 %, of the same length,
and the distance between the former point and the be-
ginning of the anal fin (the postabdominal length) in
old specimens about 21 — 24 %, in young about 18 %,
of the same. The leno-th of the ventral fins themselves

O

is about 1 3 1/2 — 14V2 % of that of the body.

The scales are of the same shape as in the pre-
ceding species, but their texture is somewhat firmer,
with denser and coarser radiating grooves, which render
their hind margin more distinctly crenulated. The la-
teral line, which slopes slightly at first, runs almost
parallel to the ventral margin, and lies much nearer to
the belly than to the back. It contains in most cases
about 57 scales, the greatest number in any Scandi-
navian Leuciscus. Above the lateral line there are 8
or 9 rows of scales, and below it 4 or 5, counting
from the dorsal and ventral fins to the lateral line, and
excluding the scales of the line itself. Between the vent
and the lateral line there are 5 or 6 rows of scales.

It is only during the spawning-season that any ex-
ternal difference between the sexes can easily be dis-
tinguished. During this period the scales of the male
are furnished with a rim, consisting of a row of clear,
yellowish, verrucose tubercles, with dark tip, which are
wanting in the female, and disappear as soon as the
spawning is over.

The coloration of the Ide is subject to considerable
variations; but the most striking alteration in its dress de-
pends on the time of year. In spring the Ide wears
its spawning-dress, and the prevailing ground-colour is
brassy yellow, which gleams through the green pigment
wherewith the scales, especially at their insertions, are
coated. This pigment grows darker towards the back,
which is almost plain grayish green, and lighter, gra-
dually disappearing, towards the belly. The head is
above of the same colour as the back, but somewhat

IDE.

767

darker; on the sides yellow, shifting into a number of
bright hues. The iris is yellow, with tine, dark dots
and a dark spot above the pupil. The dorsal and caudal
fins are dark olive gray, the former entirely plain, the
latter more or less reddish at the base of the lower
lobe. The ventral and anal fins are carnation, with
pale base and margin. The pectoral fins are pale, with
a faint tinge of red. In autumn the ground-colour is
whiter, with hardly any brassy lustre, and the pig-
ment of the scales lighter, sometimes imperceptible. The
iris is silvery white, with a fine, brassy yellow ring
round the pupil and a dark spot above the latter. The
figure (Plate XXXII, fig. 1) represents a male assuming
the spawning-dress, and is drawn from a specimen
taken in the Baltic island-belt.

The young Ide (Plate XXXV, fig. 1), known in the
islands round Morko as idplugg and lannare, in other
localities as idbarn, skall-id, gall-id , etc., differs in se-
veral respects from the adult form. It is somewhat
plumper in appearance. The head is larger in propor-
tion to the length of the body. The dorsal margin and
the occiput form an almost continuous curve. The snout
is rather prominent, extending almost in front of the
margin of the upper jaw (the intermaxillaries). The
first simple ray in the dorsal and anal fins is generally
imperceptible. The body is yellowish white, the back
darker with bright greenish tinge, the occiput dark.
The fins are light and somewhat reddish, especially
the ventral and anal, which in front shade more or
less distinctly into red.

The Gold Ide, Linnaeus’s Cyprinus or fas, a product
of Central and Southern Germany and of Austria, is a
variety often kept in ponds and aquaria.

The range of the Ide extends throughout all the
countries of Central, Northern, and Eastern Europe as
well as the west of Siberia®. In Great Britain and Ire-
land, in Switzerland and the countries bordering on the
Mediterranean, and, according to Grimm, in the Cau-
casus, the Ide is wanting. In France it is rare. In
Sweden it is spread almost everywhere, from Tornea
Lappmark to Scania. It is also common throughout
Finland, and Lilljeborg found it at Archangel; but in
Norway Lake Mjosen, according to Collett is the
northern limit of its extension. In Denmark the Ide,

which is there called Emd, has been found only on
the islands of Fiinen and Zealand.

In Sweden the Ide occurs in all the great lakes and
the waters connected with them or with the Baltic and
the Sound. But it is not equally plentiful everywhere.
In the Muonio Elf, which river, according to Laestadius
and W. v. Wright, the Ide ascends to Karesuando
(68° 30' N. lab), it is somewhat rare. This seems to
be the case wherever it occurs in Lappmark. In Jemt-
land it is common, according to Olsson, at many places
in the lowlands, but not in the highland lakes. The
Ide is most plentiful in the eastern provinces of Cen-
tral SAveden, especially to the lee of the island-belt and
in the fresh water running seaward from the coast. Off
Gothland it cannot be called rare, especially in the
month of April, according to Lindstrom, Avhen Ide are
taken in great numbers at the mouths of the rivers.

The favourite haunts of the Ide are large lakes
and the inner part of the island-belt, Avhere the Avater
is not too salt. In small lakes it is less frequently met
Avith. From these haunts, where it passes the Avinter
in deep Avater, it ascends early in the spring, soon after
the breaking up of the ice, toAvards the shore, and re-
pairs to the mouths of the rivers and brooks Avhere it
intends to spaAvn. At this season it is knoAvn by the
fishermen as the Ice-fish ( isfisk ).

The spaAvning generally takes place in Central
SAveden about the end of April. The males, mustered
in large shoals, lead the way to the spawning-place,
Avhich is chosen in very shalloAV rivers, brooks, and
meres, often Avhere narroAv ditches are the only path
open to them. During these migrations, Avhich are often
attended Avith the greatest difficulties, the Ide displays
great strength, surmounting most of the obstacles to its
progress, and skilful in avoiding the traps set in its
Avay. Like the Salmon, it leaps Avith ease over stones,
logs, and small cascades, and Avhen the Avater groAvs so
shalloAV that further advance seems impossible, to the
surprise of the observer the fish turns on its side, and
thus pursues its course. When such obstacles bar its
progress, it pauses for a Avhile, as though deliberating
Avhat path to adopt. Meamvhile several of its comrades
have come up, and Avhen one of the company has plucked
up courage and darted ahead, the others at once folloAv

a According to Pallas the Ide is common iu Siberia east to Lake Baikal and the Lena; but whether his statement really applies to
this species, is somewhat doubtful, as Pallas refers the fish in question to Bloch’s Plate XXX VI in Fisclie Deutschlands. During Norden-
skiOld’s expedition of 1876, however, an Ide 47 cm. long was taken by Theel and Trybom in the Yenisei off Goroschinskoje.

Scandinavian Fishes.

97

768

SCANDINAVIAN FISHES.

this example. Should one of them fail in its endeavour,
it then awaits the arrival of a new company before
renewing the attempt. In this manner the males press
on as far as possible until they find a spot suitable for
their object, Aveecly at the bottom and in shallow run-
ning water. Some days later, when the weather has
become mild and fair, and the water has attained a
temperature of about + 9° C. (48’2° Fahr.), the females
reach the spawning-place in a similar manner, also col-
lected in shoals, and join the males. The spawning
now begins, being always accompanied with noisy tu-
mult, and lasting about three days, night and day,
unless interrupted by a cold north wind, rain, or storm,
in which case it is postponed until finer weather sets
in. The roe is deposited on twigs, the weeds at the
bottom, stones, or some hard object, where it adheres
firmly. The eggs are numerous “ and, when deposited,
about lV2 mm. in diameter. When the spawning is
over, the fish return by the same route, male and fe-
male now in company.

The ova are hatched in 14 — 30 days6, according
to the weather; and the young, says Sundevall, are
about 7V2 mm. long when excluded from the egg.
In July, according to Cederstrom0, when the fry are
about 12 — 18 mm. long, they sometimes begin their
journey to more open water: but they generally stay
at the spawning-place, unless the water dries up, till
the end of August, Avhen they have attained a length
of about 40 — 50 mm/ They iioav rove in vast, close-
packed shoals to the shores of deeper Avater, Avhere they
remain some time among the reeds. But they seem
soon to disperse and to lead a more solitary life. At
first their groAvth is irregular and sIoav. In its second
summer, hoAvever, at the age of a year and a half, the
Ide measures about 80 — 115 mm. in length; at the
age of tAvo years, according to Marlin6, its length is
about 180 mm., and at that of three years 215 — 220
mm. According to Sundevall’s estimate the young
Ide, about 175 — 220 mm. long, sometimes to be seen
among larger Ide in the market-places of Stockholm,
are three or four years old; and the species, he says,

does not reach maturity, or a length of 3 dm., before
the age of four or five years.

In summer the older Ide are found in deep Avater
on stony shores shaded by trees, and in calm evenings
rise in companies to the surface; Avhile the young ap-
pear on fine and warm days at shallow, Aveedy places
along the shore. Towards the end of autumn the Ide
again resorts to the shallows, and often ascends the
streams and brooks visited by it in the spring during
the spaAvning-season. As soon as the lakes are frozen,
it retires to its haunts in deep Avater.

The food of the Ide consists properly of plants,
insects Avith their larvae, and crustaceans. Instances
are recorded, hoAvever, of its partiality to small fishes.
The flesh is flabby and bony, but of far from unplea-
sant taste, and retains its flavour unimpaired for a
considerable time. After someAvhat elaborate prepara-
tion it is good eating. According to Bloch it should
be stewed in beer in the same Avay as Carp, or sent
to table fried, Avith vinegar and salad-oil or a sauce
of mustard, capers, and Avine. It is also dressed in
numerous ways, either salted or dried in the sun.
When boiled, it acquires a reddish colour, Avhich may
be further heightened by alloAving the fish for some
time to lie in salt. It then resembles Salmon, and in
many places is eaten Avithout further preparation.

I11 a cauf the Ide may be kept alive for a long
time, especially if set in clear Avater with a sIoav cur-
rent. In most parts of Sweden it is knoAvn by its
common name of id; but in the southern provinces it
is called ort or art, and on the coast of Lake Malar,
in the neighbourhood of Eskilstuna, it is honoured by
the name of harp.

The most successful fisheries for Ide are those
carried on at the spawning-place and along the course
folloAved by the fish on its Avay thither. Traps ( ryssjor ,
see p. 33, fig. 7) are set in the small streams and
brooks which it passes, and also at the place Avhere it
spaAvns. They should be placed so as to close the pas-
sage entirely, for if any opening be left at the sides of
the trap, the Ide avails itself of this Avay of escape.

8 In a female weighing 1 1/2 lbs. Bloch estimated the number of the eggs at 92,720; according to Reuter a middle-sized female
lays about 70,000 eggs.

b Generally in 14 — 18 days, according to Sundevall.
c Ofvers. Vet.-Akad. Forh. 1851, pp. 163 seqq.

d According to Cederstrom and Sundevall the fry which they kept in a pond, had not attained a greater length than 20 — 28 mm.
by the 10th of September.

c See Reuter, 1. c.

CHUB.

769

Hand-nets {hdf) are also used, between two weirs or
dams built across the stream. The upper weir is kept
shut, the lower being opened at intervals, until the
fisherman sees that a sufficient number of fish have
passed, when it is suddenly closed, and the fish are
netted. The Ide is also taken in gill-nets, which are
set in spring along its route to the spawning-place,
and in autumn in shallow coves. They are seldom
used in summer; but when this is done, they are shot
in deep water off stony and shady shores. At these
latter places the Ide is caught throughout the summer
with the ‘sinking rod' (sdnkspo). The hook is baited
with grasshoppers, the shelled tails of crayfish, worms,
and, above all, beetles, the legs and wing-cases being
first removed. The seine is employed, especially during
spring, in shallow inlets near the spawning-place. In
summer the Ide is seldom taken in the seine, and then

by accident: it usually steals out of the sweep of the
seine, and if no other course be open to it, turns on
its side close to the bottom and lets the net pass over
it. The use of the ice-seine ( eckel-not ) is confined to the
island-belt, late in autumn, as soon as the ice on the
shallow, innermost coves is strong enough to bear the
fisherman’s weight. This net is hauled close in shore,
in very shallow water, where the Ide assemble before
withdrawing to their winter-quarters in the depths.
The Ide may also be speared by torchlight on dark
and calm evenings in autumn. As it never keeps still,
always moving about, considerable skill and practice
are required to strike the fish. It is seldom and only
by chance that the Ide is caught on pole lines (stdng-
krok) baited with small fish.

(Ekstrom, Smitt.)

THE CHUB (sw. FARNAN OR HARNACIvAN)-
LEUCISCUS CEPHALUS.

Plate XXXII, fig. 3.

Scales in the lateral line about 45 (44 — 46°). Branched rays in the dorsal fin 8b. Least depth of the peduncle
of the tail more than 43 % (47 %) of its length at the middle , or about equal to the length of the base of the
dorsal fin. Outer (lower posterior ) margin of the anal fin convex. Pharyngeal teeth hooked at the tip , with sharp
or worn , not very marked , and more or less distinctly granulated masticatory surface: 2 , 5 — 5, 2.

Fig. 191. Pharyngeal bones and pharyngeal cartilage of Leuciscus cephalus , natural size, «, b, and c as in the preceding figure;

d, the hindmost tooth seen from in front.

R. br. 3; D. : A. — - ; P. 1—~ V. J: C. *+1 + 17 + 1+*; L. lat. 44—46°; L. tr. - 1; Vert. 43G

(7)8 8—9° 14 — \bd 8e 3—4

a 42 — 46, according to Gunther.

43 — 48, ,, ,, Day.

39 — 45, ,, „ Steindachner.

b In exceptional cases 7.

c ,, ,, ,, 7, according to Heckel, see Kroyer.

d Sometimes 16, according to EkstrSm and Kroyer.
e In exceptional cases 10.

f 42, according to Kroyer; 43 — 45, according to Fatio.

770

SCANDINAVIAN FISHES.

Syn. Squalo Venet. et Rom., Cavedano Milan., Belon, Nat., Divers.

Poiss., p. 315. Cephalus ftuviatilis , Rond., De Pise., part.
2, p. 190 Uapito sive Cephalus , Willughb., Hist. Pise.,
p. 255; Cyprinus oblongus macrolepidotus, pinna ani ossicu-
lorum nndecim, Akt., Ichthyol. , Gen. Pise., p. 5; Syn. Pise.,
p. 7 ; + (?) Cyprinus Albo italis dictus, Syn., p. 13. Naddi
(err. pro Nadd-icl.), Lin., Act. Soc. Ups. 1744 — 50, p. 35,
tab. III.

Cyprinus Cephalus, Lin. (p. p., nec. Cyprinus cylindricus, Mus.
Ad. Fred.), Syst,. Nat., ed. X, tom. I, p. 322; Flmng ( Leu -
ciseus). Brit. Anim., p. 187 ; Ekstr., v. We. (Cyprinus),
Skand. Fisk., ed. I, p. 67, tab. XIII; Ivr. (Leuciscus), Damn.
Fisk., vol. Ill, p. 482; Dybowski ( Squalius ), Cypr. Livl.,
p. 119; Sieb., Siisswasserf. Mitteleur., p. 200; Mgrn, Finl.
Fisk., p. 46; Steind., Stzber. Akad. Wiss. Wien, Math.
Naturvv. Cl. LIV, i (1866), p. 262; Blanch, Poiss. d. eaux
donees Fr., p. 392 (+ Squ. meridionalis , p. 396, + Squ.
clathratus, p. 398); Gthr ( Leuciscus ), Cat. Brit. Mus., Fish.,
vol VII, p. 220; Coll., Forh. Vid. Selsk. Christ. 1874,
Tillsegsh., p. 181; Malm, Gbgs , Boh. Fn., p. 561; Mor.
(Squalius), Hist. Nat. Poiss. Fr., tom. Ill, p. 422; Bncke,
Fisch., Fischer., Fischz. O ., W. Preuss ., p. 137; Fatio, Fn.
Vert. Suisse, vol. IV, p. 557; Mela (Leuciscus), Vert. Fenn .,
p. 325, tab. X; Apostol. (Squalius), Pech. Gr., p. 31; Day
(Leuciscus), Fish. Gt. Brit., Irel., vol. II, p. 178, tab.
CXXXII, tig. 1; M6b., Hoke, Fisch. Osts. p. 110; Grimm
(Squalius), Fish., Hunt Russ. Wat., p. 14.

Cyprinus Idus, Bl. (syn. err.), Fisch. Deutsclil., part. I, p.
253. tab. XXXVI.

Cyprinus Dobula, Retz. (syn. err.), Fn. Suec. Lin., p. 356;
Cuv. (syn. err.), Reyn. Anim., ed. 1, tom. II, p. 195; Nilss.,
Prodr. Ichtli. Scand ., p. 26; Agass. ( Leuciscus ), Mem. Soc.
Sc. Nat. Neuch., tom. I, p. 38; Cuv., Val., Hist. Nat. Poiss.,
tom. XVII, p. 172 (+ Leuc. Albiensis , p. 194 4- Leuc. fri-
gidus, p. 234); Ivr., Damn. Fislc., vol. Ill, p. 463; Hckl
(Squalius), Stzber. Akad. Wiss. Wien, Math. Naturw. Cl. IX
(1852), p. 80, tab. VIII; Hckl, Kn., Siisswasserf. Oestr.
Mon., p. 180 (+ Squ. svalliza , p. 197 + Squ. albus, p.
198, vide Canestrini).

Cyprinus Jeses, Donov. (syn. err.), Brit. Fish., tab. CXV ;
Jur. (excl. syn.) Mem. Soc. Pliys., Hist. Nat. Geneve, tom.
Ill, part. I, p. 207, tab. XI.

Leuciscus cavedanus + L. tiberinus (= L. squalius, Val.)
+ L. Pareti + (?) L. albus, Bonap., Iconogr. Fn. Ital., part.
Ill (Pesci), tab. 112 et 113; vide Canestrini, Arch. Zool.,
Anat., Fisiol., vol. IV (1866), p. 103, et Fatio, 1. c.
Leuciscus latifrons, Nilss., Skand. Fn., Fisk., p. 309; Widegr.
(Cyprinus), Landtbr. Akad. Tidskr. 1863, pp. 202 et 208;
Lillj. (Leuciscus), So., Norg. Fn., Fisk., vol. Ill, p. 223.
Leuciscus cii, Richards., Proc. Zool. Soc. 1856, p. 375.

The Chub probably attains the same size as the
Ide, and in Sweden seems to be the largest Leuciscine,
though we do not possess such statements of its extra-
ordinary size as those given in the description of the
preceding species. Ekstrom’s largest specimen was 51/2
dm. long. Most Chub taken during the spawning-

season are of the same length as the Ide 3 or 4 dm.,
and weigh about .3 — 3 V5 kgm."

The body is shallow and thicker than that of the
Ide, and the back broader and more convex, sometimes
with a slight depression in the median line near the
occiput. The greatest depth of the body is at most
about 26 % of its length, and the greatest thickness in
adult specimens is distinctly more than half the depth,
sometimes 60 % thereof. The least depth of the body
measures in young Chub about 9 %, in old about 10
■ — 107* % °f its length. Seen in profile, the dorsal
margin rises slowly (in old specimens in a scarcely
perceptible curve) from the occiput to the beginning of
the dorsal fin, where the body is deepest, and then
slopes towards the caudal fin nearly in a straight line.
The ventral profile forms a slight curve, almost with-
out a break, from the chin to the vent.

The head, compared with the body, is somewhat
longer than in an Ide of the same size, varying in Chub
16- — -42 cm. long between about 26 and 22 % of the
length of the body. Above it is flat, with eyes com-
paratively wide apart, for the breadth of the interorbital
space, compared with the length of the head, is indeed
about equal to the corresponding measurement in the
Ide, but compared, for example, with the base of the
dorsal fin, distinctly greater, being as a rule greater
than the length of this base, or at least more than 90
% thereof. The great breadth of the forehead is highly
characteristic of the Chub at all ages, and in some
localities has gained for this species the name of I)ick-
Kopf. The frontal line is almost straight, slightly
steeper in its anterior part; and the snout is shallower
(more pointed when seen from the sides) than in the
Ide. The sides of the head are flattened and slope to-
wards the snout, which, seen from above, is broad and
more or less obtuse, but generally rather more pointed
than in the Ide. In the other structural features of the
head the Chub almost exactly resembles the Ide; but
the mouth is somewhat larger, as may most readily be
shown by a, comparison of the length of the lower jaw
with the suture between the operculum and suboper-
culum. The length of the latter is in the Ide at least
90 % (100 — 91 %, according to our measurements) of
that of the lower jaw, but in the Chub at most 90 %

“ According to Duhamel (Traite des Peches, pt. II, p. 502) the Chub attains in the Moselle a weight of 10 — 12 lbs.; in the lakes of
Upper Austria, according to Heckel, it sometimes weighs 8 or 9 lbs. Nordmann (Demid. Voy. Russ, mer., p. 484) describes an Abkhasian
variety (?) of this species that attains a length of 91/., dm.

CHUB.

771

(72 — 89 %, according to our measurements) of the same.
The gill-rakers are short and scattered, their number in
the outer row on the first branchial arch being about
the same as in the Ide (9 — 11), most of them simple,
but at least the middle ones in old specimens branched
at the tip, as in the Ide. The pharyngeal teeth (5
large ones in the inner row, and 2, a little more than
half as large, in the outer) are distinguished (fig. 191)
for their strength, their sharp, hooked tip, and the
tendency to pectination indicated by the small, terete
protuberances on the masticatory surface. The pseudo-
branchiae are well-developed.

The dorsal fin, the position of which is such that the
anterior margin lies in a line with the termination of the
insertions of the ventral fins, is rather more than twice
as high in front as behind, and its superior margin is
truncate, slightly rounded. In old specimens its shape
and backward direction are as represented in our figure;
but in young Chub it is quite upright, with pointed
front angle. It begins at a distance from the tip of
the snout measuring about 48 — 50 % of the length of
the body; its height is about 17 — 15 %, and the length
of its base about 9 or 10 %, of the same. Thus its
base is generally equal to or slightly less than the least
depth of the body", a point which among our Cypri-
nines is especially characteristic of the Chub, though
subject to exceptions, just as in young Ide the least
depth of the body may exceptionally be equal to the
length of the base of the dorsal fin. The structure of
the anal fin is unparalleled among our Cyprinines, and
thus affords a character excluding all possibility of con-
fusion between the Chub and other species. In all the

others there is a considerable difference in the height

of the fin behind and in front, the last ray being often
not half as long as the fourth or the longest anterior
ray. But in the Chub the difference is not so great,

the length of the last ray being at least about 56 — 60

% of that of the fourth. Nor is this all: the margin

of this fin in the Chub is convex, when expanded, the

middle rays too being comparatively longer than in the
rest of our Cyprinines, sometimes, in old specimens,
the longest in the whole fin. This is the case in our

figure, and we will only add that the fin may be still

more strongly expanded than it is there, and that the

margin then forms a complete curve, the greater the
older the fish. The anal fin begins at a distance from the
tip of the snout measuring about 64 — 67 %, its length
is about 8 — 11 %, and its height about 12 or 18 % (in
exceptional cases 14 %), of the length of the body. The
caudal fin resembles that of the Ide, but is generally
not so deeply forked, its middle rays occupying between
about 9 % and a little over 8 % of the length of the
body, and measuring in young Chub about half or rather
more than half, in old about 48 or 47 %, of the length
of the longest caudal rays.

The pectoral and ventral fins resemble those of the
Ide. The length of the former is about 17- — 16 % of
the length of the body, that of the latter about 15 — 14
% (in the females exceptionally 13 %) of the same. The
distance between the ventral fins and the tip of the
snout is about 46 % (in exceptional cases 47 or 48 °/o)
— 44 %1 that between these fins and the pectorals (the
preabdominal length) about 23 — 26 % (sometimes 27 %),
and that between them and the beginning of the anal
fin (the postabdominal length) about 19 — 22 %, of the
length of the body.

The scales are comparatively large and thick, but
otherwise resemble in form and texture those of the
Ide. The lateral line, which first descends in a curve
and then runs nearer to the belly than to the back, is
generally covered by 45 scales, a number which in the
Scandinavian Chub examined by us varies only between
44 and 46, in exceptional cases 47. According to other
writers, however, the variation extends, as mentioned
above, between 39 and 48. The lateral line is generally
separated from the anterior part of the base of the
dorsal fin by 7 rows of scales, and from the ventral
fins by 4 rows.

Full-grown Chub display a handsome and varied
coloration on a silvery gray ground, most like the colours
of the Roach. The back is dark olive green, the scales
are tipped with a metallic lustre. On the sides above
the lateral line the bases of the scales are dark olive
green, thus forming as it were dark frames within which
their lustrous tips are still more prominent. Below the
lateral line this dark setting fades and gradually dis-
appears, the scales acquiring a more yellowish tinge,
which grows more distinct on the belly. Behind the

a In 11 specimens 17 — 42 cm. long the average length of the base of the dorsal fin in % of the length of the body is 9‘3, and
the average for the least depth of the tail 9'4. In 5 specimens 37 — 89 mm. long, on the other hand, the corresponding averages are re-
spectively 9’9 and 91, so that the character is not distinct until the fish has attained a length of about 1 1 /2 dm.

772

SCANDINAVIAN FISHES.

gill-opening the margin of the clavicle is clyed almost
black, though this dark stripe is generally concealed by
the broad rim of the branchiostegal membrane. The
top of the head is dark olive brown, its sides and under
surface have a silvery lustre, shading into yellow with
a play of various colours. The iris is light yellow,
below silvery, above marked with a dark band. The
pectoral, dorsal, and caudal fins are nearly plain olive
brown; the ventral and anal fins, on the other hand,
of a beautiful, bright red colour, both with yellowish
base, and the latter fin with rose-coloured rays. The
coloration of the fins is, however, subject to variation.
Our figure is coloured from a specimen taken shortly
before the spawning-season. Younger Chub are paler
in hue, with the inferior fins light and colourless.

In Scandinavia the range of the Chub is confined
to the south of Sweden and the south-east of Norway.
In Denmark the Chub has not been found. In Finland
too it is a southern species, common, according to Mela,
only to the extreme south. It is one of the rarest
Scandinavian Cyprinoids, and in contradistinction to
most of them, it is taken only in small numbers and
never in any quantity, even during the spawning-season.
Retzics knew that it occurred in the Nissa, a stream
in Smaland, where it was called Bjelke. Towards the
mouth of this stream, near Halmstad, in Halland, it is
often hooked, in company with the Ide, and is here
known as harnacka. It bears the same name, according
to Lloyd0, at Falkenberg on the Atra. To Ekstrom
it was known from only three localities in Sweden,
namely, Norrkoping River, where the fishermen called
it Farna, the River Gotha off Gothenburg, where it
was entitled Dick-kopp, and Lake Hjelmar, with its af-
fluent the Stor river, where the name of the species
was Arannaren. But he remarked that its occurrence
in waters so far apart showed that these could not be
its only haunts in Sweden. It has subsequently been
found by Schagerstrom and others, according to Lillje-
borg, in the Helge in Scania; by Malm and others at
a few places in Wester Gothland and Bohuslan, e. g.
in Lake Stenstorp 20 miles east of Gothenburg, and in
the Orekil at Krokstad and Qvistrum, where it is called
Abuk; by Lloyd and others in the south part of Lake
Wener; and by Hardin and Widegren in the north of
Lake Wener and in the Klar Elf. In Norway, accord -

a Scandinav. Adventures, I, p. 64.

b Handb. Fiscliz, und Ftscherei (Max v. d. Borne), p. 134.

ing to Collett, the occurrence of the Chub is confined
to Lake Mjosen, the south part of the Glommen with
its tributaries, and the brackish water at the mouth
of the Tistedal Elf off Fredrikshald. In Central and
Southern Europe, on the other hand, the range of the
Chub is very extensive, and in England, Scotland, and
the countries bordering on the Mediterranean this spe-
cies replaces the Ide. According to Grimm the Chub
occurs throughout Russia-in-Europe, with the exception
of Transcaucasia. Richardson described it ( Leuciscus
Cii ) from Anatolia in Asia Minor. In mountainous
regions, according to Benecke6, it ascends the rivers
and brooks to a height of about 1,000 m. above the
level of the sea.

In all these southern countries the Chub has long
been better known than in Sweden. It has the repu-
tation of a strong, but timid swimmer, eagerly hiding
in the shade of any object or behind a stone, and is
accused of comparatively great voracity, not only snap-
ping up insects at the surface or such fruit as may
fall into the water, but also preying on small fishes,
frogs, and water-shrews. It possesses great cunning,
and will not bite readily, if the line be visible. From
its habit of frequenting the eddies below mill-wheels,
it has received in France the name of Meunier (miller).
According to information supplied to Ekstrom by Mr.
Lenning and Dr. Hanssen of Norrkoping and Mr.
Hamnstrom of Orebro, the Chub lives in Sweden during
the greater part of the year in lakes and extensive
pieces of water, but keeps to comparatively shallow
places, where the bottom consists of mud or weeds. In
the month of May it begins to ascend the river at
Norrkoping, and being a powerful swimmer, is met
with where the current is strongest, endeavouring with
all its might to stem the stream. About the end of
June — in more southern countries in May or even
April — the spawning-season begins, as shown by the
presence, in specimens caught in the middle of June,
of roe and milt ready to be deposited. But here as in
the case of the other Cyprinoids, this season varies, no
doubt, according to the state of the weather; and Malm
quotes a statement to the effect that in the River Gotha
the Chub spawns at the end of April. The fish as-
semble in large shoals, according to Fatio, with tu-
multuous uproar, not far from land and by preference

ROACH.

773

in slowly running water. Here the eggs attach them-
selves to the gravel and stones. According to Benecke
each female deposits about 100,000 ova. In a Chub
weighing 3 lbs. Bloch found 67,600 yellowish eggs of
the size of poppy seed.

Till the month of October the Chub stays in the
rivers, and then returns to its winter-quarters.

In Norrkoping River the Chub was taken, according
to the authorities cited above, in strong nets stretched
across the stream and in traps. As a rule, however,
this fish is caught by angling, the bait consisting of
a worm, the shelled tail of a crayfish, a fly, a cater-

pillar, a bit of cheese, or, we may almost say, any
eatable substance, for the Chub is not hard to please,
though the sight of the rod, the line, or the angler
frightens it away in a moment". The best time of day
is the morning or evening, and in the lakes the fisher-
man should choose a stony bottom, in the rivers a
rapid part or race.

The flesh of the Chub, though palatable and
not unlike that of the ‘Asp’ ( Aspius rapax), contains
numerous loose bones, and is therefore not much
esteemed.

(Ekstrom, Smitt.)

THE ROACH (sw. morten).

LEUCISCUS RUTILUS.

Plate XXXIII, fig. 1.

Scales in the lateral line about 43 {40 — 46). Branched rays in the dorsal fin 9 — 11. Least depth of the pe-
duncle of the tail more than 43 % of its length at the middle , but much less than the length of the base of the
dorsal fin. Outer margin of the anal fin concave. Pharyngeal teeth hooked at the tip or blunt , with smooth
masticatory surface , very faintly granulated , slightly pectinated , or worn into a single deep groove;

set in one row: 6(5 ) — (5)6.

a b

Fig. 192. Pharyngeal bones and pharyngeal cartilage of Leuciscus rutilus , natural size, a, b, and c as in the preceding figure.

R. hr. 3;

3

10*— 11 ’

P.

14 — 17

V.

2

8’

C. x + 1 4- 17 + 1 + x;
Vert. 40 c — 42.

L. lat. (40)43—46;

L.

tr .

8(7)
4(3) ’

Syn. Leuciscus , Rondel., De Pise ., part. II, p. 131. Rutilus sive
Rubellus fluviatilis, Gesn., Hist. Anim ., lib. IV, p. 820.
Uyprinus iride, pinnis ventralibus ac ani plerumque rubenti-
bus, Art., Ichtliyolog ., Gen. Pise., p. 3; Syn. Pise., p. 10;
Descr. Spec. Pise., p. 10; Cyprinus pinnae ani radiis duo-
decim rubicundis, Lin. Fn. Suec., ed. I, p. 124.

Cyprinus Rutilus, Lin., Syst. IVat., ed. X, tom. I, p. 324;
Odmann, Vet.-Akad. Handl. 1782, p. 163; Bl., Fiscli.,
Deutschl., part. I, p. 32, tab. 2; Retz., Fn. Suec., Lin.,
p. 357; Pall., Zooyr. Ross. Asiat., tom. Ill, p. 317; Flmng
( Leuciscus ), Brit. Anim., p. 188; Ekstr. {Cyprinus), Vet.-
Akad. Handl. 1830, p. 153; Nilss., Prodr. Ichthyol. Scand.,
p. 27 ; Agass. ( Leuciscus ndilus + L. prasinus + L. de-
cipiens), Mem. Soc. Sc. Nat. Neuch., tom. I, p. 38; Ekstr.,
v. Wr. {Cyprinus), Sleand. Fisk., ed. 1, p. 72, tab. 15;
Cuv., Val. ( Leuciscus ), Hist. Nat. Poiss ., vol. XVII, p. 130
( + L.rutiloides (ex Selys), p. 149 4- L. prasinus (ex Agass.),
p. 153 + L. Sely.sii (ex Selys), p. 198); Kr., Damn. Fiske.

0 Cf. Blanch^re, Nouv. Diet. d. Pitches, p. 174.
b Sometimes 9, according to Moreau.

c Sometimes 39, according to Fatio. 44, according to Artedi.

774

SCANDINAVIAN FISHES.

vol. Ill, p. 435; Nilss., Skand Fn., Fisk., p. 316; Sundev.
(Cyprians'), Stockh. L. Hash. Sallsk. Handl., H. 6 (1855),
pp. 81 et 174; Vet.-Akad. Handl. 1855, p. 13; Hckl, Kn.

' ( Leuciscus ), Siisswasserf. Oest.r. Mon., p. 169 (+ L. Pau-
singeri , p. 172); Dye., Cypr. Liv., p. 36; Sieb., Siisswasserf.
Mitteleur., p. 184; Mgrn, Finl. Fiskfn ., p. 46; Widegr.
(Cyprinus), Landtbr. Akad. Tidskr. 1863, pp. 201, 202, 207;
Lindstr. ( Leuciscus ), Gotl. Fisk., Gotl. L. Hush. Arsber.
1866, p. 17 (sep.); Blanch., Poiss. d. eaux douces Fr., p.
382 (+ Leuc. pallens, p. 386); Gthr, Cat. Brit. Mus .,
Fish., vol. VII, p. 212; Coll., Forh. Yid. Selsk. Christ.
1874, Tillsegsh., p. 180; ibid. 1879, No. 1, p. 96; Olss.,
Ofvers. Vet.-Akad. Forh. 1876, No. 3, p. 130; 1882, No,
10, p. 48; Malm, Gbgs, Boh. Fn., p. 557; Bncke, Fisch.,
Fischer ., Fischz., O., W. Preuss., p. 136; Mor., Hist. Nat.
Poiss. Fr., tom. Ill, p. 413; Fatio, Fn. Vert. Suisse , vol.
IV, p. 479; Mela, Vert. Fenn., p. 323, tab. X; Grimm,
Fish., Hunt. Russ. Wat., p. 14; Day, Fish. Gt. Brit., Irel.,
vol. II. p. 175, tab. CXXXII, fig. 2; Mob., Hcke, Fisch.
Osts., p. Ill; Reut., Sundm., Finl. Fisk., tab. XI; Lillj.,
Sv., Norg. Fn., Fisk., vol. Ill, p. 189.

Leuciscus Heckelii, Nordm., Voy. Russ. Merid. (Demid.), tom.
Ill, p. 491, tab. 23, fig. 1; vide Grimm, 1. c.

The Roach does not grow to any great size. Most
of the specimens caught during the spawning-season
are 15 — 20 cm. long, but many attain a length of 3
dm. The largest specimen in the Royal Museum is
from Lake Wetter, and measured 31/, dm. Nilsson
once saw a Roach 371 mm. long". At a length of
230 — 248 dm., according to Fatio, the Roach weighs
120 — 160 grammes6.

The body is rather thick, but compressed, and
generally thinner than that of the Ide. The greatest
depth varies between about 24 and 28 1/2 % of the length
of the body, the percentage being highest in old spe-
cimens and the females; while the greatest thickness
usually measures only 36 — 41 % of this depth, though
in gravid females it may rise to 54 % of the same.
The back ascends without forming any abrupt curve
from the occiput to the beginning of the dorsal fin,
from which point it slopes almost in a straight line to
the caudal fin. Throughout the greater part of its length
it is convex, but a little in front of the dorsal fin more
or less distinctly compressed or even carinated. In front
the downward curve of the ventral margin is similar
to the upward curve of the dorsal; but between the
ventral fins and the anal aperture the belly is almost

straight and slightly but distinctly carinated, and at the
beginning of the anal fin it rises at an obtuse angle.

The head resembles in form that of the Ide, save
that here the forehead is narrower0 and straighter, and
the snout more pointed, with a similar faint depression
in front of the nostrils. The length of the head is
also somewhat less, varying between about 2 1 x/2 and
20 % of that of the body. The mouth is still smaller
than in the Ide, the length of the upper jaw from the
tip of the snout being always less than that of the
latter and varying with ageJ between about 27 and 24
or 25 % of the length of the head. The length of the
lower jaw varies simultaneously between about 36 and
32 % of that of the head, and is always less than (about
95 — 84 % of) that of the suboperculum along the suture
at the lower margin of the operculum. The longitu-
dinal diameter of the eyes varies in the same speci-
mens between about 27 and 18 % of the length of the
head, or between about 75 and 48 % of the breadth of
the interorbital space. The postorbital part of the head
always measures somewhat less than half the entire
length of the same, unless the rim of the branchiostegal
membrane be taken into account, in which case it
slightly exceeds half the said length. The position of
the eyes is also such that the lower margin of the pu-
pil touches the line from the middle of the caudal fin
to the margin of the upper jaw. The nostrils resemble
those of the Ide. The gill-openings are separated be-
low, here as in the preceding forms, by the isthmus,
which is rather narrow, and to which the branchiostegal
membranes are united. The pseudobranchiae are free
and comparatively large in young Roach, in older spe-
cimens less distinct. The gill-rakers are short and
scattered, numbering 10 — 14 in the outer row on the
first branchial arch, 10 on the outer margin of the
pharyngeal bones. In contradistinction to the three
preceding species, the Roach (fig. 192) has only one
row of teeth on each of the pharyngeals, generally 6
on the left and 5 on the right, though these numbers
may vary, being sometimes only 5 on each side, some-
times 6. The first two teeth, here as in the preceding-
forms, are almost straight, blunt, conical, and without

a M6bius and Heincke give 5 dm. as the maximum length of the Roach.

b According to Isaak Walton the Roach may attain a weight of 2 lbs. (907 grammes); and in ‘The Field’ (2 Nov., 1881) we read
of a Roach weighing 3 lbs. 1 2 1 2 oz. (949 grammes), taken by Mr. Stead in Bedfordshire.

c The breadth of the interorbital space is less than 38 % (35 — 37’8 %) of the length of the head, or than 2/3 (52 — 64 %) of the
length of the base of the dorsal fin.

In our specimens, which are between 100 and 330 mm. long.

ROACH.

775

hooked tip, the others strongly curved at the tip, more
or less compressed in a transverse direction (back and
Iront), and before they are worn, more or less distinctly
crenulated (pectinated), the hindmost tooth, which is
the least worn, most distinctly. Sometimes only a trace
of this crenulation remains, in the form of small pro-
tuberances on the masticatory surface, which is eventu-
ally hollowed into a smooth, simple furrow. A strik-
ing difference between the pharyngeals of the Roach
and those of the preceding Leuciscines, and a point that
calls to mind the Tench, is the prolongation common to
the bases of the posterior teeth, the last two or three
being set on a process jutting from the pharyngeal
bone into the pharynx. The heart-shaped pharyngeal
cartilage of the three preceding species is modified here,
this organ being elongated into a tongue-like form, with
the anterior (upper) end obtusely rounded, and the
posterior, tapering extremity only slightly folded and j
elevated, scarcely marked off from the other part.

The dorsal fin, which is particularly remarkable
for its height, is obliquely truncate at the margin, with
pointed corners, especially behind. It begins at a di-
stance from the tip of the snout that- increases with
age and measures about 441/2 — 4872 % of the length
of the body, or half the length to the base of the caudal
fin. This distance is on an average less than in any
of the three preceding species. In this respect the Roach
comes nearest to the Cyprinine group; and the other
Leuciscines have evidently followed a special direction
of development from that group, reaching different
stages. The Dace has advanced furthest, as shown by
the table given below". The base of the dorsal fin
measures about 14 — 12 %, its height about 20 1/2 — 17 ,
%, of the length of the body. The anal fin, the di-
stance between which and the tip of the snout is about
63 — 68 % of the length of the body, resembles in form
that of the Ide. The length of its base is about 12 —

1 0 1 2 %, its height about 137* — 1072 of the length
of the body. The caudal fin is rather deeply forked,

its middle rays occupying about 9 — 8 % of the length
of the body and measuring about 36 — 40 % of that of
the longest caudal rays.

The pectoral and ventral fins have the same form
as in the Ide, but differ only slightly in length, that
of the former varying between about 17 and 1472 %,
that of the latter between 17 and 14 %, of the length
of the body. The distance between the ventral fins and
the tip of the snout measures about 43 — 47 %, the pre-
abdominal length about 22 — 25 %, and the postabdo-
minal length about 20 — 22 %, of the length of the body.

The scales resemble those of the three preceding spe-
cies, but are comparatively much larger than the Ide’s,
and as a rule larger even than the Chub’s, a point which
is shown by the above estimates of their number in the
lateral line and in a transverse row between the dorsal
and ventral fins. During the spawning-season they are
destitute of the rim then present round the scales of
the Ide, but are strewn in the male with the usual,
pointed, verrucQse tubercles.

The sexes are distinguished only by the more
slender body of the male and the said spiniferous tu-
bercles, which are scattered during the spawning-season
on the skin of the head, the opercula, the scales co-
vering the forepart of the body, and the first ray of
the pectoral fins. According to a note on v. Wright’s
original, our figure represents a male without this ex-
ternal sexual character.

The upper part of the head and back is of a, dark,
blackish green, which rapidly passes below into brighter
green, and then gradually into the lustrous silvery gray
of the sides, which at the middle are faintly tinged
with bronze and towards the belly shade into yellow.
Each of the scales is marked at the base with a green-
ish spot, darker on the uppermost scales, gradually fad-
ing and at last disappearing on the lower ones. In the
lateral line the scales are furnished with two similar,
but smaller spots, lying one on each side of the duct,
which is yellowish. In some specimens the iris is of

a

Aver a g c*.

Leuciscus rutilus.

Leuciscus cephalus.

Leuciscus idus.

Leuciscus

grislagine.

3 specimens
102 — 153 mm.

long.

3 specimens
167 — 327 min.
long.

5 specimens
37 — 89 mm.

long.

6 specimens
170 — 196 mm.
long.

5 specimens
244 — 424 mm.

long.

4 specimens
84 — 148 mm.
long.

4 specimens
181 — 416 mm.
long.

5 specimens
178 — 314 mm.
long.

Length of the body expressed in millimetres

127

224.7

67.4

183.4

297.8

124

305

196.2

Distance between the dorsal fin and the tip of the

snout in % of the length of the body

45.8

47.7

48.1

48.4

49.o

49.8

50. o

50.4

98

Scandinavian Fishes.

776

SCANDINAVIAN FISHES.

a handsome bright red, but this tint is not constant,
seeming, on the contrary, to have some hxed relation
to age, probably to the seasons, and above all to the
nature of the water inhabited by the fish. In some of
the great lakes, and also in the island-belt, the iris is
usually pale orange, with a spot of darker red above
the pupil. In young specimens it is very pale with a
faint dash of red; but in old Roach that live in small
lakes with thick water, the iris is often of so bright a
red that it has given rise to the saying “rodogd som
en mart ” (red-eyed as a Roach). The dorsal and caudal
fins are of a plain, light olive brown. The other fins
are yellowish, more or less tinged with red, especially
the ventral and anal, whose rays are of a, still brighter
red between the middle and the tip. The colour of the
fins is, however, highly variable, and seems to be sub-
ject to the same influences as that of the eyes.

The geographical range of the Roach embraces the
whole of Europe north of the Alps and the Pyrenees,
except Ireland and the west and north of Norway.
The species is extremely common, according to Grimm,
in the Sea of Azov and the Caspian Sea. It is also spread
throughout Siberia,, from the east of which region spe-
cimens were secured by Humboldt and Ehrenberg.
Lilljeborg found it at Archangel, where it was also
met with by Lieutenant Sandeberg. Still it is 'want-
ing, according to Reuter, in the White Sea as well as
in the Arctic Ocean. The Roach is one of the com-
monest and most plentiful fishes not only in the lakes,
rivers, and streams of Sweden, from Tornea Lappmark
to Scania, but also in the island-belt of the Baltic, as
well as in the Gulfs of Bothnia and Finland. In Swe-
den its range is roughly co-extensive with that of the
Perch, and it ascends high among the mountains. Ek-
strom found the species in a small tarn on Akkalis-
pudive, a mountain in Pitea Lappmark".

In summer the Roach frequents weedy shallows
near shore. It passes the winter in deep water, but as
soon as the great lakes are open in spring, it ascends
in large shoals to the shores, where it spawns at the
beginning of May. In their upward course the shoals
are so distributed that the males lead the way, and
consequently are the first to arrive at the spawning-

place, being hence called Ismortb (Ice Roach). The
females, which are known as Lekmort (Spawning Roach)
or Lofmort (Leaf Roach), arrive about a fortnight later.
They now join the males and commence spawning
among twigs and weeds, often “in water so shallow,”
says Sundevall, “that it seems hardly sufficient to
cover the spawning fish.” The spawning lasts from 3
to 9 days, according to the weather. During the ope-
ration the fish pack themselves in a dense mass, and
move towards the surface with such rapidity as to pro-
duce a quick hissing noise, interrupted and repeated at
brief intervals. In Lakes Som men and Wetter the older
Roach often spawn, according to Widegren, on a stony
bottom some distance from land. The roe is fine, con-
taining numerous eggs: in a gravid female 18 cm. long
the eggs, when almost ripe, were about 1 V2 nim. in
diameter; and in a female 275 grammes in weight,
with ovaries weighing 60 grammes, Lund" counted
nearly 72,000 eggs. The ova are deposited on the
twigs and the weeds at the bottom, “often so near the
surface,” says Sundevall, “that they are now and then
left dry, but are none the worse for this.” They are
hatched in 10 — 14 days. “The fry generally lie still
at the bottom, resting on their side, or supported by
and as it were suspended from plants, straws, and the
like. Gradually they begin to move and to swim a
little better, and after the yolk has disappeared, which
apparently happens in 8 — 10 days, they keep swimming
about in dense shoals among the reeds. At the age ot
two months they are 15 — 20 mm. long and fully de-
veloped in external form.” (Sundev.). When three
years old, they have attained a length of about 100 —
125^ mm.

The Roach leads a sociable life, and roves along
the shores all the summer in large and small troops.
It seems as though companionship inspires it with con-
fidence, for it is not very shy. It is indeed afraid ot
noise, but soon returns to the spot from which it has
been frightened away. The composition of the water
in which it lives, exercises great influence not only on
its appearance, as we have mentioned above, but also
on its flavour. When the Roach has its home in pure
water, the flesh is white and free from taint; in fish

“ In Switzerland, according to Fatio, the Roach in a natural state hardly ascends higher than the lakes about 700 metres above the
level of the sea; but it has been planted there in lakes 1,160 metres above the same level.

0 Also Badfislc (Bather), see Lund, Vet.-Akad. Handl. 1761, p. 186.
c L. c., p. 194.

d 135 mm., according to Reuter.

ROACH.

777

from impure and muddy water it turns red after boil-
ing- and acquires a strong, unpleasant taste of mud.
On attaining a length of 20 — 25 cm. the Roach gener-
ally grows fat, and is then known as Kartmort or
Grytmort. Its fatness depends, however, on the food-
supply, which consists of vegetable substances, insects,
larvte, and mollusks. It seldom lives long in a cauf,
unless taken during the spawning-season.

The Roach is one of our most useful fishes, both
as a welcome addition to the coarse fare of the poor",
and as a valuable bait for larger and more esteemed
tishes, in which respect it is inferior to none of our
indigenous piscine forms. Though it is most often
taken, in great or small numbers, with seines of several
kinds, other tackle is also used, consisting of traps ,
which are set during the spawning-season in weirs
(verke, see above, p. 32), or Roach-nets, which are etn- j
ployed either in summer, when they are shot off weedy
shores, or during the spawning-season, round the weirs
in which the traps are placed. At the latter season
they may also be set round the reeds or beds of weeds
where the Roach spawns, in which case the fish are
driven towards the net with the fork (see above, p.
741, fig. 185). Lastly, the Roach is taken with rod
and line, and bites during the whole summer. Winter-
angling on the ice is practised only to procure Roach
for bait. But few fish are caught in this manner.

The above characters seem quite sufficient to ren-
der the Roach easily recognisable; but both in form
and coloration it sometimes varies beyond comprehen-
sion, and several nominal species have thus originated.
Fatio has arranged these varieties in three groups:

1) Leuciscus rutilus, var. data, with body of extra-
ordinary depth, closely resembling that of the Rudd —
to this group belong the Leuciscus rutiloides of Selys-
Loxgchamps6 and Nordmann’s (1. c.) Leuciscus Heckelii;

2) X. rutilus, var. elongata, with body extraordinarily
elongated (shallow) and only slightly compressed, more

like that of the Dace — comprising IIeckel’s leuciscus
Selysii and Agassiz’ X. prasinus; and 3) X. rutilus,
var. crassa, with body of extraordinary thickness, in
appearance not unlike the Chub, but most nearly allied
to the Italian Leuciscus aula. Heckel found in Lake
Egel (Upper Austria) a form which lie called Leuciscus
Pausingeri, with comparatively high dorsal and low
anal fins. This form was again met with by Malm in
Lake Bolmen (Sweden), and hence was named by him
Leuciscus rutilus, forma bolmensis. Fatio discovered
a remarkable malformation of the Roach in Lake Bru-
nig, an Alpine lake now almost dry, and situated 1,160
in. above the level of the sea. Roach and Perch were
introduced into this lake at about the middle of the
last century. As the lake gradually dried up and
shrank into a small, deep pool of very clear water, the
Roach, which he never found to measure more than
16 cm., and whose eyes, as usual in fishes of stunted
growth, were comparatively large, had suffered an
albinotic change of colour. The back was pale green,
the sides were pure silvery white, the fins almost colour-
less, but the eyes deep red. In most of them too —
perhaps because they were compelled, like Bleak, to
seek their food at the surface — the mouth was more
oblique than usual, resembling that of the Rudd, with
more projecting chin. In other localities, in Northern
Germany (Danzig and the Frische Haff, according to
Siebold), in France (the Seille, a tributary of the Mo-
selle), and in Switzerland (the Lake of Geneva), the
said albinism has produced the Gold Roach, a form
analogous to the Gold Ide already mentioned. To con-
clude, the sociable propensities of the Roach, in Sweden
and in many other places, have induced it to join com-
pany with other Cyprinoids engaged in spawning.
Hence hybrids — or at least forms reasonably capable
of this interpretation — have been found between the
Roach and the Rudd, Bleak, Bream, and White Bream.

(Ekstrom, Smitt.)

a The Roach is equally important in South-eastern Russia, where, according to Grimm, three or four hundred millions are annually
taken in the Caspian Sea, and about a hundred millions in the Black Sea with the Rivers Kuban and Don.
h Faime Belrje, p. 212.

778

SCANDINAVIAN FISHES.

Genus SCARDINIUS.

Scales middle-sized. Lateral line complete. Lower jaw slightly projecting. Caudal lobes pointed. Length of
the base of the anal fin more than 19 % of the distance between this fin and the tip of the snout. Distance
between the dorsal fin and the tip of the snout less than 86 % of that between the anal fin and the same point.

In this genus we draw still nearer to the Abra-
inidines, a circumstance most clearly shown by the
elongation of the base of the anal tin, as remarked in
the above diagnosis. In all our specimens of the pre-
ceding Leuciscines the base of the anal tin measured
less than 7s °f the length of the body; henceforward
it is always at least somewhat longer, except where
cross-breeding impairs the natural form of the species.
When Bonaparte first established this genus (merely
as a subgenus of Leuciscus'1) , adopting its title from the
Italian name ( Scardola or Scar dine) of a species iden-
tified by Canestrini with the Rudd* 6, he laid most
stress on the deep form of the body, the position of
the dorsal fin over the space between the ventral and

anal fins, and the carination of the ventral margin
between the last-mentioned fins, characters all of which
we have seen indicated in the Roach. When Heckel
at a later period reformed the generic classification of
the Cyprinoids, he characterized the genus Scardinius c
chiefly by the pectination of the pharyngeal teeth as
in the Bleaks; but we have also seen traces of this
character in the Roach. The genus, as we understand
it from our knowledge of a single species'*, comes so
near Leuciscus that it can claim systematic recogni-
tion only as a remarkable stage of transition, in the
external form to Abramis, and in the oblique
mouth and the pectinated pharyngeal teeth to Al-
burnus.

a Iconogr. Fn. Ital ., Pesci , in the description of Leuciscus squalus. r

6 To judge by the measurements given by Canestrini (Arch. Zoo]., Anat., Fisiol., vol. IV, p. 89) of the Italian scarclola , these spe-
cies seem, however, to be distinct.

c Russeggers Reise, II Th., p. 1037.

d All the five species established by Bonaparte and Heckel, which would otherwise belong to this genus, are combined into one spe-
cies by Canestrini and Fatio; but it is quite possible that hybrid forms are thus contained within the limits of this species.

LEUCISCINES.

779

THE RUDD (sw. sarfven).

SCARDINJ US ERYTHROPHTH ALMUS.

Plate XXXIII, fig. 2.

Scales in the lateral line about 42a. Dorsal Jin situated over the space between the ventral and anal Jins , and
at a distance from the tip of the snout greater than half the length of the body. Ventral margin between the
ventral fins and the anal aperture carinated. Pharyngeal teeth set in two rows , hooked at the tip,

and pectinated: 3, 5 — 5, 3.

Fig. 193.

Pharyngeal bones and pharyngeal cartilage of Seardinius

erythrophthalmus, natural size.

a, b , and c as in the preceding figure.

R. br. 3; D. v; A.

8— 96 10 — 12'

P.

15—17

V - ■

Sd ’

7(8)

C. x + 1 + 17 + 1 + x; L. lat. 41 — 43“; L. transv. fi-f 1 ; Vert.

(3)4

37 — 39.

Syn. Rootciug id est, Erythrophthalmus Germanis dictus, Brands
affinis, Willughb., Hist. Pise., ed. Raii, p. 249. Cyprinus
iride, pinnis omnibus, caudaque rubris, Art. Ichthyol., Gen.
Pise., p. 3; Syn. Pise., p. 4; Spec. Pise., p. 9. Cyprinus
pinnae ani radiis quatuordecim, pinnis omnibus rubris, Lin.,
Fn. Suec ., ed. I, p. 123.

Cyprinus Erythrophthalmus, Lin., Syst. Nat., ed. X, tom. I,
p. 324; Osb., Vet.-Akad. Hand!. 1771, p. 152, tab. IV,
fig. 4; Bl., Fisch. Deutsclil., part. I, p. 28, tab. I; Retz.,
Fn. Suec. Lin., p. 358; Donov., Brit. Fish., tab. XL; Ascan.
(Rathke), Icon. Pier. Nat., cab. V, p. 4, tab. XLII; Flmng
{Leuciscus), Brit. Anim., p. 188; Ekstr. {Cyprinus), Vet.-
Akad. Handl. 1830, p. 162; Nilss., Prodr. Ichthyol. Scand.,
p. 28; Ekstr., v. Wr., Skand. Fisk., ed. I, p. 74, tab. 16;
Cuv., Val. {Leuciscus), Hist. Nat. Poiss., vol. XVII, p. 107 ;
Kr. Damn. Fisk., vol. Ill, p. 421; Nilss., Skand. Fn., Fisk.,
p. 313; Sundev. {Cyprinus), Stockh. L. Hush. Sallsk. Handl.
1855, pp. 81 et 174; Hckl, Kn. ( Seardinius ), Siisswasserf.
Ostr. Mon., p. 153 (+ Scard. dergle, p. 156 + (?) Scard.
scardafa (ex Bonat.), p. 157 + (?) Scard. plotizza p. 159

+ Scard. macro phthalmus, p. 160); Dyb., Cypr. Lid., p.
134; Sieb. , Siisswasserf. Mitteleur., p. 180; Widegr. {Cy-
prinus), Landtbr. Akad. Tidskr. 1863, p. 202; Mgrn {Scar-
dinius), Finl. Fiskfn., p. 45; Lindstr. {Leuciscus), Gotl.
Fisk., Gotl. L. Hush. Sallsk. Arsber. 1866, p, 17 (sep.);
Canestr. {Seardinius), Arch. Zool., Anat., Fisiol., vol. IV
(1866), p. 89; Blanch., Poiss. d. eaux donees Fr., p. 377;
Gthr ( Leuciscus ), Cat. Brit. Mus., Fish., vol. VII, p. 231 ;
Coll., Forh. Vid. Selsk. Chrnia 1874, Tilhegsh., p. 182; N.
Mag. Naturv. Chrnia, Bd. 29 (1884), p. Ill; Malm, Gbgs ,
Boh. Fn., p. 563; Fedders. ( Seardinius ), Naturh. Tidskr.
Kbhvn, ser. 3, vol. XII, p. 88; Mor. Hist. Nat. Poiss. Fr.,
tom. Ill, p. 410; Bncke, Fiscli., Fischer., Fischz. O., W.
Preuss., p. 134; Mela ( Leuciscus ), Vert. Fenn ., p. 324, tab.
X; Day, Fish. Gt. Brit., Irel, vol. II, p. 183. tab. CXXXIII,
fig. 2; Fatio ( Seardinius ), Fn. Vert. Suisse, vol. IV, p. 457;
Mob., Hcke {Leuciscus), Fisch. Osts., p. 112; Reut., Sundm.,
Finl. Fisk., tab. II; Lillj., Sv . , Norg. Fn., Fisk., vol. Ill,
p. 233.

Cyprinus compressus, Hollb., Beskr. Boh. Fisk., H. Ill, p. 66
cum tab. (efr Ekstr., Gbgs Vet., Vitt. Sarnh. Hand]., N.
Tidsf., II. 1 (1850), p. 24).

Cyprinus erytlirops, Pall., Zoogr. Ross. Asiat., tom. Ill, p. 317.

The Rudd, which is usually of no considerable
size, measuring in most cases 15 — 20 cm., seldom

° Sometimes 39, according to Ekstrom; sometimes 44 or even 45, according to Moreau.
* Sometimes 10, according to Fatio.
c 11, according to Kroyer.
d Sometimes 7, according to Fatio.

780

SCANDINAVIAN FISHES.

30 — 36 cm., still attains a weight of about 1 kgm."
In external form it seems to occupy an intermediate
position between the Leuciseines and the Crucian Carp,
which it so closely resembles in this respect that in
some localities it bears the name of Sjoruda (Lake
Crucian Carp). In Halland and Blekinge it is also
called JRitda (Rita), and on the Scanian coast of the
Sound it is known by its Danish name of Rudskalle
(Dan. skalle, Roach). The body is deeper and, in pro-
portion to the depth, more compressed than in the pre-
ceding Leuciseines, the greatest depth being about Vs 6
of the length to the middle of the caudal tin in young
specimens, in old somewhat less than 2/ 5C of the same.
In this respect, however, the Rudd undergoes a change
of growth which during youth ranges it close beside
the Roach in a more developed state A The greatest
thickness is about 32 — 42 % of the greatest depth.
The least depth of the peduncle of the tail measures
about 710 (8- — 11 %) of the length of the body. The
back rises somewhat abruptly from the occiput, form-
ing a regular curve to the beginning of the dorsal fin,
where it bends at an obtuse angle, and then descends
in an elongated S-shaped curve, or sometimes almost
in a straight line, towards the base of the caudal tin.
It is convex throughout the greater part of its length,
but in front of the dorsal tin compressed at the mar-
gin, as in the Roach. The belly is rather strongly
compressed, but Hat to the ventral fins, then carinated
to the anal aperture. It runs in an unbroken curve
from the chin to the vent, where it forms an angular
bend, and then rises almost in a straight line to the
peduncle of the tail.

The head is of moderate size, measuring about Vs
of the length of the body b and compressed, with broad

and almost straight forehead ' and small, blunt, flattened
snout6'. The mouth is small* * and turned upwards, the
lower jaw being somewhat longer than the upper, so that
the tip of the chin is the most prominent point. The
eyes are somewhat larger than in the Roach, their
longitudinal diameter measuring in middle-sized Rudd
about 23 %x of the length of the head. Their position
is such that the line from the middle of the caudal tin
to the margin of the upper jaw cuts the centre of the
pupil; and in adult specimens the postorbital length of
the head is equal to or a little greater than half the
entire length thereof, but in the fry sometimes only
about 43 % of the same. The nostrils are similar to
those of the Roach. The gill-rakers also resemble those
of the Roach; but the pharyngeal teeth (tig. 193) are
set in two rows, 5 in the inner and 3 in the outer,
their masticatory surface is coursed by transverse grooves
on the sides and granulated (pectinated) at the margin,
and the pharyngeal cartilage is furnished on the sides
with oblique, transverse grooves.

The dorsal tin occupies the same backward posi-
tion as in the Minnow, the Owsianka, the ‘Asp’, and
some specimens of the Dace, lying opposite, not to the
ventral tins, as in the other Leuciseines, but to the
space between the ventral and anal fins, as in the
Abramidines. It begins at a distance from the tip of
the snout measuring about 52 — 56 % of the length of
the body; and the distance between the ventral fins
and the same point is only about 79 % (78 %) — 84 %
(85 %) of the former distance. The form of the fin,
on the other hand, is the same as in the rest of our
Leuciseines, the length of its base measuring about
11 — 13 % (13 V2 %)■> and its height about 20 — 17 %, of
the length of the body. The anal fin is distinguished

a According to the reports sent in to the Swedish Fisheries Committee of 1881 — 83 the Rudd(?) may attain a length of 1 fg Sw. feet
(45 cm.) and a weight of 4 Sw. pounds (1,700 grammes). Day tells us of a specimen from Norfolk 3 lbs. 1 oz. (1,389 grammes) in
weight. According to Fatio the Rudd weighs at a length of 15Y2 cm. 45 grammes, at a length of 266 mm. 230 grammes, and at a
length of 3 dm. more than Y2 kilo.

* 26—36 %.

c 36 — 3 9 1 /2 /■> according to our measurements.

d In a Roach 167 mm. long the greatest depth was 26'3 %, and the greatest thickness 10'8 %, of the length of the body. In a

Rudd 63Y2 mm. long these percentages were respectively 26'8 and ll'O.

e About 23 — 20 / ( 1 9 ‘ 8 %) of the length of the body in Rudd 63 — 277 mm. long.

f The breadth of the interorbital space increases with age in the specimens just mentioned from about 40 to 47 % of the length of
the head.

v In the same specimens the length of the snout varies individually, though generally increasing, from about 28 % (in exceptional
cases 26 %) to 331/., % of that of the head.

h The length of the upper jaw from the middle of the tip of the snout varies in the same specimens between about 28 % (exceptionally

27 %) and 32 % of that of the head, and the length of the lower jaw between 39 and 36 % (34 %) of the same, being in young specimens

often equal to, in old always less than, that of the suboperculum.

1 In our smallest specimen, 63 mm. long, the longitudinal diameter of the eye measures 32 % of the length of the head, in our
argest specimen, 277 mm. long, 21 % of the same.

RUDD.

781

by its great length, its base measuring about 13 — 15 V2
% of the length of the body, its height about 16 %
(sometimes 17 7o) — IS1/, % of the same. It begins at,
a distance from the tip of the snout measuring about
59 — 66 % (in exceptional cases 69 %) of the length of
the body. The caudal tin is similar to that of the
Roach, its middle rays occupying about 1 1 1/2 — 71/2 %
of the length of the body, and measuring in our young-
est specimen about 53 %, in the oldest ones about 40 —
36 % of that of the longest caudal rays.

The pectoral arid ventral tins resemble in form
those of the Roach, but the former are always per-
ceptibly longer than the latter, the length of the former
being about 20 — 18 %, of the latter about 18 — 16 %,
of that of the body. The distance between the ventral
tins and the tip of the snout measures about 43 (42 1/2)
— 46 %, the preabdominal length about 22 — 25 (25'8) %,
and the postabdominal length about 19 — 25 (27) %, of
the length of the body.

Tire scales are also of the same type as in the
Roach, but larger and coarser, more distinctly striated,
and still more closely imbricated.

To the external sexual characters the same re-
marks apply as in the case of the Roach.

In coloration the Rudd is one of the most hand-
some Leuciscines. The sides are of a silvery lustre,
strongly tinged with golden yellow, a bright play of
colours being thus produced. The back is dark green-
ish, gradually passing first into greenish yellotv and
then into the colour of the sides. The iris is golden
yellow, shading more or less distinctly into red. The
dorsal and caudal fins are of a light olive green, tip-
ped more or less distinctly with red, especially the
latter. The pectoral fins are light and transparent, with
reddish tip. The ventral and anal fins, grayish yellow
at the base, are of a bright red hue, of varying in-
tensity and distribution in different specimens. In young
specimens, as usual, the colours are less rich.

The range of the Rudd extends throughout Europe,
except the Iberian Peninsula, and also to Asia Minor
and the interior of Siberia, where Humboldt and Ehren-
berg traced it to the Obi and Tobolsk. But both in
Siberia and Europe the species is wanting in the ex-
treme north. In Finland, according to Mela, it penet-
rates to lat. 63° 20’ N. How far north the Rudd is

found in Sweden, has not yet been ascertained; but
there are no positive instances of its occurrence in
Westernorrland, Jemtland, or Westerbotten, though Ek-
strom was told that it occurred still farther north, in
the Kalix Elf. Of its extension in Norway Ratiike
states (in Ascanius) that it is taken in several of the
Norwegian lakes, among others in Mjosen and Ojeren;
but according to Collett its occurrence is confined to
the south-east of the country, and its range scarcely
extends north of Christiania (60° N. lat,). In Ireland
the Rudd and the Minnow are the only indigenous
Cyprinines known.

In Southern and Central Sweden the Rudd is met
with in most of the lakes and rivers, as well as in
the Baltic island-belt, but nowhere in any great abund-
ance. It is known in different localities by different
names, most often by those of Sarf“ or Sarfvel, some-
times Rodfena (Red-fin) or Rodmort (Red Roach). Its
favourite haunts lie in thick and weedy water. In
spring and summer it frequents shallow, swampy coves
with weedy or muddy bottom. In winter, like fhe
other Cyprinoids, it withdraws to deep water.

The Rudd is not very active, and makes long so-
journs at its chosen haunts. Except on very warm
and fine summer days, it seldom rises in the water,
but lies at the bottom, buried in weeds and mud. It
is more cautious than timid, for though afraid of noise,
it does not retire to any distance, but hides, when
frightened, in the ooze and weeds, whence no din can
dislodge it. Being a greedy eater, and living on plants,
insects, worms, and mud, it indeed becomes fat towards
autumn, but in the Swedish lakes never attains any
considerable size. It is very sociable and cannot en-
dure solitude, intruding itself, when it cannot find any
large company of its own species, on the society of
other fishes, generally those of the preceding genus,
especially during their spawning festivities. Hence the
origin of the Swedish proverb: Scirfven i hvar lekb.
From this habit, and from the fact that large shoals
are seldom found breeding at the same place, it has
been supposed that the Rudd holds no spawning
assemblies of its own. This opinion is, however,
groundless.

The spawning-season of the Rudd occurs in Central
Sweden at the end of May or beginning of June, in

" In Finland iSorua , in Norway Seru or Flasroye (Rathke).
b Lit. “the Rudd at every spawning,” i. e. a finger in every pie.

782

SCANDINAVIAN FISHES.

more southern countries earlier, sometimes even in April.
The males and females then join company and gather
in shallow, weedy inlets, where the spawning takes
place, accompanied by a babbling or smacking noise.
This sound is produced by the fish putting their snouts
to the surface, opening the mouth, and emitting an
air-bubble, which floats on the water and bursts. The
roe is deposited on the weeds, and is hatched in 8 — 10
days, the length of time required depending on the
weather. In a female weighing 293 grammes Bloch
estimated the number of the eggs at 91,720.

The Rudd is used as food only by the poor. The
flesh is flabby, bony, and always more or less tainted
with mud. Furthermore, as the fish is hardly ever
found, even during the spawning-season, in large shoals,
it is only seldom that the fisherman pays any special
attention to it. Traps ( ryssjor , see p. 33, fig. 7) are
set at the spawning-place, in which case care should
be taken to lay them close to the bottom. Wicker-
baskets ( mjardar , see p. 32, fig. 6) may also be em-
ployed at the spawning-place, but are less useful. In
the trammel-net the Rudd is taken all the summer on
warm and tine days and at places thickly overgrown
with weeds. The trammel should be shot among the

weeds, for if it be set outside them, the Rudd refuses
to be driven into the meshes. By angling it may also
be caught throughout the summer, for it bites eagerly
at a worm. It is almost always taken in company
with other fishes.

The Rudd’s fondness for joining, as an interloper,
in the spawning of other fishes has produced several
hybrids, one of which, Jacket s“ Scardiniopsis anceps ,
is a cross between this species and the preceding one.
In this form the position of the dorsal fin is the same
as in the Roach; but the size of the scales and the
pectination of the pharyngeal teeth remind us of the
the Rudd. The dorsal fin contains 10 branched rays,
the anal 11 or 12. The number and distribution of
the pharyngeal teeth vary, being, according to Jacket.,
5 — 5; 6 — 5, 1; 1, 5 — 5; or 1, 5 — 5, 2. This hybrid
occurs in Bavaria, but has not yet been found in
Sweden. Another hybrid form, a cross between the
Rudd and the Bleak, is also unknown in Sweden;
but a third, the result of cross-breeding between the
Rudd and the White Bream, has been met with in
this country, and a brief notice of this variety will be
found below.

(Ekstrom, Smitt.)

Genus ASPIUS'.

Scales middle-sized. Lateral line complete. Lower jaw distinctly prominent , with the point fitting into an in-
dentation in the tip of the snout. Caudal lobes pointed. Length of the base of the anal fin more than 19 % of
the distance between this fin and the tip of the snout. Beginning of the do'rsal fin situated in front of the
middle of the body, and the distance between it and the tip of the snout less than 86 % of that between the

anal fin and the same point.

Thus defined, the genus contains only one spe-
cies0, and is ranged by its flatly convex belly (only
slightly carinated between the ventral and anal fins)
beside the most typical Leuciscines, but owing to
the comparatively great length of the anal fin forms
a distinct link between the Leuciscines and the Abra-

midines. The character by which Gunther distin-
guished this genus from Alburnus, the short and
scattered gill-rakers, as well as the wide gape, is an
expression of its more predatory nature, a point in
which this genus is unequalled by any of our other
Cyprinoids-

a Die Fische Bayerns , Abb. Zool. Miner. Ver. Regensburg, 9:tes Heft (1864), p. 64.

6 Agassiz, Mem. Soc. Sc. Nat. Neucli., torn. I (1836), p. 38.

c The Syrian Aspius vorax (Heckel, Russegg. Reise, Th. II, p. 1081, tab. X, fig. 3) has small scales and a short anal fin. The
Chinese Aspius spilurus (Gunth., 4$ at. Brit. Mus., Fish., vol. VII, p. 311) has large scales and an elongated body, more closely resembling
Alburnus. Kessler has described an Aspius erythrostomus from the Caspian and the Sea of Aral, and Jacowlev an Aspius hybridus from
the month of the Volga; but these forms are otherwise unknown to us.

LEUCISCINES.

783

THE ASP (sw. aspen).
AS PIUS RAPAX.

Plate XXXVI, figs. 1 (o*) and 2 (?).

Scales in the lateral line about 66 (62 — 73). Length of the lower jaw about 1/ 10 (10 or 11 %) of that of the
body , greater than the least depth of the tail , and at least 7/i0 — 82 %) of the length of the base of the anal

tin. Length of the upper jaw at least 1 /x 2 (O' 3 — S'3 %) of that of the body and greater than either the breadth
of the interorbital space or the length of the suboperculum, which measurements are about equal to each other.
Pharyngeal teeth set in two rows (3, 5 — 5, 5), hooked at the tip, with smooth, more or less sharp, and not very

distinct masticatory surface.

Fig. 194. Pharyngeal bones and pharyngeal cartilage of Aspius rapax , natural size, a, b, and c as in the preceding figure.

R. hr. 3; D.

7—8’ A' (12)13—14(15)

; P-

— - — ; V. -;
16—17’ 8’

C. x + 1 + 17 + 1 + x ; L. lat. (62)66—73; L. tr. — — 1;

(4)5—7

Vert. 48

49.

Syn. Capito jluviatilis rapax, Gesn., Hist. Anim., lib. IV, Paralip.,
p. 9; Schonev., Ichthyol. Slesv. Holst., p. 30; Marsigl.,
Danab. Pann. Mys., tom. IV, p. 20, tab. VII, fig. 2; Cy-
prinus magnus crassus argenteus; longitudine ad latitudinem
quintupla, Art., Ichthyol., Syn. Pise., p. 8. Cyprinus
maxilla inferiore longiore cum apice elevate, pinna ani ossicu-
lorum quindecim, Art., ibid., Gen. Pise., p. 6; Syn. Pise.,
p. 14; Descr. Spec. Pise. p. 14; Lin., Fn. Suec., ed. I,

p. 121.

Cyprinus Aspius , Lin., Syst. Nat., ed. X, torn. I, p. 325; Bl.,
Fisch. Deutschl., part. I, p. 48, tab. VII; Retz., Fn. Suec.
Lin., p. 359; Nilss., Prodr. Ichthyol. Scand., p. 28; Cuv.,
Val. ( Leuciseus ), Hist. Nat. Poiss ., tom. XVII, p. 265;
Nilss. (. Abramis ), Slcand. Fn., Fisk., p. 334; Widegr. (Cy-
prinus), Landtbr. Akad. Tidskr. 1863, p. 203; Reut., Sundm.
(Aspius), Finl. Fisk., tab. XIX.

Cyprinus rapax, Leske, Ichthyol. Lips, spec., p. 56; Pall.,
Zoogr. Ross. Asiat., tom. Ill, p. 311; Agass. (Aspius), 1. c.;
Kr., Damn. Fislce, vol. Ill, p. 500; Hckl., Kn., Silsswasserf.
Ostr. Mon., p. 142; Dybowski, Cypr. Livl ., p. 173; Sieb.,

a See Und. Betank. m. Forsl. Fiskeristadga 1883 , p. 154.

Silsswasserf. Mitteleur., p. 169; Mgrn, Finl. Fiskfn., p. 44,
Gthr, Cat. Brit, Mus., Fish., vol. VII, p. 310; Coll., Forh.
Vid. Selsk. Chrnia 1874, Tillregsh., p. 184; 1879, No. 1,

p. 96; Bncke, Fisch. Fischer., Fischz. O., W. Preuss., p.
130; Mob., Hcke, Fisch. Osts., p. 120; Lillj., Sv., Norg.
Fn., Fisk., vol. Ill, p. 261.

Aspius Linnei, Malm, Gbgs, Boh. Fn., p. 567.

The Asp is the largest European Cyprinoicl ; and
though it cannot rival the giants of the family in In-
dia, where Catla Buchanani, for example, may attain
a length of at least 18 dm., still it sometimes reaches
in Sweden, according to several corroborative state-
ments", a. weight of about 8 or 9 kgm. and a length
of about 1 m. (9 — 12 dm.). Its ordinary size, as it is
exposed for sale at Stockholm, is, however, about 4 or
5 dm., and its weight about 2 or 3 kgm. An Asp of
5 kgm., which may sometimes be seen, is about 8 dm.
long. An Asp weighing 1 kgm. measures about 45
cm. to the end of the lower caudal lobe.

The body is rather elongated and compressed. In
ordinary cases the greatest depth is less than 1/4

Scandinavian Fishes.

99

784

SCANDINAVIAN FISHES.

(varying between 23 and 25 %), and the greatest thick-
ness at most about 1/9 (varying between 8 and 1 1 V2
%), of the length of the body to the end of the middle
caudal rays; but in gravid females (see the figure) the
former ratio may rise to 2 7 1/2 %. The least depth of
the tail is about 7io (varying between 9 and 1072 %)
of the length of the body.

The back is arched at the occiput, but elsewhere
slopes almost in a straight line, both backwards and
forwards from its apex at the beginning of the dorsal
fin. Just in front of this fin it is very slightly
compressed at the margin, but with this exception it
is convex or even (behind the dorsal fin) flat. The
belly descends from the chin in a regular curve, rather
sharp in gravid females; and the ventral margin is
broad and flatly convex, but furnished between the
ventral fins and the anal aperture 1) in the median
line with a distinct, but low carina, composed of curved
scales of a special type, and 2) at the boundary be-
tween it and the sides with a similar carina, usually
less prominent, but sometimes quite distinct, which runs
back on each side from the outer angle of the insertion
of the ventral fin.

The head is middle-sized, but laterally compressed,
its length being in young specimens somewhat greater,
in old less, than the greatest depth of the body, and
varying between 24 and 22 % of the length of the
latter. The cheeks are flat and perpendicular; the up-
per surface is straight and slightly convex or almost
flat with a gentle slope in a forward direction; and
the breadth of the interorbital space measures in young-
specimens 31 % of the length of the head, but rises in
old to at least 35 % of the same. The eyes are set
high on the sides of the head, and are comparatively
small, being about equal in size to those of the Tench.
They invariably lie in the anterior half of the head,
for the postorbital length thereof measures in young-
specimens about 56 %, in old about 63 % of its entire
length from the tip of the snout. In young specimens
about 16 cm. long the length of the eyes, which is al-
ways somewhat greater than their vertical diameter, is
about 21 — 20 % of that of the head. In specimens
18 — 22 cm. long this proportion has sunk to 18 or
17 %, and in specimens 44 — 66 cm. long the length
of the eyes is only about 14 — 1 2 1/2 % of that of the
head. The length of the snout, which is shallow and

wedge-shaped, tapering to a sharp edge in front, mea-
sures about 28 % (27 — 29 %) of that of the head. The
lines of the mouth are characterized principally by the
sinus at the tip of the snout, into which the more or less
knob-shaped point of the lower jaw fits when the mouth
is closed. On each side of this sinus the labial mar-
gin forms a somewhat arcuate projection, thus giving
to the margin of the upper jaw on each side the elong-
ated S-shape pointed out by Nilsson. The lower jaw
expands in a similar manner on the sides in front;
and the fleshy lips, especially the lower, are tumid at
the corners of the mouth. The intermaxillaries cannot
be protruded beyond the tip of the snout; yet the gape
is large in consequence of the length of the jaws, which
is greater, compared with the size of the head, than
in any other Scandinavian Cyprinoid", the length of
the upper jaw from the tip of the snout being about
38 % (36 — 39 %) of that of the head, and the length
of the lower jaw about 48 % (45 — 51 %) of the same.
There is no free tongue; but the middle part of the
hyoid apparatus in front (the glossohyoid region) is
fleshy and soft. The nostrils are set on a level with
the upper orbital margins, about twice as near to the
eye as to the tip of the snout. The gill-openings are
large, the broad branchiostegal membranes not coalesc-
ing with the isthmus until they reach the perpendicular
from the hind margin of the eyes. Nine or ten short
and scattered gill-rakers, compressed at the tip or with
faint signs of digitation (cf. the Ide), compose the outer
row on the front of the first branchial arch. The outer
margin of the pharyngeals is furnished with 7 or 6
pointed and still shorter gill-rakers. The long pha-
ryngeals (fig. 194) are armed, as mentioned above, with
two rows of simple, but strong teeth, hooked at the
tip. The pseudobranchiae lie high on the inner surface
of the hyomandibular bone, in the hollow on each side
above the ‘carp-tongue’, and in some cases are there-
fore difficult of detection, though well-developed.

All the fins save, in some degree, the ventrals are
distinguished by a more or less incised, pointed form.
The dorsal fin begins at a distance from the tip of the
snout measuring on an average 47 V2 % (46 1/2 — 48 72 %)
of the length of the body. Its base occupies about 7io
(9'3 — 114 %) of the same length, and its height is in
young specimens about 1 8 1/2 %, in old about 16 %, of
the same. The anal fin, which lies further back in the

Compared with the length of the body, however, the jaws .of the Goldfish are equally long.

ASP.

785

females than in the males, begins at a distance from
the tip of the snout measuring 62 — 67 % of the length
of the body. The length of its base is about 1 3 1/2 —
1472 % of that of the body, and its height about 16 —

13 7 2 % of the same, the base being as a rule in the
young less than, in old specimens equal to or greater
than, the height. The base is also as a rule about equal
to the length of the ventral tins, which measures about

14 % of that of the body. The distance between the
ventral fins and the tip of the snout is about 43 — 46
% of the length of the body, being greatest in the fe-
males, where the preabdominal length is thus about 74
of the length of the body, while in the males it is only
about 79 thereof. The postabdominal length, which is
also greater in the females, varies with age between
about 20 and 23 % of the length of the body. The
outer posterior corner of the ventral tins is indeed
pointed, the second simple or the first branched ray
being the longest; but the succeeding rays are only
slightly shorter, and sometimes, especially in the young,
the outer posterior margin of these fins, when expanded,
is somewhat rounded. The pectoral fins are pointed,
with a distinct contraction at the hind margin within
the tip, calling to mind the genus Pelecus; their length
is about 17- or 18 % of that of the body. The caudal
fin with its pointed lobes, the lower somewhat longer
than the upper, is deeply forked, the length of the
middle rays being in young specimens about 9 %, in
old about 7 or 6 %, of that of the body, or in the
former about 38 %, in the latter about 31 %>, of that
of the longest caudal rays.

The scales are of moderate size and of the same
type as in Leuciscus both in form and texture, save that
the concentric strife on the free (hind) part are more
distinct, coarser, and more scattered; and as in Leu-
ciscus, their nucleus is fairly central. Their number in
an oblique transverse row between the ventral fins and
the lateral line is 7 ; but on counting vertically up-
wards we find only 6 longitudinal rows".

The coloration as a whole conforms to the Leu-
ciscine type, varying according to age, season, and en-
vironments. The colours of the young are lighter, of
old specimens darker. The olive green back with its

steely lustre passes towards the sides into a silvery
or brassy hue, and the belly is white. Especially pretty
is the brassy or golden and silvery lustre on the sides of
the head, which are finely punctated with dark green.
The iris is mostly silver white, but the pupil is encircled
by an almost golden yellow (sometimes paler) ring, and
the upper part of the iris displays a brassy tinge with
a shading of dark dots. The dorsal and caudal fins are
of the same colour as the back, though with a stronger
dash of blue (violet) and red, and with dark margins.
The other fins usually shade into red and yellow; but
in the female our figure represents in her spawning-
dress, they were darker. In this dress might also be
traced — though they became more prominent under
the action of the spirits in which the specimen was
preserved — the 8 dark stripes along the sides above
the lateral line, formed by a dark streak along the
middle of each scale’s free surface, and shown in the
above-quoted figure by Marsigli.

The Asp belongs properly to South-eastern Europe,
but is also common in the central parts of this con-
tinent. In Switzerland and Western Europe, even in
Denmark, it is wanting. It is most frequently found,
however, only in large lakes and rivers, but does not
shun the brackish water on the coast, e. g. in the Haffs
of Northern Germany and Lake Dassower (the mouth of
the Trave). In Sweden its range is almost confined to
the Malar valley and the basin of Lake Wener. It
also occurs, however, in the Dal Elf (at least in* Lake
By, a broad on this river in the south-east of Dale-
carlia6), in the Em (Calmar), and, according to Malmc,
in the Helge (Christianstad). In Lake Wetter and the
neighbouring waters it has never been found. That it
also inhabits the Baltic island-belts, is more than pro-
bable, for the large female represented in our figure
was taken at Sjbtullen outside Stockholm. Muller1*
assigned it to the Norwegian fauna under the name of
Blaa-spol; but, according to Collett, the only Nor-
wegian waters inhabited by this species are Lake Ojeren,
from which it ascends in spring the Lersundselv and
Nitelv, and that part of the Glommenelv which is below
Lake Ojeren. In Finland the Asp is known with cer-
tainty only from the extreme south-west, but according

a Sometimes only 4, according to Dybowski.

b According to information received from Dr. Steffenburg by Lilljeborg. See also the above-cited report of the Swedish Fisheries
Commission.

c See also Retzius, Fn. Saec. Lin.
d Zool. Dan. Prodr., p. 51.

786

SCANDINAVIAN FISHES.

to Mela, occurs, though seldom, in the vicinity of
Kuopio.

The Asp, as we have mentioned, is a fish of marked
predatory habits, resorting to vegetable food only by
accident, or when reduced by necessity. It is there-
fore lacking in the sociability of our other Cyprinoids,
a feeling it evinces only in the spawning-season and
perhaps during its winter sleep. “His name” (Germ.
Proppe, Lat. rapax), writes Gesner, “is derived from
his voracity, for he ranges the waters like a formidable
pirate, to other fishes a no less dangerous, but rather a
more destructive foe than the Sheat-fish or the Pike.
In headlong chase of his flying victims, which in terror
leap on dry land, he sometimes runs ashore himself.”
Bleak and Smelt are its commonest prey; but it does
not shrink from assailing larger fish or even water-
rats, nor does it disdain smaller and lower animals,
such as worms, mollusks, and the like. It is not very
tenacious of life, dying soon on dry land. Its favourite
haunts are clear lakes or gentle streams with clean,
sandy or gravelly bottom. As long as the water is
clear, it cunningly avoids all snares, and is difficult to
take with net or seine; but a Minnow set on a trolling
hook and cast enticingly before its nose is too tempting
a bait to be resisted. When the water is thick, it may
be more easily netted. Such are its habits in summer;
but they assume a different aspect under the influence
of sexual excitement.

The spawning-season of the Asp begins early in
spring, soon after the breaking up of the ice, in Swe-
den in April or May. The males are then marked by
the usual dermal eruption of small round tubercles on

the head, the pectoral fins, and the dorsal scales back
to the tail. The fish assemble in shoals, which ascend
the rivers, or proceed to shallow parts of the lakes,
where the roe is attached to stones or weeds, or simply
deposited on the bottom. A middle-sized female con-
tains, according to Benecke, 80,000 — 100,000 eggs.
The large gravid female of which we give a figure,
measured 75 cm. from the tip of the snout to the end
of the caudal lobes, and when taken off Stockholm on
the 22nd of April, 1886, had nearly the whole of the
abdominal cavity under the air-bladder and forward to
the diaphragm filled by the two ovaries with their three
or four lobes. The eggs were about l2/3 mm. in dia-
meter, and their number was computed to be about
300,000. Of the growth of the fry Norback states'2
that during the first year they attain a length of 9 cm.,
and Lilljeborg assumes specimens 15 cm. long, taken
at the beginning of May, to be one year old.

The flesh of the Asp is white and fat, but bony
and difficult of digestion. It shows a tendency in pro-
cess of boiling to separate into flakes (the muscular
sections, myomeres ), which may be obviated, however,
by putting it on the fire in cold water. Large and
fleshy as the fish is, it is one of the most important
Cyprinoids, and the head is considered a delicacy by
many. Except during the spawning-season, however,
no large catches of Asp are made in Sweden. Solitary
specimens appear in the fishmarkets of Stockholm about
Christmas; but the true season for Asp is from Febru-
ary to June. The Asp is taken chiefly in nets or traps.
Still it affords good sport to the angler, who should pre-
fer a bait of live fish, though worms may also be used.

Genus LEUCASPIUS'.

Scales middle-sized and deciduous. Lateral line incomplete. Lower jaw distinctly projecting , with the point fitting
into a shallow indentation at the tip of the snout. Lobes of the caudal fin pointed. Length of the base of the
anal fin more than 10 % of the distance between this fin and the tip of the snout , and more than P/2 times the
least depth of the tail. Beginning of the dorsal fin situated at the middle of the body or farther back, and the
distance between it and the tip of the snout more them 86 % of that between the anal fin and the same point.

An intermediate form between Aspius and Albur-
nus, Leucaspius is nearly allied to the following sub-
family. One of the most unmistakable signs of this is

the relative position of the dorsal tin to the anal. In
none of the preceding Leuciscines — except in young
males of Phoxinus — have we found the distance between

a Handl. Fislcev., F/skafu., p. 439.
6 Hckl, Kn., 1. c.. p. 145.

OWSIANKA.

787

the dorsal tin and the tip of the snout to measure more
than 86 % ol that between the anal tin and the same
point, as is generally the case in the Bleak, Bream,
White Bream, and “Ziege”. The fry of the Rudd and a
few males of the same species, however, approach the line

of demarcation very closely; but Leiicaspius is easily dis-
tinguished from all the Leuciscines that have scales of
moderate or some considerable size, by its incomplete
lateral line, which seldom extends behind the tip of the
pectoral tins. Only one species of the genus is known.

THE OWSIANKA (sw. groplojan «).
LEUCASPIUS DELINEATUS.

Fig. 195.

Coloration silvery like
slender, hooked

that of the Bleak, with a steel-blue band along the sides of the body. Pharyngeal teeth
at the tip, pectinated , and, set in one or two rows: 5 — 4(5) or 1(2), 5 — 4, 1(2).

’ *-■ w

a b

Fig. 195. Leucaspius deline atus , natural size, taken at Landskrona on the 4th August, 1871 by Lilljeborg, together with the left

pharyngeals magn. 5 diam. a and b as in the preceding figure.

R. hr.

D. 1;
8

A.

11—13'

P.

12 — 14’

V.

1(2)

C. x + 1 + (16)17 + 1 +x; L. lat. sqv. 44 — 50 (7 — 13 perfor.);
L. tr. 12 — 13; Vert. 36 c.

Syn. Aphya ( Mutterloseken ), Schonev., Ichthyol. Slesv. Holst., p. 16.
Squalius delineatus, Hckl, Rassegyers Reisen, vol. I, pt. 2,
p. 1041; Hckl, Kn., Siisswasserf. Ustr. Mon., p. 193; Sieb.
( Leucaspius ), Siisswasserf. Mitteleur., p. 171; Gthr, Cat.
Brit. Mus., Fish., vol. VII, p. 319; Lillj., Ofvers. Vet.-
Akad. Fork. 1871, p. 815, tab XVII, A; Malm, Obeys , Boh.
Fn ., p. 568, not.; Fedders., Naturh. Tidskr. Kbhvn, ser. 3,
vol. XII, p. 90; Bncke, Fiscli., Fischer., Fiscliz. 0 ., W.
Preuss., p. 131.

Leuciscus stymphalicus , Cuv., Val., Hist. Nat. Poiss., vol.
XVII, p. 295, tab. 498.

Aspius owsianka, Czernay, Bull. Soc. Natur. Mosc., tom.
XXIV, part. 1, p. 281, tab. VII; pt. 2, p. 259; Maslow.,
ibid., tom. XXVII, pt. 1, p. 442.

Leucaspius abruptus, Hckl, Kn., 1. c., p. 145; Dyb., Cypr.,
Livl., p. 147 + Owsianka Czernayi, ibid, et p. 140.

The Owsianka is one of the smallest Cypri-
noids. Its usual length is about 6 — 8 cm., measured
to the tip of the caudal lobes, the former in the males,
the latter in the females. Sometimes, however, ac-
cording to Benecke, it may attain a length of 12 cm.
Its size is thus equal as a rule to that of the Minnow,
but the form of its body is quite different, being more
compressed, like that of the Bleak, which it also re-
sembles in coloration.

The body is well proportioned, with regular dorsal
and ventral curves, the latter being the sharper. The
greatest depth of the body is about 20 — 22 % of its

a Lilljeborg, 1. c.

6 Sometimes 14, according to Dybowski.
c According to Maslowsky.

788

SCANDINAVIAN FISHES.

length to the end of the middle caudal rays, and the
greatest thickness is about half the greatest depth. The
least depth is about 8 % (7 '8 — 8'2 %) of the length.
The back is broad and convex, the belly more com-
pressed, forming between the ventral fins and the anal
aperture a sharp carina, covered with a row of curved
scales.

The length of the head is about 23 — 21 % of that
of the body. It is apparently a general rule that in
the young and the males the length of the head is
greater, in adult females less, than the greatest depth
of the body. In form the head is almost exactly si-
milar to that of the Bleak, but in old specimens its
upper surface, along the middle of the forehead and
the crown, is depressed and plane. The eyes are ver-
tically set, rather large, and situated almost entirely in
the anterior half of the head, the postorbital part mea-
suring about 43 or 44 % of the entire length of the
head. In specimens 5 — 8 cm. long the longitudinal
diameter of the eyes, which is slightly greater than or
equal to the vertical diameter, measures about 33 — 31
% of the length of the head, and is always perceptibly
greater than the length of the snout. The tip of the
snout is sharp (shallow), but broad (truncate), with a
shallow sinus to receive the point of the lower jaw.
The cleft of the mouth is turned sharply upwards and
rather large; but in consequence of its obliquity the
hind extremity of the maxillaries scarcely extends, when
the mouth is closed, to the perpendicular from the an-
terior margin of the eyes. The lips are thin. The
length of the upper jaw from the middle of the tip of
the snout is about 7 % (6*6 — 7*2 %) of that of the body,
or about 30 % (29‘2 — 32‘2 °/o) of that of the head. The
length of the lower jaw, which is generally about equal
both to the breadth of the interorbital space and the
length of the suture between the suboperculum and the
operculum, measures about 8% % (8’5 — 8‘7 %) of the
length of the body, or about 38 1/2 % (37'5 — 40 %) of
that of the head. The nostrils lie rather near the up-
per anterior corner of the orbits. The gill-openings
are fairly large, the branchiostegal membranes coalesc-
ing with the isthmus in about a line with the hind
margin of the preoperculum. The outer row on the
front of the first branchial arch contains 11 — 13 pointed
gill-rakers, small and close-set below, larger and more
scattered above. The pharyngeal teeth are slender and
almost straight, with hooked tip and pectinated masti-
catory surface. They are sometimes set, according to

Heckel, Maslowsky, and Siebold, in two rows; but
in all the specimens examined bv us they formed a
single row. The pharyngeal cartilage is elliptical, but
its hind (lower) extremity is raised, as in the preceding
Leuciscines, in an inverted canaliculate form.

The fins are of the normal Leuciscine type. The
dorsal fin begins at a distance from the tip of the snout
measuring about half (49 — 52 %) of the length of the
body. The length of its base is about 7g (12—11 %),
and its height about 1/7 (16 — 1 3 3/4 %), of the same length.
The anal fin is perceptibly longer and, especially in
old specimens, lower, and begins further back in the
females than in the males. The distance between it
and the tip of the snout measures in the males about
55 or 56 in the females about 57V2 — 5972 of the
length of the body. Its base is about 16—14 % (in
exceptional cases 13 %), and its height in young spe-
cimens about 14 %, in old 13 — 12 %, of the same length.
Its length marks the approximation to the Abramidines,
for whereas in all the preceding Cyprinoids the least
depth of the tail is more than 2/3 of the length of the
base of the anal fin — though exceptions may occur
among young Minnows — in Leucaspius delineatus the
said ratio is less than 2/3 (about 50 — 61 %). The cau-
dal fin undergoes a similar change of growth, the length
of the middle caudal rays being in young specimens
about 12 %, in old about 10 — 9x/2 %, of that of the
body. The longest caudal rays (in the inferior lobe)
measure about 22 or 23 % of the length of the body.
This fin is also subject to individual variation, an ex-
ceptional circumstance among the Cyprinoids, the num-
ber of the branched rays being either 16 or 17. The
pectoral fins are obliquely pointed and comparatively
short (16 or 15 % of the length of the body). The
ventral fins are almost triangular, and their length is
about 14 or 13 % (in exceptional cases 11 %), of that
of the body. The distance between the ventral fins
and the tip of the snout measures about 41 — 45 %, the
preabdominal length about 20 — 22 V2 %, and the post-
abdominal length about 14 or 15 % (cf) — 17 or 18 %
($), of the length of the body.

The anal aperture calls to mind the corresponding
organ in the Minnow and the ‘Bitterling’ (see above).
It is more prominent and tubiform than in the Minnow,
especially in the females, and is furnished on each side
with an oblong, compressed papilla.

The scales are thin, deciduous, and rather large,
their number in an oblique transverse row above the

OWSIANKA.

789

ventral fins being only 11 — 13. In form and structure
they show most resemblance to those of the common
Bleak, being deeper than long (of broad elliptical shape,
with longitudinal axis set across the body), with few
and indistinct radiating grooves, and with the nucleus
nearer to the anterior margin than to the posterior.
In young specimens only feAv scales (3 — 4 or even none)
are pierced by the lateral line as it descends in a curve
from the temporal region, while in old specimens this
number is generally between 7 and 13.

The coloration too is mainly that of Alburnus. The
dorsal side is olive green, more or less dark (brownish).
The sides of the body are of a silvery lustre, with a
steel-blue band, over which the silvery lustre extends,
from the upper part of the gill-openings to the middle
of the base of the caudal fin, towards which point it
grows more distinct. The scales are dotted with brown,
especially on the upper part of the sides in front.
The fins are transparent and almost colourless, the
dorsal and anal shading into grayish green, the pec-
torals into grayish white, the ventrals and anal into
faint yellow.

The Owsianka, as we have mentioned, is one
of Lilljeborg’s discoveries in the Scandinavian fauna.
In 1871 his attention was directed by Mr. Aiilbom, late
Collector in the Customs’ Department, to a “variety of
Bleak” that inhabited a small, but deep pond with peaty
bottom, in a field near Landskrona. The fish would
occasionally seem to have vanished from the pool, but
usually re-appeared in great numbers during the spring,
in the month of May. Lilljeborg recognised this Bleak
as Leucaspius ( Squalius ) delineatus, a form described
first by Heckel; and since then it has been met with
in many other peat-haggs in Southern Scania between
Landskrona and Ystad. But in Sweden, as in its true
habitat, the species has been found in running water,
even before Lilljeborg’s discovery, though it was not
correctly determined until then. Malm states in Gbgs,
Boh. Fn. that in September, 1868 he found it in the
Kjeflinge near the railway-station of < Irtofta, where

it kept to shallow water near the grassy bank of the
river; and according to Trybom it occurs both in
broads along the course of the same stream and in Lake
Vomb, the waters of which are discharged by this
river, being so plentiful that it is often used as bait.
In Denmark Fiedler and Feddersen have taken the
species in small pools on the island of Zealand.

In Germany, as Siebold has pointed out, the
Owsianka has long been known under the names
of MutterloseJcen (motherless), Moderliesken, etc. Its
sudden appearance in peat-haggs and other small col-
lections of water, which excited Ahlbom’s attention in
Sweden, had given rise in Schonevelde’s time to the
belief that it came into being without parents ( apliya ).
In Germany too it has been found both in peat-haggs
and small streams. Blasius met with numerous spe-
cimens at Brunswick; Benecke states that it occurs in
the Kurische Haff. The French Expedition to the Mo-
rea found the species in Lake Za-raco, the famed Styin-
phalian Lake of Greek mythology. It- seems to be most
common, however, in Southern Russia, where it bears
the name of owsianka , and according to Czerxay, is
used as food, in spite of its small size, and considered
fairly good eating.

The Owsianka, like the great majority of our
Cyprinoids, spawns in spring, the usual month in
Sweden being May, in Germany April. It is a lively
fish, in temperament resembling the Bleak, and also
feeding on small insects. It dies soon after it is taken
out of the water. Its apparently periodical disappear-
ance from the peat-haggs — in Southern Russia too,
according to Maslowsky, it is caught in the small
streams only from September to April — may probably
be explained either by some migration after the spawn-
ing-season, should any egress be open to the fish, or
on the assumption that at certain seasons it keeps to
the bottom of the deep pools, appearing at the surface
when the water is disturbed. In Sweden it is some-
times fried and made into ‘fish-cake’ like other small
fishes, but this is the only form in which it is eaten.

790

SCANDINAVIAN FISHES.

Subfamily ABRAMIDIN JE.

Dorsal fin much shorter than the anal, the base of the former measuring as a rule less than */3a of that of the
latter , which is more than 16 % of the length of the body. Length of the head less than half the distance be-
tween the dorsal fin and the tip of the snout. No spinous ray in the dorsal or the anal fin. Ventral margin be-
tween the ventral fins and the anal aperture sharp , but generally naked ( not covered with curved scales ) in the
median line. Mouth without barbels. Length of the lower jaw as a ride less than 47 % of that of the base of the
anal fin. Pharyngeal cartilage oval or elliptical, with the anterior {upper) extremity more or less obtusely pointed.

These characters are accompanied by the well-
known Abramidine form, a deep and compressed body
with strongly compressed or even sharp margins, along
which the scales are shed at certain parts of the me-
dian line, the two outermost rows of scales being thus
juxtaposed edge to edge, or leaving the skin at these
parts naked. This is always the case, even in Alburnus
and, at least partially, in Spirlinus, along the ventral
margin between the ventral tins and the anal aperture;
and in the true Breams the 'denudation extends, though
with varying distinctness, to the anterior part of the
dorsal margin behind the occiput.

It was not without reason that Nilsson proposed
to unite all the Scandinavian Abramidines into one
genus. Great as the difference may appear between a,
Bleak and a Bream, we find intermediate forms be-
tween them composing an almost unbroken series. A
species found in Denmark and further south, Spirlinus
bipunctatus, has also been referred by some, for ex-
ample by Gunther, to the genus Abramis, by others, for
example Heckel and Siebold, to Alburnus. The near
relationship of this subfamily to the preceding one, a re-
lationship which in its intermediate forms arid hybrids
defies every attempt by fixed characters to define the li-
mits between these two groups, has caused a like diver-
sity of opinion, the Bleak being referred by Agassiz,
Kroyer, Nilsson, and Lilljeborg to the same genus as
the Asp. If the systematic classification of forms so

closely allied is to be based on natural grounds, we are
compelled for the sake of consistency to employ as ge-
neric characters relations which elsewhere seem to be of
comparatively little weight. Hence Fatio established his
new genus Spirlinus, and for the same reason we are
obliged to divide the genus Abramis, with a view to
obtaining an expression for the points of resemblance
between the ‘Zarthe’ and preceding forms and the
respects in which it differs widely from the true
Breams.

The genera belonging to the Scandinavian fauna
may be distinguished as follows:

I: Length of the head more than 1 / 3 of
the distance between the dorsal fin and
the tip of the snout.

A: Base of the anal fin less than 1/5 of
the length of the body and shorter
than the head.

a: Lower jaw most prominent. Pec-
toral fins longer than the longest

ray in the dorsal fin Genus Alburnus.

b: Jaws about equally prominent.

Pectoral fins shorter than the
longest ray in the dorsal fin ... Genus Spirlinus.
c: Tip of the snout most prominent.

Pectoral fins shorter than the

longest ray in the dorsal fin ... Genus Leucabramis.
B: Base of the anal fin more than Vs
of the length of the body and as
a rule longer than* 6 the head Genus Abramis.

II: Length of the head less than 1/3 of the
distance between the dorsal fin and the
tip of the snout Genus Pelecus.

a Invariably less than 72 % in all the specimens measured by us.

6 In White Bream and young Bream exceptionally equal in length to the head.

ABRAMIDINES.

791

Genus ALBURNUS.

of the dorsal fin situated at a distance from the tip of the snout less than three times the length of
Length of the base of the anal fin less than 1/5 {between 16 and 18 %) of that of the body , and also
Pectoral fins longer than the longest ray in the dorsal fin. Point of the lower jaw

Scales thin and deciduous; nucleus situated in
the anterior half of the scale.

the body — we find that in this respect the Bleaks and

Beginning
the head.

less than that of the head.

projecting distinctly beyond the tip of the snout.

The genus of the Bleaks, according to Gunther’s
Cat (dog lie, contains 15 recognised species from Europe
and Southwestern Asia, most of them described by
Heckel from the latter region, which thus seems to
be the true home of the genus. They are small, but
lively fishes, readily attracting attention in rivers and
lakes, where they sport at the surface in chase of in-
sects, or in eager contest for every breadcrumb thrown
to them.

They are distinguished from the other genera of
the subfamily, with the exception of the “Ziege”, by
their comparatively low dorsal fin, generally lower even
than in the preceding subfamily, excluding the Minnow.
But on comparing the height of the dorsal fin with
the length of the pectorals — a relation which is not
without importance in preserving the equilibrium of

the Ziege rank with the adult Leuciscines, except the
Roach, where, as in young specimens of the Scandi-
navian Leuciscines in general, the length of the pec-
toral fins is less than that of the longest ray in the
dorsal fin. The majority of the Scandinavian Abrami-
dines, on the other hand, thus retain, like the Roach,
in this respect one of the characters of youth among
the Leuciscines.

The name of the genus dates even from the time
of Ausonius“. Linnaeus employed it in a specific sig-
nification; but Heckel restored it to the generic rank
it had occupied in Rondelet* * * 6, and separated0 the
Bleaks under this name from the other Cyprinoid
genera. Rondelet’s Alburnus, however, was evidently
distinct from this genus.

a The continuation of the lines quoted above (p. 751) from the Mosella of Ausonius runs:

“Et Alburnos pra?dam puerilibus hamis?”

“And Bleak, an easy catch for angling boys.’’

6 De Pise., part. II, p. 208.
c Russeggers Reisen , 1 Bd., 2 Th., p. 1036.

Scandinavian Fishes.

100

792

SCANDINAVIAN FISHES.

THE BLEAK ( SW. LOJAN ).

ALBURNUS LUCID US.

Plate XXXVI, fig. 3.

Scales in the lateral line about 50 (46 — 54), branched rays in the anal fin about 17 (16- — 20). Sides of the
body of a plain, lustrous, silvery white. Pharyngeal teeth slender, hooked at the tip, pectinated, and set in two

rows: 2, 5 — (4)5, 2.

a

Fig-. 196. Left lower pharyngeals of Alburnus lucidus, a, seen from above, b, from without, both figures twice the natural size;

C, the middle tooth in the inner row, three times the natural size.

3 3 12

R. hr. 3; D. — ; A. • P. ; V. —

8—9“ 16 — 20* 6 14 — 15c 8d

O.x + 1 + 17+1 +x; Lin. lat. 46 — 54; L. tv. el ; Vert. 42-1 — 44.

3(4)

Syn. Albula minor , Schonev., Ichtliyol. Slesv. Hols., p. 11. Cyprinus
quincuneialis, pinna ani ossiculorum viginti, Art., Ichtliyol.,
Gen. Pise., p. 6; Syn. Pise., p. 10; Descr. Spec. Pise., p.
17; Lin., Fn. Snec., ed. 1, p. 124.

Cyprinus Alburnus, Lin., Syst. Nat., ed. X, tom. I, p. 325;
Bl., Fisch. Deutschl., pt. I, p. 54, tab. VIII, fig. 4; Retz.,
Fn. Suec. Lin., p. 359; Pall., Zoogr. Ross. Asiat., tom. Ill,
p. 321; Ekstr. (subg. Abramis , ex. Cuv.), Vet.-Akad. Handl.
1830, p. 187; Nilss., Prodr. Ichtliyol. Scand., p. 31; Agass.
(Aspius), Mem. Soc. Sc. Nat. Neuch., tom. I, p. 38; Scha-
gerstr. {Cyprinus), Physiogr. Sall'sk. Tidskr. 1837, p. 295;
Cuv., Val. ( Leaciscus ), Hist. Nat. Poiss., tom. XVII, p. 272
(+ Leac. ochrodon (ex. Agass.), p. 249 + L. alburnoides
(ex Selys), p. 250); Fr., Ekstr., v. We. ( Cyprinus , subg.
Aspius), Skand. Fisk., ed. I, pp. 58 et 203, tab. 51; Kr.
(Aspius), Danm. Fisk., vol. 3, p. 485; Nilss. ( Abramis ,
subg. Aspius ), Skand. Fn., Fisk., p. 337 ; Sundev. (Cyprinus),
Stockh. L. Hush. Sallsk. Handl. 1855, p. 81; Widegr., Fiskfn.,
Fisker. Norrb. L. (1860), pp. 7 et 13; Lindstr. (Abramis),
Gotl. Fisk., Gotl. L. Hush. Sallsk. Arsber. 1866, p. 18 (sep.);
Olss. (Abramis), Of vers. Vet.-Akad. Forh. 1876, No. 3, p.
131; Fedders. (Aspius), Naturh. Tidskr., Kbhvn, ser. 3, vol.
XII, p. 90; Reut., Sundm., Finl. Fisk., tab. XIV; Lillj.,
So., Norg. Fn., Fisk., vol. Ill, p. 253.

Alburnus lucidus, IIckl (+ Alb. obtusus + A. acutus), Russegg.
Reis., 1. c.; Hckl, Kn., Siisswasserf. Ostr. Mon., p. 131
(+ Alb. breuiceps, p. 134); Dybowski, Cypr. Livl., p. 165;

Sieb., Siisswasserf. Mitteleur., p. 154; Blanch., Poiss. d.
eaux douces Fr., p. 364 (+ Alb. mirandella, p. 369 + A.
Fabrcei, p. 370); Gthr, Cat. Brit. Mus., Fish., vol. VII,
p. 312; Coll., Forh. Vid. Selsk., Chrnia 1874, Tillsegsh.,
p. 184; Mor., Hist. Nat. Poiss. Fr., tom. Ill, p. 403;
Bncke, Fiscli., Fischer., Fiscliz. O., W. Preuss., p. 127 ;
Mela, Vert. Fenn., p. 336, tab. X; Day, Fisli. Gt. Brit.,
Irel., vol. II, p. 198, tab. CXXXVIT, fig. 1; Mob., Hcke,
Fisch. Osts., p. 119; Fat., Fne Vert. Suisse, vol. IV, p. 414.

Alburnus Linnei, Malm, Gbgs, Boh. Fn., p. 568.

The Bleak is one of the smaller Cyprinoids. Its
ordinary length, including the caudal tin, is 13 — 15
cm. In Sweden it seldom measures 18 cm.; but Be-
necke states that in Prussia it attains a length of 20 cm3

The body is rather elongated and laterally com-
pressed, the greatest depth being occasionally 1/i, but
generally between 1 8x/2 and 23 %, of the length to the
end of the middle caudal rays, and the greatest thick-
ness at most about Vs the greatest depth. The least
depth of the tail measures on an average about 8 %
(7*4— 8*2 %) of the length of the body. The dorsal
margin is convex throughout its length. The belly is
flat in front, carinated from the ventral tins to the
anal aperture. The dorsal line is often nearly straight,
being, at least in front, much less curved than the
ventral.

“ 7 — 9, according to Fatio.

6 Sometimes 15, according to Fatio.

e according to Fatio.

3—4

4 Sometimes 41, according to Fatio.

g According to Bloch the Prussian Bleak may attain a length of 8 — 10 in. (20 — 26 cm.).

BLEAK.

793

The length of the head varies between 19 and 21
% of that of the body, its size being thus somewhat
below the average among the Scandinavian Cyprinoids.
It is rather pointed, of uniform thickness, and com-
pressed, at the top convex, with fairly parallel cheeks.
The facial line is straight from the occiput to the tip
of the snout. The breadth of the interorbital space
measures about 3/10 °f the head, and is generally some-
what greater than the longitudinal diameter of the eyes,
but sometimes equal to this diameter, which also differs
only slightly from the length of the snout. The nostrils
lie on a level with the upper margin of the eyes,
nearer to the eyes than to the tip of the snout; and
the distance between the anterior nostrils is equal to
that between either of them and the middle of the tip
of the snout. As usual in this family, the two nostrils
of each side are separated by a narrow dermal ridge,
raised in a lobate form, and the shape of the nostrils
varies with the position of this ridge: when it is thrown
back, the anterior is round, the posterior crescent-
shaped. The situation of the eyes is such that the
postorbital length of the head is as a rule somewhat
less, but sometimes rather more, than half (44 — 52 %)
of its entire length. The mouth is turned sharply up-
wards, its entire cleft lying in front of the perpendi-
cular from the nostrils, which line, when the mouth
is closed, touches the hind extremity of the maxillary
bones. A small notch in the sharp tip of the snout,
which is formed by the intermaxillaries (the margin
of the upper lip), receives the blunt and rather pro-
minent point of the lower jaw. The length of the
upper jaw from the middle of the tip of the snout is
often almost equal to the length of the snout, measur-
ing 29 — 26 % of that of the head. The length of the
lower jaw is 35 — 40 % of that of the head. The gill-
openings are fairly large, the branchiostegal membranes
coalescing with the isthmus almost beside each other,
a little in front of the perpendicular from the hind
margin of the preoperculum. The gill-rakers are close-
set: in the outer row on the front of the branchial
arch (16 or 17 in number) they are slender and wand-

shaped, while in the inner row on this arch and in
both rows on the other arches' and on the outer an-
terior margin of the pharyngeals they are short and
of a pointed triangular form. The pseudobranchiae are
well developed. The pharyngeal teeth are distinguished,
here as in the Rudd, by the pectination of the masti-
catory surface, at least before it is worn smooth, in
the case of the three or four posterior teeth in the
inner row. The two anterior teeth or at least. the first
tooth in this row, are as usual more conical, the se-
cond with or without pectinated masticatory surface,
or with this surface worn smooth. Such is also the
form of the two small teeth in the outer row. The
pharyngeal cartilage is oval, with grooves obliquely
crossing the masticatory surface, as in the majority of
the Leuciscines described above, and with the hind
(lower) extremity somewhat raised, a trace of the in-
verted canaliculate form more highly developed in the
larger Leuciscines.

The dorsal fin begins at about the middle of the
length of the body" or a little further back. It is of
almost the same trapezoidal form as in the Leuciscines,
with the length of the last ray at least rather more
than V3 of that of the longest ray, which here seems
hardly to exceed 15 %h of the length of the body.
The base of the dorsal fin measures about 7 nc of the
length of the body. The anal fin begins below the
posterior part of the dorsal L It is long*", low behind,
and forms a slight arch at the concave inferior mar-
gin. The first ray is hardly perceptible, the second
half as long as the third, the fourth the longest/ and,
like the following rays, branched at the tip. The
caudal fin is deeply forked, the lower lobe being some-
what longer than the upper, and measuring somewhat
more than 1/5 of the length of the body.

The pectoral fins are obliquely and bluntly pointed.
Their length is about 1/6 (16 1/2 — 1772 %) of that of
the body. The ventral fins are much shorter, measur-
ing about 12 — 13 % of the length of the body. Their
position is such that the distance from the tip of the
snout to their insertion6' does not seem to exceed

“ At a distance from the tip of the snout measuring 50 — 521/'2 of the length of the body; the latter percentage in our oldest fe-
male specimens.

b Varying, according to our measurements, between 1 31/, and 14 *U #■
c According to our measurements 8'7 — 9'7 %.

d At a distance from the tip of the snout measuring about 59 % (57'4 — 59'3 %) of the length of the body.
e The base of the anal fin measures about 17 % (16'4 — 1 7 ' 6 %) of the length of the body.
f About 11 / (11*8 — 10'5 %) of the length of the body.
g 40 — 43 % of the length of the body.

794

SCANDINAVIAN FISHES.

8 2 V2 “ of that from the same point to the dorsal fin.
The preabdominal length measures on an average 22 V2
%, and the postabdominal length 18V2 %■> of the length
of the body.

The texture of the thin scales has already been
remarked. It is fairly characteristic of the Bleaks and
their nearest relatives, in the dense, concentric striae,
most dense in the anterior (covered) part of the scale,
around the eccentric nucleus, which also lies in the
covered part. Towards the nucleus run a few (on the
scales of the sides commonly 4, on the dorsal scales
as many as 10) scattered grooves, extending forward
for a greater or less distance from the hind margin
of the scale. The scales are generally of a broad
elliptical shape (those of the lateral line more quadri-
lateral, with the posterior part rounded), the longi-
tudinal axis of the ellipse pointing up and down. They
are rather large, but deciduous. The inner surface of
their posterior part is lined with a dense layer of long,
almost filiform crystals, which give them their silvery
lustre.

The colour of the back and the upper part of the
head is greenish gray. The rest of the body is silvery.
The iris is also silvery, but above the pupil yellowish,
with dense, confluent spots of gray. The pectoral,
ventral, and anal fins are white, the first pair faintly
tinged with green at the anterior margin. The dorsal
and caudal fins are gray.

The Bleak is spread over the whole of Europe
north of the Alps, with the exception of Scotland and
Ireland6. In Sweden it is common enough, under the
names of Loja, Loga , Benloja, Pjon , Pyn, etc., up to
the neighbourhood of Quickjock0 (lat. 67° N.). It does
not seem to thrive, however, in the highlands, for it
is rare in Lapland, and in Western Jemtland, according
to Olsson, it is wanting16. In Finland, where it is
called SalakJca, a name which at Tornea is altered,
says Widegren, to Salk, its range extends, according
to Mela, to 68° 20' N. lat. In Norway, a country
not very rich in Cyprinoids, the Bleak is found
only in the south-east, no further north, accord-
ing to Collett, than Lake Mjosen. It lives in the
great lakes, in clear rivers and streams, and also in

“ According to our measurements 79 — 82'2 /.

the inner part of the Baltic island-belt, as well as in
Gothland and round the coasts of that island. Accord-
ing to Schagekstrom it also occurs in the Sound, at
least off Landskrona.

The Bleak lives in shoals, solitary specimens being
never met with, and prefers clear, running water with
a stony or sandy bottom. It is consequently found
but seldom in small lakes with a bottom of weeds or
mud. Sportive and lively in temperament, it always
keeps to the surface from spring to autumn, and
catches the insects that drop into the water. It loves
sunshine and calm, and in rainy and stormy weather
makes for sheltered shores. At such times it probably
descends some way below the surface, but it does not
stay there long, and is never found at the bottom,
except in autumn, when, like the other Cyprinoids, it
retires to its winter-quarters in the depths. Less shy
than timid, it soon returns to the spot whence it has
been driven away by some noise. On the other hand,
it is voracious, and instantly seizes any small sub-
stance thrown into the water. If it finds the morsel
unfit for food, it again disgorges its prize. Its food
consists principally of insects, small crustaceans, and
worms.

At the end of May or beginning of June the
Bleak assembles in large shoals to spawn in shallow
water with a stony or sandy bottom. The shoal presses
in serried array close to the shore, and seems like a
dark cloud in the water. The spawning now begins;
the fish leap time after time, at brief intervals, above
the surface, and meanwhile deal frequent and rapid
blows with the tail on the water, thus producing a
hissing sound like that heard when a piece of cloth is
suddenly torn in two. The roe attaches itself to stones
or twigs at the bottom. How soon the eggs are hatched
under ordinary circumstances, we cannot state with
certainty, the time allotted by different statements to
this process varying between 24 hours and as many
days; but it undoubtedly varies according to the tem-
perature of the water. Of the growth of the fry we
learn from Malm that in the middle of September he
caught young Bleak 18 — 23 mm. long in the Hoje
(a stream in Scania); and in Lake Millar Lilljeborg

411. The statement is, however, somewhat dubious, for the spawning-

b Day, 1. c.

c According to LSwenhjelm, Vet.:Akad. Handl. 1843, p.
season is said to occur in September.

d According to Fatio it ascends in the Swiss lakes to a height of about 700 m. above the level of the sea.

BLEAK.

795

took specimens 20 — 25 mm. long at the beginning of
October. Fishermen in general believe that the Bleak
spawns three times a year at short intervals; and from

o

Lomma (Scania) we are told by Astrom" that at Whit-
suntide three shoals of Bleak generally enter the stream
from the Sound, and spawn one after the other at
intervals of about a week. But the true explanation
of this fact is that the difference in the spawning-season
depends on the different ages of the fish, for the Bleak
that spawn first are always larger than those which
spawn later. Bleak are taken in numbers only during
the spawning-season. The tackle most employed con-
sists of a small and fine-meshed seine, constructed
especially for this purpose, and called lojnot or log skate
(Bleak-seine or Bleak-net), or a large circle-net (Sw.
grip), which is cast over the shoal while spawning.
In summer the Bleak may also be taken in small-
meshed nets and in traps; but the catch is seldom
large. After the spawning-season it takes a bait freely,
a fly being an especially tempting morsel.

As the Bleak is small, and is only seldom taken
in large numbers, it cannot possess any great impor-
tance as an article of food. It is generally eaten fresh,
its flavour, when fried, being not unlike that of the
Baltic Herring. When salted or dried it entirely loses
its flavour. It is most useful to the fisherman as bait,
for which purpose it is excellent, though not as live
bait, its tenacity of life being small. It is eagerly
sought after by terns and gulls, which generally flock
to the places where Bleak are to be found, and it is
one of the most important foods of our best and most
valuable predatory fishes. When pursued by them,
it may often be seen leaping in companies out of the
water. In an aquarium it is a lively, playful, and
amusing pet.

In France the Bleak has been much in request
since the year 1680* * * * * 6, when a manufacturer of beads,
J ac quin by name, discovered a method of applying

the silvery pigment from its scales to practical use.
With this substance, the so-called essence d'orient , he
coloured the inner surface of hollow glass beads, which
were then filled with wax, an excellent imitation of
the genuine pearl being thus produced. Millions of
Bleak were used in this way, and great quantities of
Bleak scales imported to Paris, the chief seat of this
manufacture. It is estimated, according to Blanchard,
that about 4,000 Bleak yield half a kilogramme of
scales, and that the proportion of the colouring matter
to the total weight of the scales is as 1 to 7.

Central Europe is inhabited by three forms whose
signification was difficult to explain, until Siebold’s
suggestion that they were hybrids between the Bleak
and other species, was generally accepted. One of
them is the form described by Holandre'* in 1836
from the Moselle under the name of Leuciscus dola-
bratus, and by Gunther from the Neckar, first d under
the same name, subsequently6 under that of Alburnus
dobidoides. This variety ivas elucidated by Siebold7
as a hybrid between the Chub and the Bleak, and be-
longs to the basins of the Maas, Rhine and Danube.
It is usually as small as a Bleak and also of the same
appearance, though with less ascending mouth, less
prominent lower jaw, and shorter anal fin with straight
or rounded (convex) margin. But it sometimes attains
a length of at least 31 cm., and is then more like a
Chub, with the scales pigmented at the hind margin
with black. The pharyngeal teeth resemble those of
the Bleak.

Another similar form has been described by Jac-
kei// from Bavaria under the name of Alburnus Bosen-
haueri , and by Benecke7' from Deutsch-Eylau (Prussia)
under that of Scardiniopsis alburniformis. Both these
authors interpret it as a cross between the Rudd and
the Bleak. Its body, according to Benecke, is deeper*
than that of the Bleak, its scales are coarser and gen-
erally fewer (45 — 47 in the lateral line), the anal fin

* Nagva iakttagelser rorcmde de vertebrerade djur , som forekomma i trakten af Lomma, disp. Lund 1859, p. 27.

b See Blanchard, 1. e. In Reaumur — Hist, de l’Acad. Roy. d. Sciences, An. 1716, p. 229 — the discovery is said to have been

made in 1656.

c Fauna du Departement de la Moselle, p. 248.

d Jahresb. Ver. Vat. Naturk. Wiirtemb., Jahrg. IX (1853), p. 314.

6 „ „ „ „ „ „ XIII (1857), p. 51, taf. II.

f Siissivasserf. Mitteleur., p. 164.
g Zooi. Garten 1866, p. 20.
h Zool. Anzeiger 1884, p. 228.

1 22 % of the length of the body to the end of the lower caudal lobe corresponds to 24 % of the length to the tip of the middle
caudal rays, and is a measurement by no means uncommon, at least in gravid females of the common Bleak.

796

SCANDINAVIAN FISHES.

is shorter with fewer rays“, the caudal less deeply
forked, the silvery lustre not so bright, and the pec-
toral, ventral, and anal fins sometimes show a reddish
tinge. But the most convincing proof of its hybrid
nature lies, it is stated, in the outer row of pharyngeal
teeth, which is frequently composed of 3 teeth, the regular
number in the Rudd, and in the sharp median line of
the ventral edge between the ventral fins and the anal
aperture, which is generally covered by a row of bent
scales, but sometimes scaleless as in the Bleak. In the
last, respect this form consequently resembles the follow-

ing species ( Spirlinus bipunctatus) ; and in the common
Bleak too we sometimes find that the scales at the
sharp ventral edge behind the ventral fins bend at an
angle over the median line. In Scandinavian Bleak,
however, three teeth have never been found in the
outer pharyngeal row.

The third of these forms was seen by Siebold4
in the fishmarket at Konigsberg. On the assumption
that it is a hybrid between the White Bream and the
Bleak, he gave it the name of Bliccopsis alburniformis.

(Ekstrom, Smitt.)

Genus SPIRLINUS.

Beginning of the dorsal fin situated at a distance from the tip of the snout less than 3 times the length of the
head. Length of the base of the anal fin less than l/6 {between 16 and 18 %) of that of the body, and also
less than that of the head. Pectoral fins shorter than the longest ray in the dorsal fin. Point of the lower jaw

situated in a line with the the tip of the snout when

situated in the anter

These characters apply to only one known species,
which is also so nearly allied to the preceding genus
that it was not separated therefrom until 1882 (by Fatio).
But this resemblance is coupled with so close an ap-
proximation to forms treated of below that Gunther

the mouth is closed. Scales thin and deciduous; nucleus
ior half of the scale.

referred the species to the genus Abramis. The generic
limitation is in many cases hardly more than a matter
of taste: its object here is among the genera to mark
the different stages in the gradual transition from the
Leuciscines to the Abramidines.

a 15 branched rays. This number falls, however, according to Fatio, between the limits of variation in the Bleak (see above).
6 Susswasserf. Mitteleur., p. 168.

ABRAMIDINES.

797

THE SPERLIN-BLEAK (sw. braxenlojan).
SPIRLINUS BIPUNCTATUS.

Fig. 197.

Scales in the lateral line about 50 (44 — 52). Branched rays in the anal fin about 15 (14 — 16). Lateral line
included between two black streaks; sides also marked with scattered black spots. Pharyngeal teeth compressed ,
hooked at the tip, with smooth masticatory surface, and set in two rows; 2, 5(4) — 4(5), 2.

a

Fig. 197. Spirlinus bipunctatus, Datural size. From the Lake of Zurich, a, left inferior pharyngeal seen from above, b, same bone seen
from without, both figures twice the natural size; c , the middle tooth in the inner row, three times the natural size.

9 6

C. x + 1 + 17 + 1 + x ; L. lat ■ 44 — 5 2 ; L. tr. — 1 ; Vert. 38 — 40 c.

4

Syn. Cyprinus bipunctatus, Bl., Fisch. Deutschl., pt. I, p. 50, tab.

VIII, fig. 1 ; Agass. {Aspius), Mem. Soc. Sc. Nat. Neuch.,
tom. I, p. 38; Cuv., Val. ( Leuciscus ), Hist. Nat. Poiss.,
tom. XVII, p. 259 ( + Leuc . Balclneri, p. 262); Bonap.
( ALburnus ), Cat. Met. Peso. Ear., p. 33 ; G-thr ( Abramis ),
Jahresb. Ver. vaterl. Naturk. Wurtemb., Jahrg. IX, p. 307 ;
Hckl, Kn. ( Albumins ), Siisswasserf. Oestr. Mon., p. 135;
Dyb., Cypr. Livl., p. 161; Sieb., Siisswasserf. Mitteleur., p.
163; Blanch., Poiss. d. eaux douces Fr., p. 371; Feddersen
{Aspius), Naturh. Tidskr. Kbhvn, ser. 3, vol. XII, p. 91;
Bncke {Alburnus), Fisch., Fischer., Fischz. 0., TV. Preuss.,
p. 128; Fatjo {Spirlinus), Fne Vert. Suisse, vol. IV, p. 392;
Lillj. {Abramis), Sv., Norg. Fn., Fislc., vol. Ill, p. 322.

The Sperlin-Bleak never exceeds the average size
of the common Bleak. According to Fatio it may
attain in Switzerland a length of 15 cm.; but the
ordinary length of adult specimens, including the whole
caudal fin, lies between 10 and 13 cm.

In form this species is deeper than the preceding
one, and consequently shows greater lateral compres-
sion, the greatest depth being at least about 1/id, the
least depth about 1/11 % of the length of the body to
the end of the middle caudal rays, and the greatest
thickness (across the opercula) at most about 2/s, on
an average 36 %, of the greatest depth.

In most other respects it resembles the Bleak;
but the mouth does not ascend so sharply, and the

“ Sometimes 17, according to Fatio; sometimes 18, according to Gunther.
. 8 — 10

0 — according to Fatio.

3 — 5

c According to Gunther and Fatio.
d 24'7 % — 26‘7 % according to our measurements.
e 8’8 / — 9’4 % according to our measurements.

798

SCANDINAVIAN FISHES.

point of the lower jaw, though it fits into a shallow in-
dentation at the tip of the snout, and projects distinctly
beyond the latter when the mouth is open, is not pro-
minent when the mouth is closed. The jaws are some-
what larger, the length of the upper jaw from the tip
of the snout measuring about 31 — 35 % of that of the
head, and the length of the lower jaw about 41 — 45 %
of the same. The dorsal fin is situated as a rule some-
what farther forward, its beginning lying at a distance
from the tip of the snout of about 50 — 47 % of the
length of the body, and the distance between the tip
of the snout and the ventral fins being about 85 — 90
% of that between the same point and the dorsal fin.
This fin is also both higher — its longest ray measures
about 19 or 20 % of the length of the body — and
longer, the length of its base being about 79 (11 ‘8 —
11 '3 %) of that of the body. The anal fin is also
higher; its longest ray measures about 15 — 14 % of
the length of the body.

The most striking difference from the Bleak ap-
pears, however, in the coloration. The ground-colour

is indeed the same; but the lateral line, which during
life is itself more or less red, is included between two
black streaks running along and close to it. These
streaks are formed by two small, elongated, black spots
on each scale in the lateral line, one above and one
below the opening duct. At the base of the scales on
the sides of the body too, at least for 3 or 4 rows
above and sometimes also below the lateral line, there
appear somewhat larger, triangular spots, one on each
scale, apparently forming longitudinal streaks along
the sides.

The continental range of the Sperlin-Bleak in
Central Europe is about the same as that of the
Bleak, and the species occurs in similar localities, but
not in such numbers. The Germans call it Aland-
bleclce, Schusslaube, Schneider , etc.; in France it is most
commonly known as the Spirlin ( Eperlan de Seine).
It has only once been found within the limits of the
Scandinavian fauna: Feddersen records (1. c.) the tak-
ing of a specimen in Fake Scanderborg (Jutland) in
July, 1877.

Genus LEUCABRAMIS.

Beginning of the dorsal fin situated at a distance from the tip of the snout less than three times the length of
the head. Length of the base of the anal fin less than V5 (77V2 — 19 %) of that of the body , and also less than
that of the head. Pectoral fins shorter than the longest ray in the dorsal fin. Tip of the snout projecting in

front of the mouth. Scales firmly attached , Leuciscine in texture.

Among the Abramidine species hitherto described
from Europe there are two — probably, however, varie-
ties of the same species — Abramis vimba and Abramis
elongatus ( melanops ), which seem to be assigned by the
development of the anal fin to a place beside Alburnus.
The form of the dorsal fin — comparatively long and
low — also divides them from the true Breams; but the
most important difference consists in a carina, formed

by the bent scales of the median line, at the dorsal
edge of the tail behind the dorsal fin. One of these
two species, if it is still to be regarded as such —
Siebold suggests that it should be interpreted as a
variety due to a constant life in rivers, without mi-
gration to lakes or the sea — belongs to Russia and
Germany (S. E. and N.). The second is our well-
known

ABRA MI DINES.

799

THE ZARTHE ( sw. yimman).
LEUCABRAMIS VI MB A.

Plate XXXV, fig. 3.

Scales in the lateral line about 60 ( 58 a — 61). The dorsal fin begins at least a little in front of the middle of
the body, and the distance between it and the tip of the snout ( about 48 — nearly 50 % of the length of the body)
is less than 4/5 of that between the anal fin and the same point. Height of the dorsal fin less than ')5 of the
length of the body. Pharyngeal teeth not hooked, compressed, with smooth ( terete or one-grooved) and almost

terminal masticatory surface, and set in one row: 5 — 5.

Fig. 198.

Pharyngeal bones and pharyngeal cartilage of Leucabramis vimba, natural size; a, left pharyngeal seen from above; b, the same

seen from without (the left); c, pharyngeal cartilage.

R. br. 3; D.

3 12

A _ p _ y _ .

17— 20r’ ' 15—17’ ' 9d’

C. .r?+l + 17 + l+ jr; L. lat. 58 a — bl; L. tr. 1:

5 — 6

Vert. 46.

Syn. Capito anadromus, Gesn., Hist. Anim ., lib. IV, Paralip., p. 11.

Nasus, Schonev., Ichthyol. Slesv. Hols., p. 52. Cyprinus
Capito anadromus dictus, Art., Ichthyol., Syn. Pise., p. 8.
Cyprinus rostro nasiformi, dorso acuminato, pinna ani ossi-
culorum viginti quatuor, Art., ibid., Gen. Pise., p. 6 ; Syn.
Pise., p. 14; Descr. Spec. Pise., p. 18; Lin., Fn. Suec.,
ed. I, p. 123.

Cyprinus Vimba, Lin., Syst. Nat., ed. X, tom. I, p. 325; Bl.,
Fiscli. Deutschl., pt. I, p. 38, tab. IV; Retz., Fn. Suec.
Lin., p. 359; Pall., Zooyr. Ross. Asiat., tom. Ill, p. 322
(+ Cypr. carinatus, p. 323); Ekstr. (subg. Abramis ), Vet.-
Alrad. Handl. 1830, p. 184; Nilss., Prodr. Ichthyol. Scand.,
p. 31; Cuv., Val. (Abramis), Hist. Nat. Poiss., vol. XVII,
p. 65; Kr., Danm. Fisk., vol. Ill, p. 400; Nilss., Skand.
Fn., Fisk., p. 322; Sundev. (Cyprinus), Stockh. L. Hush.
Sallsk. Handl. 1855, p. 82; Hckl., Kn. (Abramis), Siiss-
wasserf. Ostr. Mon., p. 109; Widegr. (Cyprinus), Fiskfn.,
Fisher. Norrb. L ., Landtbr. Akad. Hand]. 1861, p. 7 (sep.);
Dyb. (Abramis), Cypr. Livl., p. 183; Sieb., Siisswasserf.
Mitteleur., p. 125; Mqrn, Finl. Fiskfn. (disp. Helsingf.
1863), p. 41; Gthr, Cat. Brit. Mus., Fish., vol. VII, p.
303; Malm, Gbgs, Boh. Fn., p. 566; Bncke, Fiscli., Fischer.,
Fiscliz. O., W. Preuss., p. 120; Mela, Vert. Fenn., p. 329,
tab. X; Mob., Hcke, Fiscli. Osts., p. 117; Reut., Sundm.,
Finl. Fisk., tab. I; Lillj., Sc., Norg. Fn., Fisk., vol. Ill,
p. 304.

In Sweden the ‘Zarthe’ attains a length of at least
37 cm., including the whole of the caudal fin. Spe-
cimens of this size have been forwarded to the Royal
Museum from Norrkoping on the Motala River.

The body is deeper than in Alburnus, but shal-
lower than in Abramis, the greatest depth, at the be-
ginning of the dorsal fin, varying between about 26
and 32 % of the length to the end of the middle cau-
dal rays, and the least depth (least depth of the tail)
between about 9 and 1 0 1/2 % of the same. It is rather
strongly compressed, with almost vertical sides; the
greatest thickness of the head is about 2/5 °f the great-
est depth of the body, which is generally not much
thicker, though in gravid females the thickness of the
abdominal region may be half as great as its depth,
or a. little more. The dorsal profile shows a more or
less distinct break at the occiput, and at this point
rises in a faint curve which soon passes into an almost
straight line, interrupted only by a slight elevation at
the beginning of the dorsal fin. The upper and lower
profiles of the head converge in front with fair regu-
larity at an angle of about 60°. The ventral profile
is more curved in front and behind than the dorsal;
but at the middle, in front of and behind the ventral

a Sometimes 54, according to MSbius and LIeincke.

6 7 — 9, according to Krgyer.
c Sometimes 21, according to Krgyer.

d Sometimes 8, according to Dybowski; sometimes 10, according to Siebold.

Scandinavian Fishes.

101

800

SCANDINAVIAN FISHES.

fins, to the beginning of the anal fin, it is straight.
The forepart of the back, as well as the top of the
head, is transversely convex; but towards the beginning
of the dorsal fin it grows more and more compressed,
while behind this fin the dorsal margin is again con-
vex or even fiat. Along the median line of the dorsal
margin there runs, however, interrupted only by the
dorsal fin, “a thin keel, with the appearance of a coarse
thread laid under the skin. This keel begins on the
head vertically above the anterior margin of the eyes,
and ends at the caudal fin” (Ekstrom). The anterior
part of the keel, almost to the dorsal fin, is naked;
but at this point one or two scales generally begin to
overlap the margin with their lateral part. Behind the
dorsal fin, on the other hand, the keel itself consists
of curved scales. The belly in front of the ventral fins
is convex or flat underneath, but between these fins
and the anal aperture sharp, with the median line scale-
less. At the sides of the straight base of the anal fin
the lowest scales project, forming a groove in which
this fin may be partially concealed.

The length of the head measures in adult speci-
mens (19 — 34 cm. long) about 2272 — 2 1 V2 % of that
of the body. Just in front of the eyes it is more or
less tumid; but the most characteristic point in its ap-
pearance is the prolongation of the snout, as in the
Haddock or the Houting. The projecting tip of the
snout is obtusely rounded, and the length of the snout
measures in young specimens about 29 % of that of
the head, in old as much as 35 % of the same, being-
less in the former than the breadth of the interorbital
space, but in the latter at least equal to or, generally,
someAvhat greater than this breadth. In the former
too the length of the snout is somewhat less, but in
the latter perceptibly greater, than that of the upper
jaw from the middle of the tip of the snout. The
mouth thus acquires a position quite ventral and al-
most horizontal, but is fairly large, so large, according
to Artedi, that the middle finger may be inserted with
ease into the mouth of large specimens, the upper jaw
being then protruded as if from a thumb-stall. The
lower jaw, which articulates below the centre of the eyes,
measures about 36 — 33 % of the length of the head.
The lips are rather thick. The eyes are of moderate
size, their longitudinal diameter varying in the above-
mentioned adult specimens between 23 V2 and 18 % of
the length of the head, or between V2 and 2/a of the
postorbital length of the same. The postorbital part

occupies between 44 and 50 % of the entire length of
the head, the eyes being thus situated sometimes en-
tirely, and always principally, in the anterior half of
the head. The nostrils, set almost on a level with the
upper orbital margin, are farther from the eyes than
in the preceding forms; but the distance between the
posterior nostril and the eye is never more than half
of that between the anterior and the tip of the snout.
The gill-openings are large, the branchiostegal mem-
branes coalescing with the isthmus close beside each
other and in about a line with the posterior margin
of the eyes. The gill-rakers in the outer row on the
first branchial arch are not very close-set (15 or 16)
and triangular in shape, the upper and lower ones
pointed, the middle ones blunter and with a tendency
to ramification at the tip. The outer anterior margin
of the pharyngeals is set with 10 — 12 small spines.
The pharyngeals are comparatively short, and their an-
terior (lower) arm is deeply grooved (bilobate) at the
inferior margin. The pharyngeal teeth, which have
been described above, are in other respects, both in
form and arrangement, not unlike those of the Roach.

The shape of the dorsal fin also reminds us most
of the Roach, but it is somewhat more pointed, as in
Abramis, and the base is comparatively shorter. Its
position is given above. The base measures about 1/10
(10 — 10'6 %), and the height about 19 — 1 6 1/2 %, of the
length of the body. The distance between the anal fin
and the tip of the snout occupies in young specimens
about 62 %, in old sometimes nearly 66 %, of the length
of the body. The base of this fin measures in all our
specimens 18 — 19 % (according to Dybowski sometimes
only 1 7 x/2 %) of the length of the body, and its great-
est height in the males about V9, in the females about
710, of the same. The caudal fin is deeply forked, its
middle rays measuring about 7V2 — Ql/2 % of the length
of the body, and about 1/s of that of the longer (the
inferior) caudal lobe.

The pectoral fins are obtusely pointed, the second
or third ray being the longest. Their greatest length
is about 15 or 16 % (in a few males nearly 17 %) of
the length of the body. The preabdominal length is
about 22 — 24 % (in a few females 24 V2 %), and the
postabdominal length about 20 — 20 V2 % (in a few fe-
males 21 %), °f the length of the body. In percentage
of the same length the distance between the ventral
fins and the tip of the snout is about 43 — 45 (in a
few females nearly 46), and the length of these fins in

ZARTI-IE.

801

the females between 14 and 15, in the males between
15 and 16". In relation to the dorsal fin they lie so
far back that the distance between their insertion and
the tip of the snout is as a rule perceptibly more than
87 % (87 1/3 — 9472 %) of that between the dorsal fin
and the same point.

The scales are middle-sized, deeply embedded in
the skin, and imbricated. They are of an irregular
quadrilateral shape and rounded, with almost central
nucleus, dense concentric striae, and numerous and Avell-
defined radiating grooves, the latter alternating, both
in front (finer) and behind (coarser and more regular),
in their greater or less extent tOAvards the nucleus.

In coloration two varieties are distinguished* * 6, Nek-
vimma (Pale Zarthe) and svartvimma (Black Zarthe),
the latter being probably no more than the spaAvning
dress. Both on the head and the back the coloration
is above blue or greenish broAvn, at the top of the sides
of the body silvery with a dash of blue, fading beloAv
into the pure silvery white of the belly. On the sides
of the head the dark blue tint extends down to the
upper part of the gill-cover, and the silvery lustre
sIioavs stronger or fainter traces of a brassy colour. In
Germany its dark snout has gained for the Zarthe the
name of Blaunase or Bussnase. All the fins are light,
gray or grayish blue, more or less tinged, especially
the caudal, Avith orange at the base. The iris is of a
pale brassy yelloAv, shading into greenish above the
pupil. Our figure, Avhich is coloured from a sketch by
W. v. Wright, represents the Zarthe in this dress of
almost pure silver. On weedy bottoms or in dark
water the colours of the fish are greener with a more
powerful brassy lustre. Siebold has described the dark
spaAvning dress, in Avhicli the sides of the body, even
beloAv the lateral line and close to the ventral and anal
fins, acquire a singular silky lustre, this being caused
by a deep black pigment from Avhich the ventral side
itself, from the lips to a narrow strip at the margin
behind the anal fin, stands off sharply Avith its deep
orange hue. The latter colour also extends to the
paired fins and the base of the anal fin; the dorsal and
caudal fins, the upper margin of the pectorals, and the
loAver margin of the anal fin being, on the other hand,
blackish. In addition to this spaAvning dress, Avhich is

common to both sexes, the males are distinguished at this
season by the usual dermal eruption, consisting of small
round tubercles on the occiput, the opercula, the margins
of the scales, and the inner surface of the paired fins.

The Zarthe is properly a marine fish, but also lives
in the great lakes, and ascends rivers in order to spawn,
being thus a so-called anadromous species. Its geo-
graphical range comprises Eastern Europe from Finland,
SAveden, North-Avestern Germany, and the Avatershed of
the Danube to the Black Sea Avith its Russian river
systems and the Sea of Azov. It makes its Avay up
the Elbe and Weser from the North Sea, but is Avant-
ing in the Rhine. In Finland, Avhere it is common
among the southern islands and, to the east, in Lake
Ladoga, it does not go farther north, according to Mela,
than the sixty-third degree, being very rare even in
that latitude. From the SAvedish coast of the Gulf of
Bothnia Ave are told by Widegren (1. c.), though his
statement is perhaps questionable, that the Zarthe oc-
curs in the loAver course of the Lulea Elf (nearly 66°
N. lat.). The northernmost locality in Sweden to which
Ave can Avith certainty assign this species, is the neigh-
bourhood of Hudiksvall (about 62° N.), from Avhich
region Mr. Wistrom has sent to the Royal Museum a
specimen 24 cm. long, taken in the fjord of Lingaro
on one of the last days in September, 1882. In the
same neighbourhood, Ave are told, it sometimes, though
seldom, ascends the River Dekanger in spring. Ever
since Nilsson’s time the Zarthe has been knoAvn in
Blekinge; but in Scania it has never been found. It is
commonest in the Malar Valley and the Baltic island-
belt Avith the streams that fall into this part of the
Baltic. In Lake Wener it is also found, and is some-
times caught, “especially in the months of May and
June” (Malm) in the River Gotha at Gothenburg. In
Nonvay, Denmark, and the Avest of the Baltic the spe-
cies is unknoAvn; but from the Elbe it Avas described
even by Gesner and Schonevelde. From the Baltic
it enters the German Haffs to spaAvn. According to
Dybowski (1. c.) and Seidlitzc it occurs in all the riv-
ers of the Baltic Provinces of Russia. Its Baltic range
being of such extent, Ave might Avell expect to find it
both off Gothland and Bornholm; but is has not yet
been observed in either of these localities.

a According to Dybowski the same sexual distinction was present in his specimens; but the percentages are somewhat lower, this

being probably due to a different manner of measurement.

6 According to Mr. Arosenius, who forwarded to the Royal Museum specimens of these varieties from the Motala River (Norrkoping) in 1834.
c Fauna baltica , p. 101.

802

SCANDINAVIAN FISHES.

Ekstrom has described the habits of the Zarthe in
the neighbourhood of Morko. At the approach of spring
it ascends the rivers, where it remains during the sum-
mer. Towards autumn it returns to the sea, where it
passes the winter, in so deep water that it has never
been caught at this time of year. It probably chooses
its winter-quarters outside the island-belt. In the lakes
too it retires to their deepest parts0.

Cunning and shy, like most of the Cyprinoids, says
Ekstrom, it is difficult to catch, except during the
spawning-season, and dies soon after it has been taken
out of the water. It seldom attains any considerable
size. The flesh is white, but flabby and of poor flavour.
In a female 31 cm. long and 7/10 kilo, in weight Bloch
estimated the number of the ova at 28,800, whence it
appears that the Zarthe multiplies quickly enough; but
according to Ekstrom its growth is slow, and it thrives
only in clear or running water with a stony or sandy
bottom.

In the stomach of the Zarthe Ekstrom seldom found
traces of other food than crustaceans, insects, worms,

and, most often, the crushed shells of mollusks ( Neri -
tin a) ; hardly ever of vegetable substances.

At the end of May — still according to Ekstrom
— the spawning-season begins, and the Zarthe ascends
such streams and rivers with stony bottom as fall into
the sea or the great lake where it has passed the winter.
The roe is deposited on and adheres to the stones,
against which the spawning fish eagerly rubs itself in
order to get rid of its burden.

In spring and autumn, as it roves to and from
the spawning-place, small numbers of Zarthe are taken
among other fishes with net and seine in Sodermanland.
During the spawning-season it is caught in a special
kind of large hand-net.

In the localities affected by the Zarthe for the pur-
pose of spawning, continues Ekstrom, where it may
consequently be taken in numbers, it always appears
on the labourer’s humble board. It is generally but
little esteemed, and indeed requires skilful preparation
to suit a more delicate palate. Ekstrom found it best
when fried.

Genus ABRAMIS.

Beginning of the dorsal fin situated at a distance from the tip of the snout less than three times the length of
the head. Length of the base of the anal fin more than 1/5 {21 — 35 %) of that of the body , and greater as a
rule than that of the head. Pectoral fins shorter than the longest ray of the dorsal fin. Tip of the snout pro-
jecting only slightly , if at all, in front of the mouth. Scales firmly attached, Leuciscine in structure.

This genus too is mainly European, and contains
only a few species (4 determined and recognised with
certainty, excluding hybrids). Still Heckel 6 divided
it into 3 genera: Abramis, Blicca, and Bailer us, the
first with pharyngeal teeth set in a single row and the
base of the anal fin comparatively little elongated, the
second with two rows of pharyngeal teeth, the third
with pharyngeal teeth set in one row and the base of
the anal fin greatly elongated. The first two genera,
however, so closely resemble each other in all other
respects that their division even into separate sub-
genera must appear unnatural. The character drawn
from the arrangement of the pharyngeal teeth in one
or two rows has also been regarded by more recent
authors as insufficient for the establishment of distinct

genera. The great elongation of the anal fin should
rather be employed for this purpose; but the species
are so few that the division is destitute of practical
importance.

The three Scandinavian species may be distin-
guished as follows:

A: Length of the base of the anal fin less
than 3 10 of that of the body.
a: Length of the lower jaw less than 3/4

of the least depth of the tail Abramis blicca.

b: Length of the lower jaw more than

3/4 of the least depth of the tail Abramis brama.

B: Length of the base of the anal fin more

than 3 10 of that of the body Abramis ballerus.

Besides these species, however, we must not forget
three hybrids, which occur in Scandinavia as well as

a In Atter See (Austria) according to Heckel and Kner, it keeps at a depth of 10 fthms., in winter even 20; and the shoals in
which it is collected at this season betray their presence by rooting up the muddy bottom and discolouring the water.
b Russeggers Reise , p. 1032.

WHITE BREAM.

803

elsewhere, between the Breams and the Lenciscines.
Instances of a Roach or a Rudd with more than 14
branched rays in the anal tin, of a Roach with pha-
ryngeal teeth set in two rows, or of a White Bream
with less than 18 branched rays in the anal tin, have
been explained on the assumption that these forms are
hybrids between the White Bream and the Roach or

the Rudd. This explanation is indeed correct to all
appearance; but its demonstration, which can lie at-
tained only by experimental methods, has still to be
made by pisciculturists. These remarks also apply to
the assumed hybrid between the Bream and the Roach.
The last form is usually most like a Bream, but has
only 15 — 18 branched rays in the anal fin.

THE WHITE BREAM OR BREAMFLAT (sw. bjorknan).

ABRAMIS BLICCA.

Plate XXXV, fig. 2.

Scales in the lateral line about 46 (45 — 50). Anal fin with 21 — 23a branched rays. Between the lateral line
and the dorsal fin 9 or 10, in exceptional cases 11, rows of scales. Pectoral and ventral fins red or yellow at

the base. Pharyngeal teeth set in two rows: (1)2(3), 5(6) — .5, (1)2(3).

a

Fig. 199. Pharyngeal bones and pharyngeal cartilage of Abramis blicca, natural size; a, b , and c as in the preceding figure.

R. hr. 3; D.—.\ A. 3 ;

’ 8 6 2 1 c — 23(24)

P.

14—16(17)’

2

v. n

C. x + 1 4- 17 + 1 + x; L. lat. (43)45 — 50; L. tr. ^ 1;

5—6(7)

Vert. 38—40.

Syn. Ballerus, Rondel., De pise, lacustr., p. 155. Blicca , Gesn.,
Hist. Anim., lib. IV (Francof. 1620), p. 24 ( + (?) Plestya,
p. 25, ex Belon.). Gyprinus quincuncialis, pinna ani ossi-
culorum viginti quinque, Art., Gen., p. 3; Syn., p. 13;
Spec., p. 20. Cyprinus pinnae ani - radiis viginti quinque,
Lin., Fn. Suec., ed. I, p. 124 (syn. nec descr.) = Cyprinus
Bjoerkna, Syst. Nat., ed. X, tom. I, p. 326 (syn. nec. descr.).
Cyprinus Plestya , Leske, Ichtliyol. Lips. Spec., p. 69.
Cyprinus Blicca, Bl., Fische Deutschl., part. I, p. 65, tab. X;
Ekstk. (subg. Abramis ), Vet.-Akad. Handl. 1830, p. 179;
Nilss., Prodr. Ichtliyol. Scand., p. 31; Fr., Ekstr., v. Wr.,
Skand. Fisk., ed. 1, p. 64, tab. 12; Kr. {Abramis), Damn.
Fisk., vol. Ill, p. 389; Sundev. {Cyprinus), Stockh. L. Hush.
Sallsk. Handl. 1855, p. 82; Gthr. {Abramis subg. Blicca),
Cat. Brit. Mus., Fish., vol. VII, p. 306; Feddersen, Naturh.
Tidskr. Kbhvn, ser. 3, vol. XII, p. 86; Day, Fish. Gt. Brit.,
Irel., vol. II, p. 196, tab. CXXXVI; Mob., IIcke, Fisch.
Osts ., p. 118; Lillj., Sv., Norg. Fn., Fisk., vol. Ill, p. 313.
Cyprinus Laskyr, Guldenst. apud Pall., Zoogr. Boss. Asiat.,
tom. Ill, p. 326; — vide Nordm., Voy. Russ. Mer. (Demidoff),

° Sometimes 24.
b 7 — 9, according to Kroyer.
c Sometimes 19, according to Fatio.

tom. Ill, p. 504, tab. 22, fig. 1 ; — Hckl., Kn. {Blicca),
Susswasserf. Ostr. Mon., p. 123.

Abramis micropteryx et Abr. erythropterus, Agass., Mem. Soc.
Sc. nat. Neuch., vol. I, p. 39 (cf. Cuv., Val., Hist. Nat.
Poiss., vol. XVII, pp. 44 et 58).

Abramis Bjoerkna , Nilss., Skancl. Fn., Fisk., p. 328; Sieb.
{Blicca), Susswasserf. Mitteleur., p. 138; Mgrn, Finl. FisJcfn.,
disp. Helsingf. 1863, p. 42; Blanch. {Abramis, subg. Blicca),
Poiss. eaux douces Fr., p. 359; Malm {Abramis), Gbgs, Boh.
Fn., p. 565; Mor., Hist. Nat. Poiss. Fr., tom. Ill, p. 398;
Bncke {Blicca), Fisch., Fischer., Fischz. O., W. Preuss.,
p. 123; Fat., Fn. Vert. Suisse, vol. IV, p. 358; Mela
{Abramis), Vert. Fenn ., p. 333, tab. X; Reut., Sundm.,
Finl. Fisk., tab. IV.

Blicca argyroleuca , Hckl. Kn., 1. c., p. 120; Dybowski, Cypr.
Livl., p. 202.

Obs. Artedi described this species with fair accuracy in De-
script. Spec. Pise. (p. 20, No. 9), under its Upland name of Bjorkna,
and also included it in Synonymia Piscium (p. 13, No. 27) and Gen.
Pise. (p. 3, No. 3). But he failed to perceive that it was the spe-
cies called Ballerus and Blicca by former ichthyologists, referring
these names instead to his own Blicca {Descr. Pise., No. 11), to
which Linnaeus on his authority applied the specific name of Ballerus,
which it has since retained. Artedi’s Bjorkna remained unknown to
Linnaeus, though the name is given both in the Fauna Suecica. and

804

SCANDINAVIAN FISHES.

the Systema Naturce. This is evident from the appended note in the
Fauna, where Linnaeus expressly states that he had counted 35 rays
in the anal fin of the Bjorkna ■ — a number which, among the species
indigenous to Sweden, is to be found in Abramis ballerus alone. In
Retzids’s edition of the Fauna Suecica the error was not corrected,
although Bloch had already given a lucid description and a good
figure, considering the date at which it was executed, of Artedi’s
species, under the name of Cyprinus Blicca . which it still retains.
Nilsson was the first Scandinavian writer to restore the name of
Bjorkna to its original signification, and to recognise Bloch’s nomen-
clature (see Prodr. Ichth. Scand.); but Ekstrom had previously (Vet. -
Akad. Handl. 1830) expressed the same opinion on the strength of
information privately supplied to him by Nilsson.

The White Bream is the smallest Scandinavian
member of the genus Abramis. It never attains any
considerable size. Its ordinary length is between 15
and 23 cm., rising, however, sometimes to 30 or, in
exceptional cases, 35 cm., including the whole of the
caudal fin.

In most respects the White Bream comes nearest
our common Bream, and in appearance so closely re-
sembles young Bream that it is frequently confounded
with them, under the common name of Braxenpanka.
In our description we shall therefore lay special stress
on the differences brought to light by a comparison
between them.

In the form of the body it is hardly possible to
detect any constant difference between the White Bream
and the Bream. The body of both is compressed and
deep, though generally a little less so in the former.
In White Bream preserved in the Royal Museum, and
measuring between 106 and 239 mm. in length to
the end of the middle rays of the caudal fin, the great-
est depth of the body is between 31 % (in young spe-
cimens) and nearly 40 % (in old) of its length, and the
greatest thickness (with more individual variations) be-
tween V3 and V4 of' the greatest depth. The least depth
of the tail increases during these changes of growth
from 9‘4 to 11*3 % of the length of the body. The
back, which rises somewhat abruptly from the occiput,
then ascends in a regular curve to the beginning of
the dorsal fin, where it forms an obtuse angle, and
afterwards slopes almost in a straight line to the caudal
fin. Though carinated from the occiput to the dorsal
fin, it is thicker and more convex than in the Bream.
The belly is flat from the pectoral region to the vent-
ral fins. From this point to the vent it is sharply
carinated. The ventral profile is almost straight from
the pectoral region to the anal aperture, where it- forms
an obtuse angle, more acute, however, than the dorsal

angle, and advances in a somewhat incurved line to
the caudal fin.

The head, the length of which is about 1/5 (21 x/2 —
20 %) of that of the body, tapers uniformly forwards
from above and below, but the snout is fairly thick and
obtuse, projecting a little beyond the mouth. The fore-
head is broad and convex, the breadth of the inter-
orbital space being about 772 — 8 % of the length of
the body, or 3572 — 3872 % of the length of the head.
The frontal profile is straight from the occiput to the
nostrils, where it descends abruptly, and slopes towards
the snout. The sides of the head are moderately com-
pressed, the thickness being equal to the depth, mea-
sured at the anterior orbital margin. The mouth is
small and ascends only slightly, but may be projected
to some distance in a tubular form. The corner of the
mouth falls short of the perpendicular from the anterior
orbital margin. The length of the snout, which mea-
sures about 5 — 6 % of that of the body or about 23 —
31 % of that of the head, is as a rule equal in young
specimens to the length of the upper jaw (from the tip
of the snout), in old a little greater. The length of
the lower jaw measures about 6x/2 — 7 % of that of the
body, or about 32 — 34 % of that of the head, is less
than that of the suture between the suboperculum and
operculum, and also than 71 % (in the said specimens
70 — 57 %) of the least depth of the tail. The eyes,
on the other hand, are comparatively larger than in
the Bream, their longitudinal diameter varying in our
specimens between about 34 and 27 % of the length of
the head, while in young specimens it is perceptibly
greater, in old slightly less, than the length of the snout.
Their size is often so considerable that the White Bream
may be at once distinguished thereby from the young
Bream in its company. The position of the eye is such
that the postorbital length of the head in young spe-
cimens is less, in old somewhat more, than half its
entire length. The nostrils are set much as in the
Zarthe, the distance between the posterior nostril and
the upper anterior part of the orbital margin being
about 7s °f that between the anterior nostril and the
tip of the snout. The anterior nostril is smaller than
the posterior, but circular; while the latter is obliquely
set and elliptical or crescent-shaped. The gill-openings
are smaller than in the Zarthe, the branchiostegal mem-
branes coalescing below with the isthmus at a greater
distance from, each other and farther back, in a line
with the hind (vertical) margin of the preoperculum.

WHITE BREAM.

805

The gill-rakers are short and scattered, the outer row
on the front of the first branchial arch containing 14
or 15, and the outer anterior margin of the pharyngeals
being furnished with almost as many (13). The pha-
ryngeals are armed as a rule with two rows of teeth
(fig. 199), the inner row being always composed of 5
compressed and nearly straight teeth, with distinctly
hooked tips in most cases only in the hindmost (upper-
most) tooth and the middle one, the outer generally of
two (in exceptional cases one or three) smaller, more
cylindrical teeth. According to Ekstrom the outer row
is often wanting, which probably depends on the shed-
ding of the teeth, when the pharyngeal teeth readily
drop out, or lie loose in the gums".

The dorsal fin lies on the hind arm of the angle
formed by the dorsal profile, so high that the first rays
occupies the apex of the back. The distance between
it and the tip of the snout measures more than half
(about 51 — 54 %) of the length of the body, but less
than 88 % (81 — 87 %) of the distance between the tip
of the snout and the anal fin. Its height is about twice
its length, the longest ray measuring about 22 — 26 %
of the length of the body, and the base about 11 — 12
% of the same. The fin is pointed in front, with the
upper posterior margin obliquely truncate, the height
behind being only l/3 of that in front. The anal fin, in
consequence of its length, is not so obliquely truncate,
but more deeply concave at the margin. The distance
between it and the tip of the snout is about 58 — 64 %,
the length of the longest ray about 15 (exceptionally
14) — 18 %, and its base about 23 (exceptionally 21) —
2672 the length of the body. The caudal fin is

deeply forked, with the lower lobe somewhat longer
than the upper. Its middle rays measure about 9 or
10 % (exceptionally 8 72 or H °A of the length of the
body, or rather less than 2/5 of that of the longest ray
in the lower lobe.

The pectoral fins are normal in form, but com-
paratively short, always shorter than the preabdominal
length. Their length is about 17 (16 x/2) — 18 (1877 0/0
of that of the body. The ventral fins are inserted at
a distance from the tip of the snout measuring about
40 — 45 % of the length of the body, and less than 87 %
(77 7g- — -86 7g 0/,°) °f the distance between the dorsal fin
and the same point. Their length is about 15 (14 1/2)

— 18 % of that of the body. Both the preabdominal
length and the postabdominal, the latter as a rule rather
less than the former, measure about 1/5 (20 — 23 or
24 %) of the length of the body.

The body is covered with large, striated, and im-
bricated scales, very like those of the Roach, and with
more distinct striae than those of the Bream. This
peculiarity combined with their relative size, is especi-
ally characteristic of the White Bream, and renders it
easily recognisable, if only attention be paid hereto.
The most trustworthy standard of the size of the scales,
however, is their number. In the lateral line, which
lies rather low, nearer to the belly than to the back,
we generally find only 46 scales — the ordinary varia-
tions lie between 45 and 48 — and above the lateral
line only 8 or 9 large rows and one smaller row, the
latter at the very base of the dorsal fin (the Bream
has at least 12 such rows). Between the lateral line
and the insertion of the ventral fin we usually find 6
rows of scales. The distribution of the scales on the
anterior part of the dorsal margin and on the ventral
margin behind the ventral fins has already been described.

The coloration of the White Bream changes some-
what with age and according to the season of year.
The young are of a lighter colour and have paler fins.
In spring during the spawning-season, or as shown in
our figure, the ground colour of old specimens is sil-
very white with a strong dash of yellow, darker above
and gradually passing into the olive gray of the back.
The sides of the head are bright with a play of many
handsome colours. The iris is of a faint golden yellow,
densely punctated with fine, dark green dots, collected,
especially above, into a broad, dark band. The dorsal
and caudal fins are of a plain olive gray. The pec-
toral, ventral, and anal fins are pale, with more or less
ruddy rays and a reddish tinge at the base. By this
red or orange hue of the inferior fins it is always easy
to distinguish the White Bream from young Bream, in
which these fins are pale and colourless. On the con-
clusion of the spawning the sides of the body resume
their dress of silvery white.

In the White Bream, says Fatio, the back and the
sides of the body are often strewn with some roundish,
black spots. In normal Scandinavian specimens we have
never found such spots; but in a variety (hybrid) which

a Unless it be the case that EksteSm met with hybrids between the White Bream and the Bream, with the external characters of
the former and the dentition of the latter.

806

SCANDINAVIAN FISHES.

occurs in the River Helge, and of which we shall have
more to say below, these spots, according Mr. Svens-
son, Preparator at the Royal Museum, are very com-
mon. To judge by their appearance in one stuffed
specimen, however, they seem to be morbid symptoms,
concretions of pigment on abraded scales or more or
less injured fin-rays.

The White Bream — the German Blicke or Glister,
the Danish Flire, BJege , or BJaafinne, and the French
Bordeliere a — has an extensive range in fresh water
north of the Alps, from Ireland eastwards throughout
Russia-in-Europe6. In Finland, according to Mela, it
goes north to lat. 63° 40'. In Norway it has not been
found. It is common in most of the lakes of South-
ern Sweden and in the inner part of the Baltic island-
belt. As it is a species generally despised, and while
confounded by most fishermen with young Bream, goes
by different names in most of the localities where it
occurs and is known to be a distinct form, we are as
yet unable to fix the limits of its range in Sweden,
especially to the north. We know that it is common
in the Malar Valley, Lake Wener, and the southern
provinces. Nilsson found it in the Dal Elf at Soder-
fors in 1829. Wistrom supplied Lilljeborg with in-
formation of its occurrence in the fjords near Hudiks-
vall (about 62° N. lat.), and Steffenburg of its pre-
sence in the basin of the Dal Elf up to Lake Oje in
Dalecarlia (61° N. lat.). According to Malm it is pretty
common in Bohuslan, according to Nilsson common in
Scania. It is known by many names, e. g. Bjdlke,
Blecka, Bjorkfisk , Bjorkare , Kjcirta , Panka, Bldpanka,
with several different dialectic pronunciations of these
words.

The White Bream thrives best in lakes and rivers
with a clayey and sandy bottom overgrown with weeds.
Early in spring it repairs to shallow and weedy shores,
where it spawns periodically in June or even at the

middle of May. In favourable weather each spawning
lasts about three days, at longer or shorter intervals.
The old fish spawn first, the young some time later.
The roe is deposited on the weeds, to which it adheres,
During the operation of spawning the fish plunge and
splash about at the surface, where they are seldom seen
on other occasions. The White Bream usually keeps
to the bottom, sometimes swimming in mid water.
Though shy and greatly afraid of noise at other times,
it is tame and fearless during the spawning, when it
may be caught with ease.

It passes the whole summer in shallow water, and
does not retire to the depths until autumn, when it de-
scends to its winter-quarters. It is one of the most
voracious Cyprinoids, feeding on weeds, insects, and
worms, and biting so freely that it is a nuisance to the
fisherman, for it often secures the bait without being
hooked. It has therefore received the nickname of dtare
(glutton).

Though it thieves so greedily, and is always on
the alert where there is anything to be got, it is al-
ways lean. As it is also of small size, and the flesh
very bony, it is never in request, being eaten only for
want of better fish or by the poor. Like most of the
Cyprinoids, it is very prolific — Bloch estimated the
number of the eggs in a female weighing 117 grm. at
about 108,000 — and thus yields quite a considerable
supply of food to the larger and better flavoured fishes-
of-prey.

No special fishery is carried on for the White Bream
to the best of our knowledge; but as it is, so to speak,
omnipresent, it is taken with most of the tackle em-
ployed for other fishes. The greatest quantity is caught
in spring, at the approach of the spawning-season, in
the baskets (Sw. lanor ) set for Eel in rivers and large
streams.

(Ekstrom, Smitt.)

As we have mentioned above, the difficulties of the
systematist are increased by the occurrence in this fa-
mily of forms whose characters incapacitate them for a
place either within the Leueiscine or the Abramidine
subfamily, though their whole nature distinctly indicates

“ “Because it always keeps to the shore” (Rondelet).
b Gkimm, Fishing and Hunting in Russian Waters, p. 15.

their near relationship to one or other of the species
already described. These forms have been explained as
hybrids; and in Southern Sweden (Scania and Blekinge)
we find two of them, descended probably on one side
from the White Bream and on the other from the Rudd

ABRAMO-LEUCISCINE HYBRIDS.

807

or the Roach. These forms were first discovered in
Germany, and Heckel, who did not regard them as
hybrids, bestowed upon them the special generic name
of Bliccopsis ". Much may be urged against this name
as co-ordinating hybrids with true species and genera in

the systematic nomenclature; and Fatio6 therefore coined
the compound names Scardo-blicca Erythro-Bjorkna and
Leucisco-Blicca Butilo-Bjorkna. But the forms have
been most accurately defined in Bavaria by Siebold and
Jacket, who have retained the Heckelian generic name.

BLICCOPSIS ERYTHROPHTHALMOIDES.

(hybrid between the white bream and THE RUDD.)

Fig. 200.

Body deep as in the deepest Rudd, the greatest depth being 34 — 39 % of the length, with the forepart of the
back convex and covered with scales in the median line. Dorsal fin with 8C or 9 branched rays , anal with 15d
or 16. Distance from the tip of the snout to the dorsal fin more than half the length of the body, and per-
ceptibly greater than that to the foremost point in the insertion of the ventral fins. At most 10 scales in a
transverse row above the lateral line. Pharyngeal teeth set in two rows and more or less distinctly pectinated.

Fig. 200. Bliccopsis erigthrophthalmoides, taken in a Herring-seine off Karlskrona, in May, 1876, by Mr. Lundberg, Inspector of Fisheries.
Natural size, a, left pharyngeal, seen from above, natural size; b, the same, seen from without, natural size; c, the last tooth

in the inner row, twice the natural size.

a Russeggers Reisen, Bd. 2, p. 1032.

b Faune des Verte'bre's de la Suisse, vol. IV, pp. 376, 387
c Sometimes 7, according to Fatio.
d Sometimes 12 — 15, according to Fatio.

Scandinavian Fishes.

102

808

R. br. 3; I).

8° — 9

A. 3 : P. 1

14 — 1 6 ' 16—17

SCANDINAVIAN FISHES.

v. 2

8—9’

9 io

C . #+l + li + 14- sc L. lat. 42 — 4b ; B. tv. 1.

4 — 5

Syn. Bliccopsis Buggenliagii, p. p. Heckel, 1. c. (uee Bloch);

Qvennerstedt ( Cyprinus ), Of vers. Vet.-Akad. Forh. 1877,
No. 7, p. 13, tab. VII et VIII; Malm ( Abramidopsis ), Gbgs
Naturh. Mus. Arsskrift, III (1881), p. 25.

Abramis abramo-rutilus, p. p., Holandre, Fne Mos., vide Sieb.
( Bliccopsis ), Susswasserf. Mitteleur., p. 142.

Bliccopsis evythrophthalmoid.es, Jackel, Fiscli. Bay., Abb. Zool.
Miner. Ver. Regensburg, H. 9 (1864), II, p. 49. Hybrid
between Leuciscus erythrophthalmus and Abramis blicca,
Gthr, Cat. Brit. Alas., Fislt., vol. VII, p. 233.

The hybrid assumed on the above grounds to exist
between the White Bream and the Rudd attains in
Sweden a length of at least 27 cm., including the whole
caudal tin. It has the convex dorsal margin of the
Rudd — though the dorsal profile, as in the White
Bream, forms a sharper break at the occiput — the
broad forehead of the Rudd — the breadth of the inter-
orbital space sometimes about 45 % of the length of
the head — and the comparatively low dorsal fin of the
same species — the length of the longest ray less than
V5 of that of the body. But the length of the anal
fin, which at least exceeds 16 % of that of the body,
at once shows, apart from the number of the rays, that
the fish cannot be referred to this species. The pha-
ryngeal bones and teeth are also most like those of the

Rudd; but the only essential difference in this respect
between the Rudd and the White Bream lies in the
more elongated form of the teeth, with longer and more
lateral masticatory surface, and their more distinct
pectination in the former. In the only specimen of
this hybrid whose pharyngeal apparatus we have been
able to examine, the teeth numbered 3, 5 — 5, 3.

In Sweden, as far as we know, this hybrid has
been found only at two localities in Scania and one in
Blekinge. It was first taken in 1864 by Prof. Berg-
gren in Lake Ring; but Prof. Qvennerstedt was the
first (1876) to notice and determine this find, describing
at the same time two other specimens from the same
locality, one procured through Berggren, the other
taken by Prof. Naumann, who has also presented a
specimen to the Museum of Gothenburg. In the mean-
time (1869) a specimen denoted by Qvennerstedt (1. c.,
p. 21) as ‘Ex. 1’, had been caught in the River Helge,
and stuffed by Mr. Svensson for the Museum of Kristian-
stad. The original of our figure was taken at the be-
ginning of May, 1876, by Mr. Lundberg, Inspector of
Fisheries, among a haul of Herrings caught in a seine
off Lagervik at Karlskrona. According to Qvennerstedt
two more specimens have been found in Lake Ring, but
not preserved, so that in all 8 examples have hitherto
been met with in Sweden, most of them in fresh water,
but one in the brackish water of the Baltic island-belt.

a Sometimes 7, according to Fatio.

6 Sometimes 12 — 15, according to Fatio.

ABRAMO-LEUCISCINE HYBRIDS.

809

BLICCOPSIS ABRAMO-RUTILUS.

(hybrid between the white bream and the roach).

Fig. 201.

Body more elongated , deep as in the Rudd or Roach — greatest depth about 31 % of the length. Dorsal margin

more compressed , but covered with scales , though not regularly arranged. , in the median line. Dorsal fin with 9a

branched rays, anal with 16b. Distance from the tip of the snout to the dorsal fin about half the length of the
body, and only slightly greater than that to the foremost point in the insertion of the ventral fins. At most 10

scales in a transverse row above the lateral line. Pharyngeal teeth set in one or two rows and with smooth or

only faintly pectinated masticatory surface.

201. Bliccopsis abramo-rutilus, cf\ taken at Ivristianstad, 4th May 1872

of the natural size. From a stuffed specimen

3 3 12

B. br. 3 ; D. — j d. — — , ; P. — - — — — j V. — ; C . .c+l-f-17 + l+ .zu

’ 9° 1G6 16(15) 8

10rf

L. lat. 4 7 c ; L. tr. 1.

5 — 6

Sgn. Abramis ( Bliccopisis ) abramo-rutilus, p. p., Holandre, Siebold —
vide supra.

Abramis Baggenhagii, Sel.-Longcu., Fne Beige , p. 216.
Bliccopsis abramo-rutilus, Jackel, 1. c., p. 53.

Hybrid between Abramis blicca and Leuciscus rutilus, Gthr,
Cat. Brit. Mus., Fish., vol. VII, p. 215.

In its most pronounced form, the form with the
influence of the Leuciscine type most distinct, this
hybrid most resembles a Roach with the anal fin con-
siderably elongated and the forepart of the back extra-
ordinarily compressed (carinated). It thus differs at
the first glance from the preceding hybrid partly in
the more elongated body, partly in the apparently
greater size of the scales. These are not so densely

a According to Fatio 8, or more seldom 7.
b According to Fatio 11 — 15, more seldom 16.
c 41 — 46, according to Fatio.

, according to Fatio.

4—5

810

SCANDINAVIAN FISHES.

imbricated, and a. greater part of their surface is thus
left bare, the rule being that in this hybrid most of
the scales, at least those on the middle of the sides of
the body, have the free (hind), externally visible por-
tion half (or more) as long as deep, while in the pre-
ceding hybrid, as in the Rudd and the White Bream,
the length of the said part of each scale is less than
half its depth. The same difference in the position
of the dorsal tin as appears between the Roach on one
hand, the Rudd and White Bream on the other, is
also maintained here, the distance between the tip of
the snout and the beginning of the said tin being less
in this hybrid than half the length of the body. In
the specimen examined, it is true, the dorsal tin con-
tains 9 branched rays; but the last of them is slender
and branched only at the tip, not, as is generally the
case in the Cyprinoids, divided almost to the base.
The pharyngeal teeth we have not been able to exa-
mine; the above remark on this head is taken from
Jackel, who found them to number: 0(1 or 2), 6(5)
— 5(6), 0(1 or 2).

This form, according to Mr. Svensson, is the most
common hybrid in the River Helge and the lakes
through which this river flows as it approaches the
sea. Kristianstad Museum possesses three stuffed spe-
cimens, one of which ive have been enabled to examine
by the kindness of Lecturer Wahlstedt. It is a male,
caught on the 4th of May, 1872, with the small, ver-
rucose tubercles characteristic of the spawning season,
on the sides of the head (thickest on the gill-covers),
the lateral scales on the forepart of the body, and the
inner surface of the pectoral tins. The coloration seems
to have been that typical of the Roach, the front part
of the anal fin, however, being of the black colour
frequently present in the White Bream. The entire
length to the tip of the lower caudal lobe is 29 cm.,
but to the end of the middle caudal rays — the measure-
ment constantly termed in this work, unless other-
wise specified, the length of the body — only 256 mm.
The greatest depth is 30'7 %, and the least depth 10 %,
of the length of the body. The greatest thickness —
in the stuffed specimen just in front of the dorsal tin

— measures 37*6 % of the greatest depth. The length
of the head is 20'7 % of that of the body, and the
postorbital length of the same part half the entire
length of the same. The distance from the tip of the
snout to the beginning of the dorsal tin is slightly less
than half the length of the body. The base of the
dorsal tin measures 12*5 %, its longest ray 20'3 %>, of
the length of the body, ahd its last ray rather more
than 1/3 (3472 %) of the length of the longest ray.
The beginning of the anal tin lies at a distance from
the tip of the snout equal to 65*6 % of the length of
the body; its base measures 16*8 %, its longest ray
15 %, of the same length. The length of the pectoral
fins is 16 % of that of the body. The ventral fins
are only slightly shorter than the pectoral, and the
distance between them and the tip of the snout is 93
% of that between the dorsal tin and the same point.
The length of the middle caudal rays occupies 9 % of
that of the body. Apart from all the other resem-
blances to the Roach, the position of the ventral tins
in relation to the beginning of the dorsal tin shows
that one of the parent species of this hybrid must have
been the Roach; while the length of the pectoral tins
and the number of the scales admit of the assumption
that the White Bream was the other parent species,
but exclude the Bream from the possible progenitors
of this hybrid.

From the same localitv the Museum of Kristian-
stad has received two other hybrids, similar to each
other, one of which is mentioned by Qvennerstedt
(1. c.) under the head of “Ex. 2”. The other (tig. 202)
was caught on the 17th of March, 1869, and has been
lent to me for the purpose of examination by Mr. Wahl-
stedt. It has the coloration of the White Bream, in
particular the above-mentioned black spots'1 on the
sides of the body, the dorsal, anal, and caudal fins, and
the right ventral tin. Two similar spots of somewhat
larger size, with the same symptoms of skin-disease,
are situated on the right side of the head, the lower
on the hind margin of the preoperculum, the upper
on the front margin of the operculum. The specimen
evidently has more White Bream blood in it, a cir-

a These spots, which are reproduced in our figure, cannot be minutely examined in a dried specimen, but are probably caused by
Holostomum cuticola (Nordmann, Mikrographische Beitrcige zur Naturgeschichte der icirbellosen Thieve, Heft. I, p. 49, taf. IV, fig. 1 — 4), a
flatworm occurring in the skin and eyes of many fishes, and belonging to the Trematod order — cf. for example fig. 211, p. 209 in Max
von dem Borne, Handbuch der Fisrhzucht und Fisclierei. The spots themselves are pure black; and the scales over which they extend are
partially abraded. At the centre of some spots is a blister, concealing a cavity which was probably inhabited by the parasitic worm.

ABRAMO-LEUCISCINE HYBRIDS.

811

cumstance especially distinct in the form of the body
and the situation of the dorsal tin. The fins are some-
what damaged; but the length to the tip of the lower
caudal lobe was probably almost 23 cm. The length
of the body (to the end of the middle caudal rays is
199 mm. Greatest depth 32*2 % of the length of the
body, least depth 9'5 % of the same. Greatest thick-

tin 17*1 %, of the length of the body. Pectoral tins

equal in length to the base of the anal tin, ventral
fins 15 7 2 % of the length of the body. Distance be-
tween the latter fins and the tip of the snout 87'2 %
of that between the dorsal tin and the same point.
Length of the middle caudal rays 7*5 % of that of
the body. Above the lateral line 10 scales, below

Fig. 202. Bliccopsis rutilo-blicca from the R. Helge at Kristianstad, taken April 17, 1869. Natural size. Among the stuffed

specimens in the Museum of Kristianstad School.

ness 39 % of the greatest depth. Length of the head
21*1 % of that of the body, postorbital length of the
head 47'6 % of its entire length. Distance from the
tip of the snout to the beginning of the dorsal tin
51 '3 %, base of the dorsal tin L 2*3 %, of the length
of the body. Distance from the tip of the snout to
the beginning of the anal tin 62‘2 %, base of the said

it 6, in an oblique transverse row from the inser-
tion of the ventral tin. Scales in the lateral line 47.
Fin-formula identical with that of the preceding
hybrid.

This latter hybrid is thus distinguished from the
White Bream mainly by a thicker body and a shorter
anal fin.

812

SCANDINAVIAN FISHES..

THE BREAM (sw. braxen).

ABRAMIS BRAMA.

Plate XXXIV, fig. 2.

Scales in the lateral line about 53 (51 — 56°). Anal fin with 25 — 27h branched rays. Between the lateral line
and the dorsal fin 12 — 14 (in exceptional . cases 11) rows of scales. Pectoral and ventral fins grayish white at

the base. Pharyngeal teeth set in a single row: 5(6 ) — 5.

Fig. 203. Pharyngeal bones of Abrarnis brarna , a and b natural size; a , left pharyngeal, seen from above; b, the same, seen from without

(the left); c, the last (uppermost) tooth, seen from behind, twice the natural size.

R. br. 3; D.

3 ld 2

• p • V —

25 — 27 b' ' 16—18’ ' 8'

C. x + 1 4- 17 + 1 + x; L. lat. 51 — 56“ ; L. tr. — - 1;

Vert. 43 — 45.

Sijn. Abramis , Belon, Nat., Div. Poiss., p. 318. Cyprinus latus
sive Brama , Rondel., Be Pise. Lacustr., p. 154; Schonev.,
Ichthyol. Slesv. Hols. p. 33. Cyprinus pinnis omnibus ni-
grescentibus, pinna ani ossieulorum viginti septem, x\.rt.,
Ichthyol., Gen. Pise., p. 6; Syn. Pise., p. 4; Descr. Spec.
Pise., p. 20; Lin., Fn. Suec., ed. I, p. 121.

Cyprinus Brama, Lin., Syst. Nat., ed. X, tom. I, p. 326; Bl.,
Fisch. Deutschl., part. I, p. 75, tab. XIII; Retz.. Fn. Suec.
Lin., p. 360; Pall. Zoogr. Ross. Asiat., tom. Ill, p. 325;
Ekstr. (subg. Abramis ), Vet.-Akad. Handl. 1830, p. 169;
Nilss., Prodr. Ichth. Scand ., p. 30; Ekstr., v. We., Skand.
Fisk., ed. I, p. 175, tab. 42; Cuv., Val. ( Leuciscus ), Hist.
Nat., Poiss., tom. XVII, p 9; Kr. ( Abramis ), Danm. Fisk.,
vol. Ill, p. 369; Sundev. {Cyprinus'), Stkhlms L. Hush. Sallsk.
Handl. 1855, pp. 81 et 173; Nilss. {Abramis), Skand. Fn.,
Fisk., p. 324; Hckl, Kn., Siisswasserf. Ostr. Mon., p. 104;
Widegr. {Cyprinus), Fiskfn., Fisker. Norrb. L ., 1860, pp.
7 et 13; Dyb. {Abramis), Cypr. Livl., p. 190; Sieb., Suss-
wasserf. Mitteleur., p. 121; Mgrn, Finl. Fiskfn., disp.

Helsingf., p. 40; Widegr. {Cyprinus), Landtbr. Akad. Tidskr.
1863, pp. 201, 203, 207; Gthr {Abramis), Gat. Brit. Mus.,
Fish., vol. VII, p. 300; Coll., Fork. Vid. Selsk. Chrnia
1874, Tillaegsh., p. 183; Malm, Gbgs, Boh. Fn., p. 565;
Mor., Hist. Nat. Poiss. Fr., tom. Ill, p. 395; Bncke, Fisch.,
Fischer., Fischz. O., W. Preuss., p. 118; Day, Fish. Gt.
Brit., Irel., vol. II, p. 193, tab. CXXXV; Mela, Vert. Fenn.,
p. 332, tab. X; Mob., Hcke, Fisch. Osts., p. 115; NorbAck,
Handl. Fiskev., Fiskafv., p. 433; Reut., Sundm., Finl. Fisk.,
tab. VII; Lillj., Sv., Norg. Fislc., vol. Ill, p. 274.

Cyprinus Farenus, Ekstr., Vet.-Akad. Handl. 1830, p. 175;
Nilss., Prodr., 1. c. ; — juvenem hujus speciei judicarunt
Fries et Ekstr., Skand. Fisk., ed. I, p. 57 (anno 1837).

Abramis vetula, PIeckel; Abramis microlepiclotus + Abr. argy-
reus, Agassiz; Abramis Gehini, Blanchard, vide Siebold, 1. c.

Obs. After the determination given by Fries and Ekstrom of
Artedi’s Faren, later writers — Nordmann {Voyage of Demidoff, 3,
p. 503), Valenciennes, Nilsson, Siebold, etc. — have adopted this
designation as a synonym of Abramis brama. Here as elsewhere,
however, the Swedish name proves the safest guide to the correct
interpretation of Artedi's species, though a slip of the pen (27 rays
in the anal tin, instead of 37) has been overlooked both in Syn. Pise.
and the short diagnosis in Descr. Spec. Pise. See the note to the
following species.

a Sometimes 49, according to Lilljeborg, or even 45 (in specimens 16 cm. long), according to Valenciennes; sometimes 63, ac-
cording to Day.

b 23 — 28, according to Siebold. Sometimes 29, according to Czernay.

c Sometimes 7, according to KrOYER. Sometimes (in our largest specimen) 10, though here the last ray is not divided, in accordance
with the general rule, to the base, and is besides slender, being thus best reckoned as a part separated from the ray in front of it.

. 1

“ — . according to EkstrOm.

15' &

e Sometimes 7, according to KrOyer.

BREAM.

813

The Bream is one of the largest Swedish Cypri-
noids. The ordinary length, including the whole cau-
dal tin, is 23 — 35 cm.; but it is not very unusual to
meet with Bream 5 dm. long, and according to the
reports sent in to the Swedish Fisheries Commission
of 1881 — 83 the species attains in many parts of Swe-
den a length of about 59 cm. and in Scania even of
74 cm. The weight, on the same authority, is some-
times 1 8 1 /2 Sw. lbs. (nearly 8 kilo).

The body is compressed, thin, and deep, the
greatest depth, which occurs at the beginning of the
dorsal fin, being about 34 — 38 % of the length, and
the greatest thickness about 27 — 36 % of the greatest
depth. The least depth, just in front of the caudal
fin, is about 9J/2 (in young specimens sometimes 973)
— 10y3 % of the length of the body. The dorsal pro-
file, which rises more or less sharply from the occiput,
most abruptly if the fish be fat, from this point forms
a regular curve to the first ray of the dorsal fin, where
it makes an obtuse angle, afterwards descending almost
in a straight line to the base of the caudal fin. The
back is compressed and thin, but not carinated, all the
way to the dorsal fin; from this fin to the caudal it
is somewhat plumper and convex. The belly, as usual,
is flat to the ventral fins and carinated from this point
to the anal aperture. From the isthmus to the ventral
fins the ventral line is almost straight, but slopes a
little towards the anal aperture, where it forms an angle
more obtuse than the dorsal; from the vent it rises al-
most in a straight line to the end of the anal fin, be-
tween which point and the caudal fin it curves inwards.

The head is as a rule equal in size to that of the
White Bream, measuring 2 1 a/2 — 20 % of the length of
the body; but in young specimens, even those about 2
dm. long, its length may be 23 or 24 % of that of the
body. It is so much compressed that the greatest thick-
ness, across the opercula, is about equal to the post-
orbital length, which in young Bream is somewhat less,
in old rather more, than half of its entire length. The
forehead from the occiput to the nostrils is straight,
broad, and convex, the breadth of the interorbital space
being about 8 — 9 % of the length of the body or 36 — 41
% of the length of the head. Both on the forehead and
the sides of the head the pores belonging to the system
of the lateral line are quite distinct. From the occiput
two parallel rows of pores (the frontorostral branches
of the system, cf. fig. 104, p. 368) run along the fore-
head to a point almost vertically above the eyes: these

pores are 8 or 9 in number, and the first two or three
lie somewhat within the line, as if they composed an
independent row. Another row of similar pores (the
suborbital branch of the system), the number of which
is still greater, also begins at the occiput, a little be-
low the former. This row bends up towards the fore-
head above the cheek, then descends below the eye,
and ends half-way between the tip of the snout and
the nostrils. Similar pores also occur at the posterior
and inferior margins of the preopercula and on the
under surface of the lower jaw (the mandibular branch
of the system). Above the nostrils lies the depression
usually present in the Cyprinoids, terminating the mo-
bile portion of the snout (see above), which is obtuse
and projects only slightly beyond the mouth. The
mouth is small and turned slightly upwards, but when
it is opened, may be protruded to a considerable di-
stance in a tubular form. When the mouth is closed,
the upper jaw projects some way beyond the lower.
The corner of the mouth lies in the perpendicular from
the anterior margin of the nostrils. In this species too
the length of the snout, which varies between about 6
and 7 1 /2 or nearly 8 % of the length of the body, or
between 28 and 38 % of that of the head, is as a rule
in young specimens about equal to, in old somewhat
greater than, the length of the upper jaw from the
middle of the tip of the snout. The length of the lower
jaw varies between about 8 and nearly 10 % of that
of the body, or between about 35 and 45 % of that of
the head, and is greater than that of the suture be-
tween the suboperculum and the operculum, and also
more than 75 % of the least depth of the tail (76 —
95 % in our specimens, which are between 17 and 47
cm. long). The eyes are as a rule comparatively smaller
than in the White Bream, their longitudinal diameter
varying in our specimens between about 28 and 19 %
of the length of the head. But this great reduction
depends to a considerable extent on the fact that our
largest Bream are more than twice the size of our
largest White Bream. The position of the eyes, on the
other hand, is essentially the same as in the White
Bream. The nostrils, which in form are exactly like
those of the White Bream, occupy in young Bream the
same position as in the White Bream, but in old spe-
cimens are removed by the prolongation of the snout
comparatively nearer to the eyes. The gills, the oper-
cula, the branchiostegal membrane and rays, resemble
those of the White Bream. The gill-rakers are more

814

SCANDINAVIAN FISHES.

numerous than in the said species, their number in the
outer row on the front of the first branchial arch being
22 or 23, on the outer anterior margin of the pha-
ryngeals 15 or 16. The pharyngeal teeth (fig. 203)
are always set in one row, except during the shedding
of the teeth, when one or two may be found in the
gums within the main row, which usually consists of
5 compressed teeth, obliquely truncate at the crown,
with hollowed masticatory surface and slightly hooked
tip. The pseudobranchiae are distinct in young Bream,
in old concealed.

The dorsal fin begins at the apex of the angle
formed by the dorsal profile, almost vertically above
the middle point in the postabdominal part, and at a
distance from the tip of the snout which increases with
age, measuring about 52 — 58 % of the length of the
body, or (in our specimens) about 88 — 94 % of the
distance between the tip of the snout and the anal fin.
In this species too its height is about twice its length
— longest (first branched) ray about 21 — 26 % (some-
times 27 %) of the length of the body, and the base
about 1 1 1/2 (sometimes 11) — 1272 % of the same. The
upper posterior margin of the fin is so obliquely trun-
cate that in front the height is more than three times
as great as behind. The anal fin is of about the same
form, and occupies about the same position as in the
White Bream; but is generally somewhat longer. In
proportion to the length of the body the distance be-
tween this fin and the tip of the snout is about 59 —
62 %, its base about 241/ 2 (sometimes 24) — 27 %, and
its height in front about 20- — 17 % (in our largest spe-
cimen 1572 %). The caudal fin is deeply forked, the
lower lobe being longer than the upper. The middle
rays measure about 10 — 772 0/o of the length of the
body, or about 1/.i of that of the longest ray in the
lower lobe of the fin.

The tips of the pectoral fins extend (in the males
and sometimes in the females) to the insertions of the
ventrals or a little farther, the length of the former
pair being about 20 — 2 1 1/2 % of that of the body. The
ventral fins are set at a distance from the tip of the
snout measuring about 44 — 42 72 % of the length of
the body, or less than 83 % (82 — 74 %) of the distance
between the tip of the snout and the dorsal fin. Their
length is about 17 % (16— 1 7 x/2 %) of that of the body.
The postabdominal length is always less than the pre-

abdominal, the former varying between about 17 and
20 %, the latter between about 23 and 20 %, of the
length of the body.

The scales are large, finely striated, broader than
long, and of a rounded quadrangular shape, with the
free (hind) part more than twice as deep as long, and
the anterior (inserted) margin irregularly undulating or
even smoothly rounded. They are densely imbricated
in regular rows, and the lateral line runs much nearer
to the belly than to the back.

The coloration, which varies greatly according to
the age of the fish and the water in which it lives, is
darker in old and fat specimens than in young and
lean. In a full-grown fat Bream the upper parts of
the head and the back are yellowish gray, suffused
with a brassy lustre most distinct on the sides, which
are lighter and become yellower towards the belly.
The belly is of a more or less whitish yellow. The
opercula and cheeks are yellowish white with a brassy
lustre. The lips are white. During the spawning-
season the scales of the milters are studded on the front
part of the sides and the head with whitish tubercles.
All the fins are blackish gray. The pectoral and ventral
fins have lighter, almost white bases; but these parts,
as well as the belly, become suffused with blood on the
death of the fish (cf. above, on the Sheatfish). The
iris is yellow, strewn with extremely fine, dark dots,
and is divided from the pupil by a ring of still deeper
yellow.

North of the Alps the Bream is more or less com-
mon in most of the great European lakes, in places
where the water is clear, and where at least a few
shelving, wreedy shores are to be found. It also lives
in large rivers where the current is not too strong.
In such waters it occurs in all the provinces of Sweden
save Gothland". In Finland it is equally common, and
according to Mela goes north to lat. 67° 25'. It does
not thrive in the highlands, and in Lapland and even
in Jemtland it is rare. In Norway, according to Col-
lett, it occurs only in the south-eastern tracts of the
country, hardly going north of lat. 61°. Though it
does not attain the same size in salt water, it is found
in the inner parts of the Baltic island-belts.

The favourite haunts of the Bream invariably lie
in clear and moderately deep water with a weedy
bottom, especially where it can find Merlirfs-grass h

“ Cf. UndercL. Bet. m. Forslag till Ny Fiskeristadga, Stklilm 1883, p. 154.
b Sw. Braxengrds , Bream-grass.

BREAM.

( Isoetes lacustris), which it roots up like the pig. Float-
ing bits of this weed betray to the fisherman the re-
sorts of the Bream. Though during summer the Bream
stays in water of a moderate depth, adult individuals
never ascend to the shores except in the spawning-
season. Only small Bream repair to the shallows in
early spring and remain there all the summer. Late
in autumn, when the weather is very stormy, the Bream
is sometimes driven into shallow inlets, and is then
known by the fishermen as Ovddersbraxen (Storm
Bream). The individuals that ascend rivers during the
spring spawning and in summer are generally young.
Their habits also differ from those of their fellows in
general, for the shoals of Bream do not move against
the stream, but always swim with the current. As the
autumn draws to a close, the Bream retires to deep
water, and chooses its winter-quarters in the deepest
place it can find. Here it lies packed in countless
multitudes, and such places are called Braxenstand
(Bream-stands). A famous pool of this description lies
in Lake Hallbo in Sodermanland, and has annually
yielded a take, at the same spot and at a single haul,
of 4,250 — 17,000 kgrm“.

The Bream is cautious, cunning, gregarious, and
very timid. It is seldom found alone, but almost
always in large or small companies. The best proof
of its cunning is its habit of burrowing in the mud
or lying on one side that the seine may pass over
it. Loud noises, thunder, the sound of bells, the re-
port of firearms, etc., always drive it into deep water,
from which it does not return for several days. Its
tenacity of life is such that, embedded in grass, it may
be conveyed alive for considerable distances. A method
long employed in order to keep the fish alive during
as long a transportation as possible, is to place in its
mouth a bit of bread previously dipped in spirits.

The Bream may be planted in ponds, where it
thrives well and grows fat if not stinted in food. In
a state of nature the fish lives on weeds, mud, worms,
and insects. In the stew it may be fed and fattened
on brewers’ grains, pellets of dough, and the like.

Like the other Swedish Cyprinoids the Bream is
subject to great variations of form and colour, these
depending on the age of the fish, the season of the
year, the nature of the water, etc. The difference is
often so great that some ichthyologists — as shown by

“ Cf. Lloyd, Scandinav. Advent., vol. I, p. 45.

815

our list of synonyms — have been in doubt whether
these forms did not constitute distinct species. If the
supply of food is plentiful, and the fish becomes fat,
the body is always deeper in relation to its length than
in specimens which are lean for lack of food. In the
latter case the body is more elongated, 'with more
pointed head, and the back usually rises in a less sharp
curve. Young specimens are always much shallower
than old, and more elongated in form. At this age
they are regarded by the fishermen in certain localities
as a distinct species, and are called Banka, Bldpanka,
Flira, Svartspoling, etc. The colouring too, as we have
mentioned above, is much darker in fat Bream than in
lean or young specimens.

At the end of May or beginning of June the Bream
repairs to the shores in order to spawn. The spawning
always takes place when the juniper is in flower, a
circumstance which is duly observed by the fishermen.
The first shoal to arrive at the spawning place, which
is never changed, being the same from year to year,
consists of males alone. The females come later to
join their mates, and the spawning now begins, the
silent hours of night being preferred for this purpose.
The operation is accompanied with great noise, for the
fish rove to and fro at the surface in dense shoals,
lashing the water with their tails and displaying their
activity in many ways. The roe is deposited on rushes
and weeds, against which the female rubs her body
while spawning. The ova are small and yellowish.
The spawning last 3 or 4 days, according to the wea-
ther. The eggs are hatched in three weeks, and the
fry grow quickly. When the old Bream have finished
spawning, the young fish commence. If no convenient
spawning-place can be found in a lake, the shoal ascends
some large stream in quest of a suitable spot. In the
latter case the choice always falls on some weedy bight
at the side of the channel.

The Bream is very prolific. Bloch counted 137,000
eggs in a female weighing 6 lbs. (2,811 grammes), and
Benecke estimates the number at 200,000 — 300,000
ova about 1 1/2 mm. in longitudinal diameter.

The flesh of the Bream, though bony, being of
good flavour, the fisherman has many different methods
of taking this species. During the spawning-season gill-
nets are generally used, the seine being less suitable,
as the fish are frightened by its use, and often disturbed

103

Scandinavian Fishes.

816

SCANDINAVIAN FISHES.

in their spawning operations. The katsa (tig. 204), a
fish trap set at the spawning-place or in the channels
which the Bream must pass on its way thither, is a

often caught, as we have mentioned, in ice-seines, when
the winter- quarters of the fish have been discovered.
In August and September the Bream is taken by ang-

Fig. 204. Katsa , a fish trap in general use in Central Sweden, and chiefly employed in the taking of Cyprinoids. Seen obliquely from
the side with the shore-arm to the right, and in the plan with the shore-arm turned downwards.

better kind of tackle, the fish being thus afforded an
opportunity of depositing their roe on the shore-arm
of the trap. In winter great numbers of Bream are

ling in 6 — 12 fathoms of water, the bait used being
the common earthworm.

(Ekstrom, Smitt.)

A form which has long been known, but was first
recognised in scientific nomenclature by Bloch, the
German Letter and Yarrell’s Pomeranian Bream, has
been explained by Siebold and other later writers as
a hybrid between the Bream and the Roach. For the
sake of uniformity and on the analogy of Bliccopsis,
however, Siebold gave this form the generic name of

Abramidopsis. The most striking character — a com-
paratively short anal fin in conjunction with a Bream-
like body — is common to Abramidopsis and Bliccopsis ,
and we therefore consider it most appropriate to com-
pare this form in our diagnosis thereof with the said
hybrids of the White Bream.

ABRAMIDINES.

817

THE POMERANIAN BREAM.

ABRAM J DOPSI S BUGGENH AG II.

(hybrid between the bream and the roach.)

Fig. 205.

Body slialloiver than in the Bream, the greatest depth being about 31 — 33 % of the length. Dorsal margin in
front of the dorsal fin compressed, but covered with, scales in the median line. Dorsal fin with 10 branched
rays, anal with 15 — 18. Distance from the tip of the snout to the dorsal fin slightly less than half the length
of the body, but perceptibly greater than that to the foremost point in the insertion of the ventral fins. At least
10 scales in an oblique transverse row between the beginning of the dorsal fin and the lateral line. Pharyngeal
teeth set in a single row , hooked at the tip, but with smooth masticatory surface: G or 5 — 5.

Fig. 205. Abramidopsis Baggenhagii, -/3 of the natural size, from Dalecarlia, Steffenburg. The property of the Zoological Museum of
Upsala University, a, the left lower pharyngeal, seen from above, natural size; b, the same, seen from without, natural size;

C, the hindmost tooth, twice the natural size, a, b, and c from a somewhat smaller specimen in the Royal Museum.

B. br. 3; D. — ;

10

A.

3

15 — 18’

P.

9

C. x + 1 + 1 7 + 1 + x; L. lat. 44 — 51a; L. tr.

10 — 11
1.

5— G

Byn. Cyprinus Buggenhagii, Bl., Fisch. Deutschl., part. Ill, p. 137,
tab. xcv ; Cuv., Yal. ( Leuciscus ), Hist. Nat. Poiss., vol.
XVII, p. 53; — (?) Thomps. ( Abramis ), Nat. Hist. Irel.,

vol. IY, p. 137; Ytarr., Hist. Brit. Fish., ed. 2, vol. I, p.
391; Couch, Fish. Brit. Isl., vol. IY, p. 42, tab. C'LXXXIX;
— Mor., Hist. Nat. Poiss., Fr., tom. Ill, p. 400.

Abramis Leuckartii, Hckl, Ann. Wien. Mas. Naturg., Bd. I,
p. 229, tab. XX, fig. 5; Nordm., Voy. Pass. Mer. (Demid.),
tom. Ill, p. 508; Cuv, Yal., 1. c., p. 59; Hckl, Kn., Siiss-
ivasserf. Ostr. Mon., p. 117; Sieb., ( Abramidopsis ), Siiss-
wasserf. Mitteleur., p. 134.

45 — 54, according to Siebold.

818

SCANDINAVIAN FISHES.

Abvamis Heckelii , Sel. Longch., Fne Beige , p. 217.

Hybrid between Abramis brama and Leuciscus rutiJus, Gthr,
Cat. Brit. 9 fits., Fish., vol. VII, p. 214; Lillj. (p. p.), Sv .,
Norg. Fn., Fisk., vol. Ill, p. 287.

Of this form, which according to Siebold attains
a length of about 31 cm. (to the extreme tips of the
caudal lobes), we have been in a position to examine

three specimens 17 — 22 cm. long (to the end of the
middle caudal rays), all three taken in Dalecarlia by
Dr. Steffenburg, and two others, resp- 20 and 32
cm. long, from the Helge River (Scania). Instead of a
description at length we shall here give those average
measurements of the first-mentioned three specimens
which express the most important relations of form.

Length of the body expressed in millimetres.

Length of the head in % of the length of the body

Greatest depth of the body „ „ ,, „ „ „ „

Greatest thickness of the body i „ „ „ „ „ „ „ „

Least depth of the tail „ „ „ „ „ „ ,, „

Distance between the dorsal tin and the tip of the snout ,, ,, „ „ „ „ „ „

Length of the base of the dorsal fin „ „ „ „ „ „ „ „

„ „ „ longest ray of the dorsal fin „ „ „ „ „ „ „ „

Distance between the anal fin and the tip of the snout „ „ ,, „ „ „ „ „

Length of the base of the anal fin „ „ „ „ „ „ „ „

„ „ „ longest ray of the anal fin „ „ „ „ „ „ „ „

„ „ „ middle rays of the caudal fin „ „ „ „ ,, „ ,, ,,

,, ,, 5, longest rav ,, ,, ,, ,, - ,, ,, „ ,, ,, ,5 ,, ,,

55 55 55 pectoral fins — — — ,, ,, ,, ,, ,, ,, ,, ,,

„ „ „ ventral „ „ „ „ „ „ „ „ „

Distance between the ventral fins and the tip of the snout „ „ „ „ „ „ „ „

Preabdominal length „ „ „ „ „ „ „ „

Postabdominal ,, — — ,, ,, ,, ,, ,, ,, ,, „

Length of the peduncle of the tail behind the perpendicular from the termination of the base of the anal fin „ „ „ „ „ „ „ „

Diameter of the eyes... ,, „ „ „ „ „ „ head

Length of the snout „ „ „ ,, „ „ „ ,,

Breadth of the interorbital space „ „ „ ,, ,, „ „ „

Length of the upper jaw from the tip of the snout ,, „ „ „ ,, „ „ „

55 55 55 lower ,, ,, ,, ,, ,, 5, ,, ,, ,,

„ ,, „ suboperculum at the upper margin „ „ „ „ „ „ „ „

193

21.9

31.8

(9.3)

10.5

49.7

13.5

(23.0)

62.8

17.2
(15.5)

9.4

(25.4)

(18.1)

(16.5)

42.8

20.9

(20.5)

15.9

23.9
(30.3)

37.5

29.5

35.2

35.6

The characters of the Roach here find expression
in the comparatively short lower jaw and pectoral fins, |
in the forward position of the dorsal fin and its com-
paratively long base, and in the comparatively great
postabdominal length. The influence of the Bream type
may be traced, on the other hand, not only in the
great depth of the body and, relatively, of the pe-
duncle of the tail, but also in the height of the dorsal
and anal fins and in the comparatively great preab-
dominal length.

Bloch described this form from Swedish Pomerania,
Heckel and Siebold from Central Germany and Austria,
Nordmann from the Lower Danube, Valenciennes from
France, Selys-Longchamps from Belgium, Yarrell and
Gunther from England. According to Thompson it
also occurs in Ireland; but this statement seems to re-
quire further corroboration, for, as Day" has remarked,

the Roach does not belong to the Irish fauna. In Swe-
den this hybrid had as yet been found, to the best of
our knowledge, only in Dalecarlia and, according to
Lilljeborg, in the Klar Elf, until we received very re-
cently (May, 1893), through Fisherman Billing of Kris-
tianstad, two splendid specimens from the broad below
that town on the lower course of the River Helge.

The German name of Letter (Leader) has been con-
ferred on this hybrid as well as on several other more
or less singular fishes that occur in solitary specimens
among a shoal of some common species, and are there-
fore supposed to pilot their comrades. A catch of this
kind. is enough to give the fisherman hope of a plenti-
ful take of Bream.

Like the Cyprinoid hybrids in general this form
too is prolific, the spawning-season occurring in Ger-
many, according to Siebold, at the end of April.

Fish. Gt. Brit., Irel ., vol. II, p. 195.

ABRAMI DINES.

819

THE ZOPE (SW. FLIRAN OR faren).

ABEAM IS BALLERUS.

Plate XXXIV, fig. 1.

Scales in the lateral line about 70 ( 69a — 73). Anal jin with 34 — 42 branched rays. Between the lateral line
and the dorsal Jin 14 — 16 rows of scales. Mouth ascending , with the tip of the lower jaw slightly projecting.

Pharyngeal teeth set in a single row: 5 — 5.

R. hr. :

1

3

P.

' 34

—42’

15

.v + 1 + 17

+ 1 + x ;

L. lat.

69“—

-73;

L.

V.

7—8’

14—16

8—10(11)

i;

Vert. 47—48.

Syn. Cyprinus admodum latus et tenuis; pinna ani ossiculorum
quadraginta, Art., Iclithyol., Gen. Pise., p. 3; Syn. Pise.,
p. 12; Descr. Spec. Pise., p. 23 4- Cyprinus iride flava:
pinna ani ossiculorum triginta (viginti) septem, ibid., Gen.,
p. 3; Syn., p. 13; Spec., p. 23; Cyprinus pinna ani ossicu-
lorum quadraginta, Lin., Fn. Suec., ed. I, p. 122 + Cypri-
nus pinnae ani radiis triginta septem, ibid., p. 123 + Cypri-
nus pinnae ani radiis viginti quinque (p.p.), ibid., p. 124, obs.
Cyprinus Farenus , Lin., Syst. Nat., ed. X, tom. I, p. 326;
Retz., Fn. Suec. Lin., p. 361.

, Cyprinus Dnllerus , Lin., Syst., 1. c.; Retz., 1. c., p. 361; Bl.,
Fisch. Deutschl., part. I, p. 62, tab. IX; Pall., Zoogr. Boss.
Asiat., tom. Ill, p. 327 ; Nilss., Prodr. Ichthyol. Scand.,
p. 30; Ekstr., v. We., Skand. Fisk., ed. I, p. 112, tab.
26; Cuv., Val. ( Leuciscus ), Hist. Nat. Poiss ., vol. XVII,
p. 45; Lillj. (Cyprinus), Vet. Akad. Handl. 1850, p. 305;
Kr. ( Abramis ), Damn. Fisk., vol. Ill, p. 411; Sundev.
(Cyprinus), Stockh. L. Hush. Sallsk. Handl. 1855, p. 81;
Nilss. (Abramis), Skand. Fn., Fisk., p. 331; Lloyd
(Cyprinus), Scand. Adv., vol. I, p. 51; Kessl. (Abra-
mis), Bull. Soc. Natar. Mosc., tom. XXIX: 1 (1856), p.
377; Hckl, Kn., Susswasserf. Oestr. Mon., p. 113; Dyb.,
Cypr. Livl., p. 196; Sieb., Susswasserf. Mitteleur., p. 130;

Widegr. (Cyprinus), Landtbr. Akad. Tidskr. 1863, p. 203;
Gthr (Abramis), Cat. Brit. Mus., Fish., vol. VII, p. 302;
Malm, Gbgs, Boh. Fn., p. 566; Bncke, Fisch., Fischer.,
Fischz. 0 ., IE. Preuss ., p. 122; Mob., Hcke, Fisch. Osts.,
p. 116; Grimm, Fish., Hunt. Puss. Wat., pp. 15 et 22;
Lillj., Sv., Norg. Fn., Fisk., vol. Ill, p. 294.

Cyprinus Biorkna (neo synon.), Lin., Fn. Suec., ed. II, p.
130; Retz., 1. c., p. 360.

Obs. When Ekstrom wrote his Morko Fiskar and Nilsson his
Prodromus, they had both received from Lake Malar young Bream
under the name of farnar (Zope); and the parenthetic remark in
Artedi’s diagnosis of the Faren (Gen. Pise.) that in Syn. Pise, (as
well as in the diagnosis of Faren in Descr. Spec. Pise.) a misprint
had escaped observation (27 anal rays instead of 37), was supposed
to be an insertion of no importance made by Linnjeiis. Up to the
present day EkstrSm’s followers have therefore regarded the Cyprinus
Farenus of Artedi and Linnzeds as a young Bream'. But Artedi’s
minute description of this fish plainly shows that he really referred
to the form still known by most of the Malar fishermen as the Faren.
Not only do we read in Clause 12 of this description “pinna ani . . .
ossiculorum triginta septem”; the measurements given by Artedi
(Longitudo tota ad initium caudae unc. 8, lin. 8; ad pinnam dorsi
unc. 4, lin. 8; ad pinnam ani unc. 5, lin. 3°) demonstrate beyond
doubt that his specimen was a Faren. The following table contains
our measurements of Bream and Zope preserved in spirits — the di-
mensions given by Artedi were probably those of fresh specimens —
ranged in parallel columns with Artedi’s measurements of the Bream
and Zope in Descr. Spec. Pise., pp. 22 and 25.

“ Sometimes 65, according to Valenciennes.
b 7 — 9, according to Kroyer.

c Whether these measurements are duodecimal (12 lines to the inch) or decimal (10 lines to the inch), I am unable to determine;
but the difference is of no importance to the solution of the question. For the sake of completeness I have included in the table (in paren-
theses) the percentages which the decimal system would involve.

820

SCANDINAVIAN FISHES.

Length of the body from the tip of the snout to the base of the caudal fin, ex-
pressed in millimetres

Distance between the dorsal fin and the tip of the snout in % of the above length of the body
„ ,, „ ventral fins „ „ „ „ „ „ „ „ „ „ „ „ ,, ,, „

„ „ „ anal fin „ ,, „ „ „ „ „ „ ,, „ „ ,, „ „ ,,

Royal Museum specimens
Abramis brama.

of

Artedi’s

Royal Museum
Abramis

specimens

ballerus.

of

I slat) (1-belt of Stock-
holm, Sept., 1890.

Island-belt of Stock-
holm, Sept., 1891.

Tsland-belt of Stock-
holm, Sept., 1890.

Lake Oppmanna,
Sept.., 1882.

tr

zr pr
n rs

r o

GO c

cc 5
a

B

'ZX

1 r

cr. b

CC P
LO

Average
in the
preceding
six

specimens.

S

rs

S

Average
in the
following
four

specimens.

CH C/2

~Loq’

S'

i— - 1 ^
CX

cy

o

a

a>

M-

CX

cr.

Sigtuua (Millar)
July 11, 1831.

ZX

> b
^ aq
2. B

| — 1

cr. ^

cr. ~

H_L 2^

152

166

169

180

391

424

247

330

216

189

92

199

203

263

63.4

59.4

60. o

59.4

63.0

63.o

6 1 . 4 ( > 59)

60.0(59.5)

54.3(54.0)

52.9( < 55)

53.2

52.8

52.2

53.2

46.7

48.2

47.9

51.7

46.5

47.o

48.o(>46)

48.7(48.5)

42.8(42.3)

40.9( < 43)

42.9

40.4

39.9

40.3

b4.5

65.5

66.2

65.8

66.7

67.9

6G.i( 64)

67.5(67.2)

60.0(59.7)

57.o(<61)

59.8

56.8

57.i

58.i

No further evidence seems necessary to show that Artedi’s and
Linnaeus’s Fareri and Braxnaflicka ( Blicka , which they erroneously
identified with Rondelet’s Ballerus ) and Linnaeus’s (not Artedi’s)
Bjorkna , all belong to the same species. The specific name of fare-
nus has consequently as great claims to recognition as ballerus , and
has the advantage, as compared with the latter, of not being based
on a false explanation of names already in use.

The ordinary length of the ‘Zope’ is between 20
and 25 ein., but may rise, according to Lilljeborg, to
45 cm., including the whole caudal fin.

The body is strongly compressed and deep, but
more elongated than that of the Bream, the greatest
depth, which occurs at the beginning of the dorsal fin,
being about 29 % (27 ’/, — 31 %) of the length of the
body, measured to the end of the middle caudal rays.
The greatest thickness, which lies as a rule across the
opercula, is less than 1/3 of the greatest depth, and at
most half of the length of the head. The least depth
of the tail is about 81/, — 8 % of the length of the body.
The back, which has only a faint break at the occiput,
then forms a regular curve to the beginning of the dor-
sal fin, where it makes a very obtuse angle, and from
this point slopes almost in a straight line to the base
of the caudal fin. It is moderately compressed, with
a faint, convex, longitudinal ridge, especially near the
dorsal fin. The part between the dorsal and caudal fins
is somewhat plumper. The belly is of a broad convex
shape or flat from the isthmus to the ventral fins, from
this point to the vent sharply carinated (with the very
edge, as in the other true Abramidines, naked, i. e.
not covered, although concealed, by the scales at its
sides), and then very strongly compressed and thin
throughout the length of the base of the anal fin. The
ventral profile forms an unbroken curve from the chin
to the anal aperture, then runs in a straight line to
the end of the anal fin,_ and finally bends slightly in-
wards to the base of the caudal fin.

The head, the length of which is about 20 — 19 %
of that of the body, is comparatively smaller than in
the preceding Abramidines. It is pointed, but the snout
is thick and blunt, and does not project beyond the
tip of the lower jaw. The forehead is broad and con-
vex, with straight frontal line and a slight slope from
the nostrils to the tip of the snout. The breadth of
the interorbital space measures about 2/s (39 — a little
more than 40 %) of the length of the head. The sides
of the head are so compressed that its greatest thick-
ness is equal to the perpendicular drawn through the
anterior orbital margin. The mouth is not large, but
turned sharply upwards and protruded, when the mouth
is opened, in a tubular form. The anterior margin of
the upper jaw is somewhat less prominent than that of
the lower, and the length of the upper jaw from the
tip of the snout to the hind extremity of the maxil-
laries is in young specimens somewhat greater than
that of the snout to the anterior margin of the eyes,
in old equal to, or somewhat less than, the latter, and
measures about 28 — 29 % of the length of the head.
The lower jaw is valvular in shape, contracted at the
base, expanded in front; it points upwards, and is ar-
ticulated in the perpendicular from the anterior margin
of the eyes. Its length is as a rule equal to the breadth
of the interorbital space. The eyes are rather large,
their longitudinal diameter varying in specimens 10 —
29 cm. long between about 32 and 26 ‘/2 % of the length
of the head. They are set fairly low, the upper mar-
gin of the pupil (sometimes of the iris) being touched
by the line joining the margin of the upper jaw at the
tip of the snout to the middle of the tail at the base
of the caudal fin. The postorbital length of the head
measures in young specimens somewhat less than half
its entire length; in old specimens these two measure-
ments are equal to each other, and the postorbital part

ZOPE.

821

is twice as long as the snout. The nostrils are similar
to those of the preceding forms, but lie a little higher
in relation to the eyes. The gill-openings are large,
the branchiostegal membranes coalescing with the isth-
mus rather near each other, just behind the perpendi-
cular from the hind margin of the eyes. The gill-
rakers are long, thick-set, and pectinated, with more
or less distinct serrations. The outer row on the first
branchial arch contains about 30 — 37, that on the outer
anterior margin of the pharyngeals 24 — 26. The pha-
ryngeals, which are remarkably thin, are each furnished
with a row of five teeth, these being compressed,
obliquely cut, and hooked at the tip, with concave
(one-grooved) masticatory surface.

The dorsal fin begins at a distance from the tip
of the snout measuring 47 — 49 % of the length of the
body. It is very obliquely truncate, with straight, or
even slightly convex margin, and its height in front is
more than twice its length, the longest ray measuring
about 20 — 21 % of the length of the body, the base
about 9 % of the same. The anal fin is very long,
highest in front, then sinking in a small hollow, and
afterwards proceeding with the margin almost straight.
Its beginning lies at a distance from the tip of the snout
measuring about 51 — 53 % of the length of the body;
the length of its base is about 1/3 (32 1 ,/2 — nearly 35 %),
and that of its longest ray about 13 x/2 — 15 %, of the
same. The caudal fin is deeply forked, the lower lobe
being the longer; its middle rays measure about Vio
of the length of the body and about 2/s of that of the
longest caudal rays.

The pectoral fins are long and pointed, and their
tips extend, when the fins are at rest, sometimes
scarcely to the perpendicular from the insertion of the
ventral fins, sometimes a little further back. The fins
are longest as a rule in the males; but this rule is
subject to exceptions which render it useless as a sexual
character. Their length is about 1/e (16'/2 — nearly
19 %) of that of the body. The tips of the ventral fins
commonly extend, at least in the males, quite to the
anal aperture. The length of these fins is generally
greatest in the males, and measures about 13 — 15 % of
of that of the body. The distance between the tip of
the snout and their insertion is greatest as a rule in
the females, and occupies about 36 — 37 % of the length
of the body. Both the preabdominal length and the
postabdominal measure 16 (l51/2) — 18 % of the length
of the body.

Idle body is covered with comparatively small and
thin scales, set in very oblique rows and densely im-
bricated. They are in general more rounded than the
scales of the Bream; but the true distinction lies in
their still finer concentric striation and the smaller
number of radiating grooves.

The coloration is indeed subject, as usual, to great
variations dependent on the age of the fish and the
season of the year; but it closely resembles that of young
Bream. The sides of the head display many colours,
shot with silver and brassy yellow. The silvery white
iris is dashed with yellow above the pupil, where the
fine, green dots with which it is strewn are collected
in a dark spot. The forehead is olive gray, and a
somewhat lighter shade of the same colour extends to
the back and the upper part of the sides, gradually
fading below and passing, somewhat above the lateral
line, into silvery gray. This last colour grows lighter
and lighter towards the belly, which is almost white.
The dorsal, caudal, and anal fins are of a plain olive
gray, though the outer margin, especially of the last-
mentioned fin, is dark. The pectoral and ventral fins
are light gray with a dash of brown, red, or yellow
at the base.

In its habits the Zope seems nearly allied to the
Bream, ft occurs in our largest lakes, Malar and We-
ner, but is common only in the former, which appears
to be its true habitat in Sweden. Sometimes it descends
the River Gotha from Lake Wener (Malm), and it, no
doubt, wanders from Lake Malar, as Lilljeborg has
remarked, out into the island-belt of Stockholm (Sunde-
vall). But in Sweden it does not seem to belong to
the common fishes of the island-belt. The case is the
same in Finland, though to the south of this country
the Zope is common, according to Mela, in the interior.
In the south of the Baltic, however, in the German
Half’s and in the Gulf of Riga off the mouth of the
Dima, it seems to lie a stationary fish (Mob., Hcke).
According to Lixxnsus it occurs in Smaland, being there
known by the name of Braxenflidca; but no trust-
worthy information on this head has since been pro-
cured, and the name may easily be due to a confusion.
In the River Helge at Kristianstad, on the other hand,
two specimens about 3 dm. long were taken in 1869,
the first in March, the second in May; and they are
now preserved among the stuffed collections in the mu-
seum of the said town. Lilljeborg assumes that these
examples were stray visitors from the south or east of

822

SCANDINAVIAN FISHES.

the Baltic; but their occurrence in the lower course of
the Helge, which there widens into broads, finds a pa-
rallel in many places, the Zope often ascending rivers
both from the Baltic and the Black Seas. The Zope is
further known from the lower parts of the Rhine, We-
ser, and Elbe (Siebold), from the above-mentioned
German rivers and Half’s connected with the Baltic, from
the Danube, up to Hungary and the Hungarian lakes
(Hckl., Kn.), from upper Austria (Sieb.), from the whole
of Russia-in-Europe, with the exception of the river ba-
sins sloping to the Arctic Ocean, and from the Black
and Caspian Seas (Pallas, Kessler, and Grimm).

The temperament of the Zope seems in the main
to resemble that of the Bream. Its diet, however, in-
cludes less vegetable substances, and consists more of
insects and worms. The spawning-season occurs in
April and May, much earlier than that of the Bream,
and sometimes, according to Baron Gyllenstjerna as
quoted by Nilsson, even before the ice has melted in
the inlets of Lake Malar. The spawning-place is chosen
in shallow water with a weedy bottom. At this season
the Zope often ventures even upon flooded meadow
land. The operation of spawning is performed as we
have described in the case of the Bream, but is said
to last no more than 1 — 3 days, during which time the
Zope is no less afraid of noise than the said species.
The course of its development and its growth are little
known; but of its fecundity Bloch states that in a fe-
male weighing 468 V, grin., with ovaries 164*7 grm. in
weight, the number of the ova, which were of the size
of a poppy seed, was about 67,500.

When the spawning is over, the Zope retires to
deeper water, and is seldom found near shore through-
out the rest of the year, but only in the depths. In

autumn and winter it assembles in large shoals, like
the Bream, where it can find a deep pool.

The flesh of the Zope, is flabby, white, bony, and
of sweetish, indifferent flavour, being poorer in summer
than in winter, and fattest previous to the spawning in
April. Still the fish is eaten fresh, in which case it is
boiled, and is also salted and dried for future con-
sumption.

During the spawning-season, at which time the
Zope, as we have mentioned, approaches the shore and
ascends into the shallows, it is taken in traps ( ryssjor ,
see p. 33, fig. 7), nets, and seines hauled from the shore,
where the bottom admits of this operation. It is occa^
sionally speared by accident. Late in autumn it is
sometimes caught in a kind of seine shot in deep water,
the hauls being always large, if made at the right
place, as the fish have now assembled in their winter-
quarters. The Zope may also be taken with rod and
line baited with a common earthworm.

On the south coast of Lake Malar throughout its
length the Zope is called Fame or Fana, in the central
and western parts of the lake Faren, and in the neigh-
bourhood of Upsala, according to Lilljeborg, Vimma.
In Lake Wener, according to Lloyd, it bears the name
of Ldngstjert (Longtail) or, according to Widegren,
Langhala (Norw. Hale = stjert). But it is often con-
founded by the fishermen with White Bream and young
Bream, being included with these under the names
quoted by Artedi and Linnaeus, Blecka ( Blicka ), Bra-
xenpcmka , Braxenflia (Braxenflira). In order to pre-
clude the error into which he had himself been be-
trayed by the name of Faren , Ekstrom preferred to
use FUra.

(Ekstrom, Smitt.)

Genus PELECUS.

Beginning of the short dorsal fin situated vertically above that of the long anal fin , and the base of the former
less than 1J i of that of the latter. Dorsal musculature extending forward across the inter orbital space. Lateral
line bent sharply downwards behind the pectoral fins , and in the posterior part of its course undulating , or even
externally interrupted. Scales like those of the Bleak, but with the nucleus situated in the posterior half . Mouth

sharply ascending , and the tip of the lower jaw projecting.

Among the Cyprinoids to which the Scandinavian
fauna can lay claim, this genus is in several respects
the most remarkable. With a form of body reminding
us of the Herrings, it has several characters that range
it beside the Bleaks, others again being Abramidine

characters at their highest point of development. The
characters of Alburnus may be seen in the form of the
mouth, with the point of the lower jaw fitting into a
sinus at the tip of the snout, the slender and prolong-
ated pharyngeals, and the thin, comparatively deciduous

ZIEGE.

823

scales. The Abramidine type, on the other hand, may
be traced not only in the long anal tin and the naked-
ness of the median dorsal line throughout a consider-
able part of its length in front, but also in the similar
appearance of the median ventral line, which is sharply
carinated and naked all the way from the isthmus to
the vent, the latter being situated, as usual, just in
front of the anal tin. The most natural course is there-
fore to regard the genus as the type of an offshoot
from the common stock of the Bleaks and Breams.
But in the structure of the mouth, the pharyngeals, and
the scales the Zope shows signs of transition from the
true Breams to this genus. Pelecus is further distin-

guished by its long and pointed, almost scythe-shaped
pectoral fins, which recur, however, in some other ge-
nera, notably Chela, a genus belonging to Indian and
the East Indian Archipelago, fairly rich in forms, and
very nearly approximated to Pelecus. This latter
genus was established by Agassiz" in 1835, and con-
tains only one species, essentially differing from Chela
merely in the irregular, undulating lateral line.

Fries and Ekstrom adopted the genus — though as
a subgenus entitled Chela — in the previous edition of
“ Scandinavian Fishes" (1837), under the Swedish name
of Skdr-Braxnar ; but for the species they retained the
Linnaean name of Skarknifven (Carving-knife).

THE ZIEGE (sw. skaricnifyen).
PELECUS CULTRATUS.

Fig. 207.

Pharyngeal teeth slender , compressed , hooked at the tip , with pectinated masticatory surface , and set

in two rows: 2 , 5 — 5, 2.

Fig. 207.

a

Pelecus cidtratus, from Finland, Nyi.ander, 1850. '/2 °f the natural size, a, the left lower pharyngeal, seen from above, natural

size; b, the same seen from without, natural size; c , the hindmost tooth in the inner row, twice the natural size.

S'yn. Sarachi congener , Marsigl., Danub. Pann. Mys ., tom.
IV, p. 21, tab. VIII. Lucius, sp. 2 et 3, Klein, Hist.
Pise. Nat. Miss., V, p. 74, tab. XX, fig. 3. Cyprinus pinnae
ani radiis XXX, linea laterali declioata, ventre acut'ssimo,
Lin., It. Scan., p. 82, tab. II, fig. 1.

a Mem. Soc. Sc. Nat. Neucli., tom. I, p. 39.
b 6 — 7, according to Gunther.
c According to Gunther.
d ,, ,, Krcyer.

E. hr. 3; I). — ;

7 b

25—21

1 2

P. -; V. — ;

14—16 7

14—15

U. x + 1 + 17 + 1 + x; L. lot. c:a 100; L. tv. — 1;

4 — 6

Vert. 48 c — 51d.

Scandinavian Fishes.

101

824

SCANDINAVIAN FISHES.

Cyprinus cultratus, Lin., Syst. Nat., ed. X, tom. I, p. 326;
Bl., Fiscli. Deutschl., pt I, p. 255, tab. XXXVII; Retz.,
Fn. Suec., Lin., p. 360; Pall., Zoogr. Ross. Asiat., tom.
Ill, p. 331; Nilss., Prodr. Ichthyol. Scand., p. 32; Abass.

( Pelecus ), 1. c.; Cuv., Val. ( Leuciscus ), Hist. Nat. Poiss.,
vol. XVII, p. 330; Ivr. (Pelecus), Damn. Fisk., vol. Ill, p.
511; Nilss. ( Abramis , subg. Chela ex. Buchan.), Skand.
Fn., Fisk., p. 340; Kessl. (Pelecus), Bull. Soc. Natur. Mosc.,
vol. XXIX, I (1856), p. 376; Hckl, Kn., Siisswasserf.
Osir. Mon., p. 126; Dyb., Cypr. Livl., p. 153; Sieb.,
Siisswasserf. Mitteleur., p. 152; Mgrn, Finl., Fiskfn. (disp.
Helsingf.), p. 43; Gthr, Cat. Brit. Mits., Fish., vol. VII,
p. 330; Bncke, Fiscli., Fischer., Fischz. 0., W. Preass.,
p. 125; Mela, Vert. Fenn., p. 337, tab. X; M6b., Hcke,
Fiscli. Osts., p. 121; Grimm, Fish., Hunt. Russ. Wat., pp.
16 et 22; Mela, Sundm., Finl. Fisk., tab. XXVII; Lillj.,
Sv., Norg. Fn., Fisk., vol. Ill, p. 324.

The ‘Ziege’ attains even in the North a length of
4 dm.; but in Southern Russia, where it apparently
has its true home, it sometimes measures, according to
Kessler, as much as 6 dm. The elongated and com-
pressed form of the body is most suggestive of a lean
Herring, but the most characteristic external features
are the almost straight dorsal margin and regularly
arcuate belly, without the more or less sharp angles
which we have seen in the protiles of most of the pre-
ceding Cyprinoids. Furthermore the form differs from
that of our other Cyprinoids in the strong elongation
of the forepart of the trunk, for in no other Scandi-
navian member of the family is the preabdominal length
more than 3/10 of the length of the body. The greatest
depth in adult specimens is somewhat less than 7 5 of
the length of the body (about 19 % in our specimens,
which are 22 — 36 cm. long); the greatest thickness
about '/a — 4/<)> and the least depth of the tail about 1/3 ,
of the greatest depth of the body.

The head also measures somewhat less than 7s of
the length of the body, and is characterized principally
by the above-mentioned relation to the dorsal muscles,
the origin of which extends forward over the forehead.
The eyes are large and set rather low, almost as in
the Zope. Their longitudinal diameter, which even in
adult specimens is somewhat, though only slightly,
greater than the length of the snout, measures a little
more than 74 (28 — 26 %) of the length of the head.
The postorbital length of the head is about half its
entire length. The position of the nostrils is normal.
The sides of the snout (the preorbital bones) are re-
markable for the distinct ducts belonging to the sub-
orbital branch of the cephalic system of the lateral
line. The shape and position of the mouth we have
already described. The length of the upper jaw from

the tip of the snout is somewhat less than the least
depth of the tail, but the length of the lower jaw is
perceptibly greater than the latter, measuring about
42 % of the length of the head. Both the operculum
and the preoperculum are comparatively large, and the
latter is distinguished by the rectangular form of its
hind inferior corner. The gill-openings are large, the
branchiostegal membranes coalescing with the isthmus
rather near each other and in a line with the posterior
orbital margin. The gill-rakers are strong and rather
scattered. The outer row on the first branchial arch
contains about 17 or 18, the inferior pointed, the su-
perior flat and branched or lobulate at the tip.

The dorsal fin is remarkable both for its small
size and its backward position. It begins almost ver-
tically above the origin of the anal fin, at a distance
from the tip of the snout measuring about 64 % of the
length of the body. Its base is equal to the longi-
tudinal diameter of the eye (in our specimens 5‘3 — 5 %
of the length of the body) and measures about half of
its height in front, which is nearly 1 1 — 972 % °f the
length of the body. It is sharply and obliquely trun-
cate, with the upper posterior margin someAvhat con-
cave. The anal fin is more deeply concave, reminding
us of the Garpike, with the anterior part prolongated
to a lobe, but with the posterior part of almost uni-
form height. Its base measures about 2 1 1/2 — 22 %, and
its height in front about 12 — 11 %, of the length of
the body. The caudal fin is deeply forked, with pointed
lobes, the lower somewhat longer than the upper. Its
middle rays occupy about 7 % of the length of the
body, and measure a little more than 1/3 of that of
the longest caudal rays.

The shape of the pectoral fins too — expanded at
the base and incised at the lower margin — calls to
mind the Garpike or, still more, the Horse Mackerel;
and they are remarkable, as in the latter of these two
species, for their great length, 29 — 28 % of that of
the body and only slightly less than the preabdominal
length. The ventral fins, which are inserted half-way
along the body, are also more pointed than in the rest
of the Scandinavian Cyprinoids. Their length, which
measures about 12 — 1272 % of that of the body, is less
than the postabdominal length, though the latter is
remarkably small, being only about 1 6 1/2 — 15 % of the
length of the body.

The scales are thin and deciduous; they vary
greatly in size and shape on different parts of the

ZIEGE.

825

body, being small on the back and belly, largest at the
middle of the sides. The large scales are of a rounded
quadrilateral shape or almost circular; the small ellip-
tical, with the longitudinal axis set across the body.
The concentric striae are extremely fine and dense; the
radiating grooves few and faint on the hind part of
the scale, hardly visible on the inserted part, though
the latter, at least in the large scales, is usually si-
nuate at the margin.

The coloration resembles that of Alburnus and
Clupea. According to Hf.ckel and Knee “the occiput
is steel-blue or bluish green, the back grayish brown,
the cheeks of a nacreous lustre, the sides of a light
silvery lustre; the dorsal and caudal tins grayish, the
others with a dash of red; the iris silvery.”

The Ziege, like the Zarthe and, in some localities,
the Zope, is an anadromous fish, ascending rivers from
the sea or the great lakes in order to spawn. Its best
known haunts, where it is taken in millions, are the
Black Sea, the Sea of Azov, the Caspian, and the Sea
of Aral, with all the large rivers flowing into these
waters. But the species also occurs in the Baltic,
chiefly to the south and east, west to Greifswald, and
in Lake Ladoga. From these localities it makes its way
into the North German Haffs and estuaries and into
most Russian and some Finnish rivers. Within the
basin of the Danube it is common in Hungary, and in
summer some Hungarian lakes yield large catches of
Ziege; but in Bavarian waters it is rare. At the middle
of the last century, according to Linnaeus, it frequented
the River Helge off Kristianstad, in spring, till the end
of May. The specimen taken in this river and de-
scribed by LinnvEUS (length 22 cm.) is still preserved
in the Zoological Museum of Upsala University. The
species has never since been caught in the Helge, but

off the mouth of tins river it is well known to some of

o

the fishermen of Alius, according to their own asser-
tion, being frequently taken in the Herring-nets. In
Denmark it has never been found.

The Ziege is an active and roving fish, not unlike
the Herring. It lives on miscellaneous small animals,
such as crustaceans, insects, the fry of other piscine
species, and small fishes. It seeks its food, as indicated
by the position of the mouth, in the same manner as
the Bleak, mostly at the surface of the water. In spring
and early summer, from May to July, it spawns
in shallow water on a weedy bottom. In a female
weighing 117 grin, and with ovaries weighing 77
grm., Bloch estimated the number of the eggs at
105,740.

Bony, lean, and thin — hence, it is said, the Ger-
man name of Ziege (goat) — the species is held in
little esteem where other fish is to be had; but where
this is not the case, or where the Ziege is taken in
enormous quantities, as in Southern Russia, it is used
as human food. Its scales are employed, like those of
the Bleak, in the manufacture of the so-called essence
d’orient, the colouring matter of imitation pearls. The
nearest fishing-stations to Scandinavia where the Ziege
is taken in any considerable quantity, lie on the
Prussian coast and in the German Haffs, the tackle
used consisting of gill-nets, which are set at the sur-
face, or drift-nets. “It has numerous foes,” says
Bloch, “in predatory fishes and waterfowl, to which
it often falls a prey, its silvery colour rendering it
easy of observation.” Thus the Ziege shares the fate
of the Herring, and where these two species live in
company, the former has as good means of escape
as the latter, if not better, its sharp tins being well
adapted for rapid flight and speedy doubling.

826

SCANDINAVIAN FISHES.

THRISSOMORPHI.

Pliysostoms with the shoulder-girdle suspended from the head (as usual in the Teleosts). Scapular disk inter-
nally strengthened by an arch formed by a special bone ( os prcecoracoideum) from the coracoid bone to the clavicle.
The first four abdominal vertebrae of normal form and development. Hyomandibular and pterygopalatine arches
complete, as well as ( in most cases) the opercular apparatus. Maxillary bones fully developed.

That the Salmons and Herrings have much in
common, was declared by Agassiz in 1843", when he
united them into one family, Halecoides; and Cope
followed up this opinion in 1871* by the establishment
of the order Isospondyli, answering to the series of the
Thrissomorphs, and distinguished from the preceding
series mainly by a negative character, the absence of
the so-called acoustic bones. The connexion between
the organs of hearing and the air-bladder, however, is
not foreign to this series. In the Clupeoids Weber0
discovered that the air-bladder, which tapers forward
for some length, is divided in front into two branches,
one to the right and one to the left, which pass through
the occipital bone into the cranium, and there, as Hasse^
has shown, touch with their tips the outer wall of the
lymphatic chamber surrounding the auditory apparatus.
Sagemehl has given good reason0 for his assumption
that this mode of communication is the more primitive,
from which the more complicated connexion of the
preceding series has been evolved, and which has been
persistent not only in the Clupeoids and some other
Pliysostoms, but also in some Physoclysts of ancient
type, as for instance the Berycoids (cf. above, p. 66),
the Macruroids, some Gadoids and Balistoids. We have
also seen a reminiscence of this connexion in the two
tubular processes which ascend from the anterior ex-
tremity of the air-bladder of the Gadoids (cf. above,
p. 476) in vermicular curves towards the head, one
on each side of the anterior renal mass. The absence
of the so-called acoustic bones is thus explained as an

earlier stage of development; and a negative character
of this signification can hardly be sufficient to define
a natural evolutionary series. We also find that a great
number of the Thrissomorphs are without the character
adduced by Agassiz as one of the most important in
his definition of Halecoides, namely the participation
of the maxillary bones in the formation of the margin
of the upper jaw. Thus we might reasonably treat
these forms as a distinct series of families ( Scopelo -
morphi); but the resemblances in other respects are
sufficiently great to render the Thrissomorphs a natural
whole, with the same limitations as Cope’s Isospondyli,
although the said resemblances do not constitute cha-
racters applicable to the Avhole series. Within this
series too the variation of form and structure is great
and affects most of the organs. Scales and a distinct
lateral line may be present or wanting. The maxil-
laries may be simple, as in the rest of the Teleosts,
or composite, furnished with so-called supplementary
bones answering to the cheek-bones ( ossa jugalia ) of
the higher vertebrates. The teeth may be numerous
and well developed or wanting. A so-called adipose
fin is present on the hind part of the back in many
forms, but wanting in others. Luminous organs, so-
called phosphorescent spots, may be present or absent.
The ovaries may be furnished with an oviduct or with-
out a complete tube of this description. The air-bladder
may be present or wanting. Such great differences in
forms which nevertheless show their natural affinity
in some way or other7, render the characterization of

° Recti. Poiss. Foss., tom. V, pp. 3 and 96.

6 Iclitliyol. Less. Ant., Trans. Amer. Philos. Soc., Philad. N. ser., vol. XIV, pp. 452 and 454.
c De anre cett., p. 73, cett. ; figg. 6.3, cett.
d A.nat. Stucl., Bd. I, p. 599, cett.
e Morpliol. Jahrb., Bd. X, p. 51.

f Thus for example the Haplochitoiridce — - a family of fresh-water fishes from South America and Australia — are almost typical
Salmons, but have the margin of the upper jaw formed by the intermaxillaries alone; and similarly the Sternoptycliidce and Scopelidce are
families so like each other that they ought hardly to be kept apart, though in the former the osseous framework of the mouth follows the
Halecoid type, while the latter preserve in this respect the ordinary Teleosteous character.

THRISSO MORPHS.

827

the series extremely difficult. Within the Scandinavian
fauna the series contains the following families.

I: Hind part of the back furnished with an adipose fin.

A: Ovaries without special oviduct com-
municating directly with their cavity :
the ripe eggs fall into the abdominal

cavity and pass thence through a geni-
tal pore. No phosphorescent spots on

the sides of the body Farm Salnionida •

B: Ovaries furnished with a complete ovi-
duct. Sides of the body furnished in
all Scandinavian forms with phosphore-
scent spots'* Fam. Scopelidce.

II: No adipose fin Fam. CAupeidae.

Fam. S A L 31 0 X I I) E.

Body of the typical Salmon form or more compressed , approximated to the Boacli or the Herring form. Scales
middle-sized or small, rather firmly attached1'. No phosphorescent spots on the sides of the body. Dorsal margin
of the tail furnished with an adipose fin. Margin of the upper jaw formed in front by the intermaxillaries
and behind by the maxillaries. No barbels. Air-bladder simple and not connected with the cranial cavity.
Branchial cavity furnished with pseudobranchice. Ovaries open on the outside, without oviducts or with oviducts

opening inwards ( into the abdominal cavity).

The family of the Salmons and Gwyniads is well-
known everywhere in the frigid and temperate regions
of the northern hemisphere. In the southern hemi-
sphere, on the other hand, only two genera occur, a
Capelin form ( Betropinna Richardsonii ) belonging to the
fresh water of New Zealand, and two (?) Argentine
species, marine iishes of the same locality. The family
possesses more than ordinary interest both from an eco-
nomical and a scientific point of view, to the economist
because the flesh of these fishes is one of the most
highly esteemed and most remunerative foods, to the
scientist because there is hardly any other part of the
system where he is confronted with such difficulties in
defining the limits of the species. This is due in most
cases to a plasticity and inconstancy of form, caused
beyond doubt by the periodic migrations of the Salmo-
noicls from the sea or the great lakes to running fresh
water. The economical value of these fishes has also
contributed to the said difficulties. Where they appeared
at certain seasons and certain places in enormous mass-
es, with one or other of their various dresses predomi-
nant, the fishermen gave these dresses different names
which the systematist had to explain, often merely after
examining solitary specimens that were supposed to
represent distinct species. “Kart barn har manga namn"%

says the Swedish proverb; and this rule has repeatedly
asserted itself in the nomenclature of the Salmonoids.
The investigations of recent years, however, have elu-
cidated the significance of the variation of form within
this family and the relationship to each other of the
more or less constant forms, arid have thereby thrown
light upon the comprehensive question of the origin of
the species.

The geological researches of Agassiz d have traced
the Salmonoid family back to the Cretaceous Period. At
that time there lived, where the green sand now lies in
modern Westphalia, and where the chalk subsided over
Southern England, several forms, which Agassiz referred
to the genera Osmerus and Osmeroidesj the latter fur-
nished ■with scales resembling those of the modern ge-
nus Argentina. The evolution of the Salmonoid family,
we might hence conclude, began with marine fishes of
the Smelt and Capelin type, with few pyloric appen-
dages or none at all, and its development advanced in
two directions, 1) towards the Salmon and Charr, fur-
nished with teeth and with comparatively small scales,
2) towards the toothless and comparatively large-scaled
genus Coregonus. In both these directions of evolution
the pyloric appendages were more and more developed,
to assist these fishes in storing up a reserve supply of

a An extremely rare form, Sadis atlanticus , is an exception.

b The genus Salanx ( Albula chinensis , Osbeck, Ostind. Resa, p. 237), however, is scaleless or has small, extremely thin, and deci-
duous scales (Gunther).

0 “A dear child has many names.”
d Rech. Poiss. Foss., vo). V, p. 103.

e Gunther, however, considers that this genus perhaps belonged to the Scopeloid family. Introd. Study Fish., p. 582; Handb.
Iclithyol., p. 417.

828

SCANDINAVIAN FISHES.

fat for the time when they must resort to fresh water
for the reproduction of their species.

The Scandinavian genera of this family are the
following:

A: Dentition strong, even the tongue fur-
nished with teeth. Length of the maxil-
laries more than 53 %a of that of the head
reduced (from the Lind extremity of the
intermaxillary bone to the posterior mar-
gin of the preoperculum). Base of the
dorsal fin proper shorter than the head
reduced. ( Salmonince )

1 : Branched rays in the anal fin lessthanlT.
a: Transverse rows of scales on the

sides of the body more than 100.. Genus Salmo.
b: Transverse rows of scales on the

sides of the body less than 80 Genus Osmerus.

2 : Branched rays in the anal fin at least 17 . Genus Mallotus.
B: Dentition weak or teeth wanting, at least
the tongue toothless6. Length of the ma-
xillaries less than 53 % of that of the
head reduced. ( Coregonince )

1: Base of the dorsal fin proper more
than 15 % of the length of the body

and longer than the head reduced Genus Thymallus.

2: Base of the dorsal fin proper less than
15 % of the length of the body.
a: Base of the dorsal fin proper more
than half the length of the head

reduced Genus Coregonus.

b: Base of the dorsal fin less than half

(<45%) the length ofthe head reduced Genus Argentina.

In this manner we have endeavoured to find an
expression for the said two directions of development.
The development, however, has not been so regular that
a sharp and fixed line can be drawn between two di-
stinct subfamilies; and the intermediate position occupied
by the Graylings (genus Thymallus) may perhaps justify
the establishment of a third subfamily, characterized
principally by the advanced development of the dorsal
fin. The community of origin in the case of all these
genera affords an explanation of the fact that the fry
of the most differentiated forms in one group show
points of resemblance to the other group.

In order rightly to comprehend the relations be-
tween the genera, it is indeed necessary to have a
knowledge of the exotic forms as well. But two of the
Scandinavian genera are so rich in forms, even within
the limits of our fauna, that in them we may trace the
development of form in its past course and, seemingly,
in its present continuation.

ovd

Fig. 208. A gravid female Smelt, taken at Stockholm on the 19th
of April, 1892. Natural size, ovs, left ovary, with eggs arranged in
transverse rows; ovd, lower (posterior) part of the right ovary; m,
margin of the mesoarium, which envelops the ovary both above (in
front) and below (behind) even on the left (outer) side, but below,
owing to the coalescence of the margin with the wall of the abdomi-
nal cavity, forms a funnel-shaped duct, open above at a; r , rectum;
an, vent; gu , situation of the genital and urethral apertures, the for-
mer in front of the latter, drawn as a common opening because the
wall between them could not be distinguished externally.

a Exceptions occur among Salmon fry.

„ Grayling „

b

SALMONOIDS.

829

In the structure of the ovaries the Salmonoicl fam-
ily stands almost alone among the Teleosts. Only
among the Eels, in a New Zealand family ( Galaxiidce )
and an East Indian and West African family ( Notopter -
idee), are the eggs deposited in the same manner. Yet
the SaJinonoids cannot be said to be entirely destitute
of oviducts. The true difference between the ovaries
of this family and those of most other Teleosts is that,
while in the latter the said organ is a closed sac, open
only behind (through the oviduct), the outer wall of
which is formed by a fold of the peritoneum (meso-
arium), and round the inner surface of which the ovi-
ferous layer is set in transverse or longitudinal folds
(lamellae), in the Salmonoid family this layer extends
only over the outer side (the side facing the lateral wall
of the abdominal cavity) of the simple, pendent meso-
arium (peritoneal fold). Or in other words, we may
imagine the otherwise closed ovary (together with the
oviduct) to be split throughout the length of the outer
side (the side facing the lateral Avail of the abdominal
cavity). Both at the anterior extremity of the ovary

and at the loAver (ventral) margin, however, the meso-
arium bends toAvards the outer side, and behind the ovary
the mesoarium is continued in the form of a simple
lamella, either hanging betAveen the intestine and the
Avail of the abdominal cavity, Avith the loAver (ventral)
margin free, or Avith the said margin curved upAvards
and outwards and coalescent Avith the Avail of the ab-
dominal cavity, thus forming an oviduct (abdominal
funnel) open even in front. These oviducts (fig. 208)
— one on each side in front, but united at the extreme
end into a common passage behind the intestine — Avhich
occur in the Smelts and Capelins, are indeed imperfect
in comparison Avith those of the other Teleosts. But
in their anterior opening they sIioav a remarkable ana-
tomical resemblance" to the corresponding structure both
in the Ganoids (Avith the exception of Lepidosteus ),
Sturgeons, and Cartilaginous fishes and in the higher
vertebrates, where the oviducts (Mullerian ducts) are
separate from the ovaries and open into the abdominal
cavity, receiving in this orifice the ripe eggs Avhen de-
tached from the ovaries.

Genus SALMO.

Mouth armed with strong and pointed , rather scattered teeth , set in one row on the lower jaw, the inter maxillaries,
the maxillaries , and the palatine hones, in one or two rows ( hut deciduous and sometimes wanting) on the shaft of
the vomer, in an irregular group on the head of the vomer, in two rows on the tongue. Length of the maxillaries
behind the intermaxillaries as a rule 53 — 63 %b, and of the lower jaw as a rule 75 — 95 %c, of that of the head
from the hind extremity of the intermaxillary hone to the posterior margin of the preoperculum. Number of rays
in the anal fin as a rule less than in the dorsal d and at most 14, of which 7 — 10 are branched, and the base
of the anal fin shorter than that of the dorsal. Pyloric appendages well-developed and numerous ( about 25— -90).

Scales small, more than 100 in the lateral line, which is complete.

In the Tertiary Period, when the Rocky Mountains
rose and divided the arctic and temperate Avaters of
North America into tAVO regions, an eastern and a Avest-
ern, the genus Salmo Avas differentiated in two distinct
directions of development, of which in the above dia-
gnosis Ave have taken into account only one, the Sal-
mons and Charr of the eastern region. The other course
of development ran side by side Avith this — branched

like it into Salmon and Charr — but acquired a cha-
racter peculiar to itself in the long anal fin, with a
greater number of rays, (15?) 17 — 19, of Avhich 14 —
16 may be branched. This direction of development,
whether Ave choose to regard its forms as a genus or
subgenus, has received in recent times6 the name of
Oncorhynchus. To the European faunist it is not Avith-
out interest, for its best representative, the Quinnat

° Cf. Rathke, Beitr. z. Gesch. d. Thierw., Abtli. 2. Uber den Darmkanal und die Zeugungsorgane der Fische , pp. 123 and 159;
Huxr.EY, Proc. Zool. Soc. London 1883, p. 132; Weber, Morphol. Jahrb., XII, p. 3G6. Oviducts of the same description occur in the Cape-
lin; but there only one (the left) ovary is fully developed. In Salmo and Coregonns the mesoarium passes over the rectum, forming on
each side thereof a short abdominal funnel (peritoneal duct).

h In Parr and individual exceptions 43 — -53 % or, on the other hand, up to 78 %.

C (IK 7 PC -111 0/

?? 9? •)•> ) ?? ?? n ?> x x

d Sometimes the number is the same.

e Suckley, Ann. Lyc. Nat. Hist., New York 1861, p. 312.

830

SCANDINAVIAN FISHES.

Salmon ( Oncorh . tschawytscha ), has been introduced
with success by piscicultural methods not only into the
Eastern States of North America, but also into Australia
and Europe, though not yet, to the best of our knowledge,
into Scandinavia. The Scandinavian fauna has, how-
ever, been enriched in very recent years, according to
a newspaper paragraph at the beginning of 1892, by
the introduction from Germany of a form planted in
the latter country, the Californian Rainbow Salmon
( Salmo irideus). The said form, together with a Charr
form — or rather an intermediate form between the
Salmon and Charr — has its original home west of the
Rocky Mountains and in the Pacific Ocean, although
they both belong to the course of development other-
wise represented only to the east of this mountain
range. A closer examination of these forms leads us,
however, to the result" that in them we have a blending
of characters most naturally to be explained, if not on
the assumption of hybridism, as a reminiscence of the
time when the Salmon and Charr, in their common de-
velopment, were not fully differentiated from each other.
Another American form, a Charr from the Eastern
States (Salma fontinalis), has been cultivated with suc-
cess in England, Wales, and Scotland. Salmo fontinalis
is also an intermediate form between the Salmon and the
Charr, the only character — though not of great vali-
dity — in which it differs from the Greenland Charr,
the absence of teeth on the basibranchial bones (hyoid
teeth), being a character of the Salmon6. The Brook-
Trout (S. fontinalis) has been crossed both here and
in America with the Salmon as well as with European
Charr; and these experiments have taught us, among
other things, that the character which otherwise com-
poses a constant distinction between Salmon and Charr,
the feeble ossification and toothlessness of the vomer in
the latter, loses its validity in these hybrids'. Conse-
quently, in elucidating the relations of the forms to each
other, here as in the preceding family, we have to
reckon with hybridism as an important factor in the
modifications of the types. To all appearances the ge-
nus has its original home in the American seas, where
Salmon and Charr still occur in masses almost incon-
ceivable to the European.

" Smitt, Riksmuseets Salmonidcr , Vet.-Akad. Hand]., Bd. 21,
h Cf., however, Smitt, 1. c., lab. metr. V and VI, Nos. 371,
e Cf. Day, British and Irish Salmon idee, pp. 261 and 270,
cl Day, British and Irish Salmonidce, p. 143.
e Day, ibid., p. 51.

From the preceding family we have learnt that
cross-breeding may take place not only between differ-
ent species, but also between different genera. Hybri-
dism alone cannot, therefore, prevent the distinguishing
of the species. But the variability of form within the
genus Salmo has hitherto rendered it impossible to de-
fine with certainty the numerous species that have been
adopted; and we doubt whether the observer who con-
sistently requires trustworthy and distinct characters can
recognise more than three European species of the ge-
nus: the Salmon, the Huch, and the Charr. As yet,
however, to the best of our knowledge, only Nilsson
and Sundevall — the latter in the titles of the figures
in PI. 58 and 59 in the former edition of “Scandinavian
Fishes” — have ventured to advance this simple opinion.
Still, both from a scientific and an economical point
of view, it is of importance to know the conditions that
involve the said inconstancy; and to this end it has
been necessary to denote by special names the more
or less constant forms that appear under different cir-
cumstances and in different localities. As regards the
Salmon, general credence has been given in recent times
to the opinion first advanced by Lilljeborg and after-
wards by Widegren, namely that we can distinguish
between two “species”, the Trout d ( Salmo trutta, Sw.
grdlaxen ) and the Salmon f! (S. salar , Sw. blanldaxen).
The same applies in essential points to the Scandinavian
Charr, among which we can generally distinguish with
ease two forms, the Northern Charr ( Salmo alpinus , Sw.
L app lands-rodi ng en) and the Saddling ( S . salvelinus,
Sw. Vetterns-rodingen). Of the Huch, which, at least
up to the present, cannot be claimed for the Scandina-
vian fauna, there are also two forms, the Siberian Huch
( Salmo fluviatilis ) and the Danube Huch (S. hueJio).

The Scandinavian forms of the genus Salmo may
in general be distinguished most readily in the follow-
ing manner:

A: Number of scales in a longitudinal row
above the anal fin, for an extent of V10
of the length of the body, at least 23
(23 — 30). Salmo umbla.
a: Distance between the ventral fins and
the tip of the snout more than half
the length of the body Salmo salvelinus.

). 8, p. 143.

386 and 458.
ig. 52, 2 and 2a.

SALMONS.

831

b: Distance between the ventral lins and
the tip of the snout less than half the
length of the body. Salmo alpinus.

B: Number of scales in a longitudinal row
above the anal fin, for an extent of 1/10
of the length of the body, as a rule at
most 19 (15 — 19, exceptionally 13 or 21).

Salmo salar.

a: Least depth of the tail less than 27 %
of the preabdominal length (the di-
stance between the first rays of the

pectoral and ventral fins) Salmo salar.

b: Least depth of the tail more than 27

% of the preabdominal length Salmo trutta.

The natural relation between these forms asserts
itself both in their changes of growth and their external
differences of sex. These differences are most distinctly
reflected in the characters here given for the said forms.

Artedi, the father of modern ichthyology, on whose
opinions and method Linnaeus, his contemporary and

o

friend, in great part based Ins system, was born in An-
germanland. As a native of Northern Sweden, he was
undoubtedly well acquainted with the opinions that pre-
vailed at his time in this part of the country, where
the Salmon forms are most abundant, as to the diffe-
rences between them and as to their nomenclature.
Where he applies to the Swedish Salmon forms their
current Swedish names, we have in the latter, beyond
doubt, the best possible clue to a correct under-
standing of his scientific views on this head.

In Aktedi’s Genera Piscium we find the following
Salmons enumerated.

1 : Salmo rostro ultra inferiorem maxillam ssepe prominente, in Swe-
dish, Lax and Blanklax.

2 : Salmo maeulis cinereis, caudfe extreme asquali, in Swedish,
Gralax.

3: Salmo latus; maeulis rubris nigrisque, cauda sequali, in Swedish,
Laxoring, Borting etc.

4: Salmo cauda bifurca; maeulis solum nigris, sulco longitudinali in
ventre = Salmo lacustris , auett., from the Lakes of Geneva and
Garda.

5 : Salmo maxilla inferiore paullo longiore, maeulis rubris, in Swe-
dish, Forell, Stenbit , Rdfislc, Backro etc.

6 : Salmo oblongus duabus dentium lineis in palato, maeulis tan-
tummodo nigris = the Huch of the Danube.

7: Salmo pede minor, quinque dentium ordinibus in palato = a con-
fusion, originating with Willughby, of Salmo carpio, auett., from
Lake Garda, with the Gilt Gharr , an English form.

8: Salmo vix pedalis; pinnis ventris rubris, maxilla inferiore paullo
longiore = Lapplands-Rddingen.

9 : Salmo lineis lateralibus sursum recurvis, cauda bifurca = Salmo
ivmbla , auett., from the Lake of Geneva.

10: Salmo pedalis; maxilla superiore longiore = Salmo salvelinus,
auett., from the Lake of Geneva.

In Artedi’s Synonymia Nominum Piscium the Sal-
mons are given in the following order:

1 = 1 in Genera Piscium
1 1

,, , = Trutta , auett.

11
11

6: Salmo dorso fulvo: maeulis luteis, cauda bifurcata = Salmo sal-
marinus, auett., from Trent (the Tyrol)

7 = 9 in Genera Piscium

8 = 6 „ „

9 = 4,, „ „

10 = 8 ,, ,, ,, , here with the Swedish names of Rotele

and Boding.

n = io„

12, Salmo minor, vulgari (1) similis.

In the Descriptions Specierum Piscium, quos vivos
prcRsertim dissecuit et examinavit etc., from which title, as
well as from Linnaeus’s preface, it appears that Artedi
there included all the species which he had personally been
able to examine, we find only five species of the genus:
1 = 1 in Genera Piscium.

2=12 in Synonymia Nominum Piscium , is here explained as a young
Salmon, though often more than 12 inches (i. e. about 30 cm.)
long, bearing the name of Laxunge at Elfkarleby, and differing
from the Blanlclax only in the comparatively larger and blunter
head, the denser black spots, and the slight bifurcation of the
caudal fin.

3 = 3 in Genera, Piscium.

4 5 ,, ,, ,,

5 = 8 „ ,, „

When LiNNiEUS framed on this foundation his
scheme of the Swedish Salmons, he adopted in his
Fauna Suecica (1746) the following:

Sp. 306 = No. 1 in Art., Gen. Piscium.

,, 307 = ,, 2 ,, ,, ,, ,, , with the remark: “It oc-

curs in company with the preceding form, and I should
scarcely consider it to be specifically distinct therefrom'1.
,, 308 = No. 3 in Art., Gen. Pise, according to the synonymy, but

with a new diagnosis: Salmo maeulis nigris brunneo cin-

ctis: pinna pectorali punctis sex, and with the Swedish

name of Laxoring altered to Orlax.

,, 309 = No. 5 in Art., Gen. Pise., but with the word paullo

omitted from the diagnosis, and with the addition of the
Swedish name of Laxoring.

„ 310 = No. 8 in Art., Gen. Pise.

Now Gisler tells us in a series of papers (in the
Transactions of the Swedish Academy of Science for
1751 and 1752) which are well worth reading even at
the present day, that the Blanklax was called Grdlax in
Norrland, “when it had spent itself in the rivers and had
become quite lean and gray, with a long hook at the
tip of the lower jaw, especially in the males, after the
flesh had wasted away”. Gisler, it is true, recognised
only three Norrland species, namely

1 Spec. Lax,

2 ,, Laxoring,

3 ,, Stenbit etc.

Scandinavian Fishes.

105

832

SCANDINAVIAN FISHES.

But as varieties of Laxoring he adopted 1) from Lin-
naeus’s Fauna Spec. 307, which he calls Laxoring, Lax-
ockel, Fjerd-lax, or Eriksmdss-lax, and Spec. 308, which
he calls Borting, Sikmatk, Tajmen, or Lodjor, and 2)
from Artedi the above-mentioned No. 3 in Gen. Pise.,
which he calls Tvdrspol, and No. 12 in Syn. Nom. Fisc.
which he calls Laxunge , Smdlax, or Padrifvare. Gis-
ler’s third species, Stenbit etc., is indeed equivalent,
according to his synonymy, to No. 5 in Artedi, Gen.
Pise.; but all that he says of it is that it “has its con-
stant haunts in small streams, tarns, and lakes, and is
quite rare further down the country in the great cas-
cades. Deposits the roe in small streams at the middle
of September. In clear water with a stony or sandy
bottom its colour is light, but on a bottom of ooze or
mud quite blackish”. It is not improbable that Gisler
meant the Charr, which was of less interest to him and
perhaps, in consequence, less known, for both the Swe-
dish name of Boding and the Lapp Baud (according
to Linnaeus) might easily be confounded with Bofisk.
If this be the case, we find in Gisler the opinion now
held by most ichthyologists, that within the Scandina-
vian fauna there are only three species of the genus
Salmo, namely the Lax (Salmon), Oring (Trout), and
Boding (Charr).

This opinion was subsequently advanced first by
Lilljeborg in Ofvers. Vet.-Akad. Forliandlingar for the
year 1849, where he points out that “the characters
which divide Salmo salmulus and Salmo fario have much
in common with those which distinguish Salmo salar
and Salmo eriox. That S. salmulus is the young of
the Blanklax ( S . salar), and S. fario that of the Okla
or S. eriox, is therefore almost unquestionable, especially
as they occur in the same waters, and connecting links
between them are known”. A more explicit statement
of this opinion was Widegren’s principal object in
his u Bidrag till kdnnedomen om Sveriges Salmonider”
( Ofvers . Vet.-Akad. Fork. 1863).

But another, still more radical reduction of the
species had been proposed, conditionally so to speak,
by Nilsson in Ofvers. Vet.-Akad. Fork, for 1848.
“Under such circumstances, and seeing that all the cha-
racters prove variable in so high a degree, Ave are
almost tempted to ask Avhether there are here more than
two species of Salmons: Trutta and Salvelinus-, or the
very two which I have designated in my Prodromus as
the representatives of distinct groups”. Nevertheless he
retains the Blanklax, Grdlax, Laxoring, and two Charr

forms as “the most differentiated” or “least intercom-
municating” species. In his Skandinavisk Fauna he
remarks, hoAvever (p. 395), that the Grdlax “is 'most
probably nothing more than an old outgroAvn form of
Salmo Trutta or Odd’ / and Ave may thus trace even
in Nilsson’s writings the opinion “that in the division
of the true Salmons Ave have only two species certainly
distinct from each other” (Lillj.). While Nilsson, how-
ever, in his last-mentioned Avork, divided the Trouts
( Oringarne ), including the Grdlax, into six (conditio-
nally seven) species, Sundea^all adopted the above opi-
nion, that the true Salmons of Scandinavia belong to
one single species. This appears from the titles given
by him to the figures in PI. 58 and 59 in the first
edition of “ Scandinavian Fishes'. The male Salmon
represented in PI. 58 is evidently a Blanklax, probably
from Norrkoping; fig. a in PI. 59 is a Laxoring ( Bor-
ting) from the Ljusne Elf, and fig. b in the same plate
a Grdlax from Lake Wetter; yet all three bear the
specific name of salar.

All that different vieAvs as to specific determination
has been able to accomplish, has thus been devoted to
the elucidation of the Scandinavian Salmon forms. But
here as elseAvhere the point seems to be, not so much
the establishment of a certain number of species, as the
explanation of the natural relationship between the
forms selected as distinct types. Before these forms
can rank as distinct directions of development, it is of
course necessary to sIioav that the characters employed
do not coincide with those Avhich mark the common
variations of the Avhole genus under the influence of
different developmental, sexual, and other circumstances.

We begin with the investigation of these circum-
stances in the true Salmons, assuming at first the di-
stinction between Salmon and Trout, as separate spe-
cies, to be true.

In order to find an expression for those common
variations of the Salmons Avhich depend on their growth
(their developmental circumstances), Ave have divided
our Scandinavian material, 316 specimens, into five
groups, arranged according to age (the length of the
body). These five groups represent different stages of
development Avhich are in general Avell defined both by
form and coloration, though the rule is often infringed
OAving to a cause long known, especially as a result of
Widegren’s observations, namely the fact that the de-
velopment is not uniform, one or other of the stages
being often persistent in an individual specimen longer

SALMONS.

833

than in its fellows of the same age. In average cal-
culations, however, such individual exceptions may be
regarded as eliminated, and the said classification is
observed, generally speaking, even in popular phraseo-
logy, the first group containing the Laxyngel , Salmon-
fry (length under 70 mm.), the second group the Stir-

rar, Parr j length ^ rara'j, the third group the Fo-
reller or Stenbit | length ^ inni'|, the fourth group

the Oring ar or Borting | length ^ mm‘ J, the fifth

group the full-grown Laxar or Salmon (length > 450
mm.). The average percentages calculated as expres-
sions of the different relations of form in these groups,
lead us to very singular results". Out of 59 distinct
relations of form' 41 follow a fixed course in their
changes of growth, i. e. they either decrease with in-

creasing age (as for example the average length of the
head in proportion to that of the body), or increase
with age (as for example the average postabdominal
length in proportion to the length of the body).

In order to obtain an expression for those common
variations of the Salmons which are due to external
sexual differences, we have compared 17 males of the
different stages of growth with 17 females of about the
same size as the males and with the sexual organs about
equally ripe. The result is that 16 of the said 41 changes
of growth run in the same direction as the sexual
difference between male and female, i. e. a change of
growth that shows decreasing percentages with increas-
ing age answers to a sexual difference with greater
percentages in the males than in the females, and vice
versa. The two examples cited above appear in the
following percentages :

Average

Pry.

Parr.

For ell.

Oring.

Lax.

c?

9

Length of the body expressed in millimetres

48

108

179

318

595

218

218

„ head in % of the length of the body .

25. o

23.o

22.5

21.9

20.9

23.4

21.8

Postabdominal length „ „ ,, ,, „ „ „ „

16.9

19.7

19.7

20.4

21.2

19.6

20.9

The parallelism between the numerical alterations
l) in the first five columns and 2) in the last two
columns, may ' evidently be expressed as follows: the
younger Salmon are to the older as the males to the
females.

If Ave noAV examine the difference of form betAveen
the tAvo Scandinavian Salmon species that have received

most general recognition, Salmo trutta and S. solar avc
arrive at exactly the same result: the difference between
the averages for the said characters runs from the Grd-
lax (S. trutta) to the BlanJdax (S. salar ) in the same
direction as the averages for the changes of groAvth and
also for the external sexual differences. Some of the most
important of these characters may here be tabulated.

A

y e r a

g e

Pry.

■>

Parr.

Forell.

Oring.

Lax.

c?

9

1

Salmo

trutta

47

108

182

313

5871

Length of the body expressed in

millimetres..

I

218

218

1

Salmo

salar

50

109

161

403

609|

„ „ „ head

.. in % of the

length of the body

Salmo

trutta

25.2

23.5

22.6

22.o

22.o \

23.4

21.8

55 55 5? 55

- „ „ „ „

77 77 77 77

Salmo

salar

24.6

23.5

22.2

20.i

20.2 f

Postabdominal length

Salmo

trutta

16.5

19.8

19.7

20.3

21.21

19.6

20.9

55 5)

-- 77 77 77 77

„ „ „ „

Salmo

salar

17.7

19.5

19.5

21.3

21.i |

Length of the maxillaries

Salmo

trutta

9.8

9.2

9.i

9.o

8.6)

9.G

8.6

55 55 55 55

-- 77 55 75 77

77 75 77 57

Salmo

salar

8.i

8.0

7.6

7.o

7.o f

Least depth of the tail

Salmo

trutta

9.5

9.i

9.4

8.7

8.7 1

8.7

8.4

)! n n » »

-- „ „ „ „

55 55 55 55

Salmo

salar

8.4

8.o

8.2

7.i

6. 9 J

Length of the middle caudal rays „ „ „ „

55 55 55 5)

Salmo

trutta

11.4

9.0

8.o

7.7

7.7)

ft 1

„ » „ „ „ „

77 77 77 77

55 55 55 55

Salmo

salar

8.8

7.i ■

0.7

5.5

5.8 |

Least depth of the tail

■ 77 77 77 77

preabdominal length

Salmo

trutta

36.3

32.4

32.7

30. o

30.i)

30. o

29.4

55 77 77 77 77

- 77 77 77 77

55 ?5

Salmo

salar

30.4

27.8

28.i

23.6

23.2 1

° As we cannot give here all the particulars of our investigations, we refer the reader to our Kritislc Forteckning ofver de i Rilcs-
museum befintliga Salmonider, Vet.-Akad. Handli, Bd. 21, No. 8.

834

SCANDINAVIAN FISHES.

Many other such relations might here be cited to
prove the statement that the relation between Salmo
trutta and Salmo solar is the same as that between young
Salmon and old and between the males and the females.
To judge by all analogous cases there must be some
causal connexion between the difference of species, the
difference of age, and the difference of sex.

We find, however, among the changes of age cer-
tain peculiarities which indicate that another factor has

also asserted its influence in the differentiation of the
species. We see something of the kind even among
the above averages, for example in the average least
depth of the tail: — a comparatively large gap appears
in the numerical series, usually after the Forell stage,
and this gap is greatest in Salmo solar. The most
striking example, however, is perhaps the relation be-
tween the height of the anal fin (the length of its
longest ray) and the length of the body:

Average

Fry.

Parr.

Forell.

Oring.

Lax.

Length of the body expressed in millimetres Salmo trutta

47

108

182

313

587

,, ,, ,, „ ,, ,, ,, Salmo salar

50

109

161

403

609

Height of the anal fin in % of the length of the body Salmo trutta

13.c

13.2

13.i

12.i

11.5

„ » „ „ „ „ V „ ), „ „ „ ?) - Salmo salar

12.9

11.8

11.2

8.7

9.0

Now it is a well-known rule that after the Forell
stage the Salmons change their habitual place of abode.
The Grdlax ( Salmo trutta) leaves the brooks for meres
and large lakes, sometimes directly for the sea; but the
Blanklax ( S . salar ) must necessarily repair at this pe-
riod to the sea, else its development is arrested, and it
persistently retains a greater or less similarity to the
Grdlax. Thus the wide gap in the averages is probably
connected in some way with an alteration in the abode
and habits of the Salmons. Here we have assumably a
third factor in the differentiation of the species.

A fourth cause of difference in form undoubt-
edly lies, here as among the preceding family, in hy-
bridism, of which we have just given instances.

The relation of the Charrs to the Salmons leads
us in most characters to the conclusion that the former
should be regarded as more advanced stages of deve-
lopment of the Salmon type. To exemplify this, we
shall only refer the reader to the above-mentioned
characters, which afford the following averages:

Avers

g e

316 Salmon.

108 Charr.

Length of the body expressed in millimetres

279

333

v head

. in

% of

the length of the body

22.5

21.5

Postabdominal length

- 55

55 55

55 55 55 55 55

19.9

20.9

Length of the maxillaries

- 55

5 5 55

55 55 55 55 55

8.8

8.5

Least depth of the tail

55 55

55 55 55 55 55

8.8

7.2

Length of the middle caudal rays

55 55

55 55 55 55 55

7.9

6.9

Height of the anal fin

55 55

55 55 55 55 55

12.3

10.5

Least depth of the tail

55 55

,, preabdominal length

30.7

24.9

In every case where the percentage decreases dur-
ing growth in the Salmons, it is less in the Charrs,
and vice versa. This applies also to an internal cha-
racter which usually possesses great validity, the num-
ber of the gill-rakers. In the outer row on the front
of the first branchial arch Salmo trutta has on an
average 14—16, Salmo salar 17—20, Salmo umbla
23 or 24.

Hence it is clear that, in the great majority of
cases, the Blanklax must come nearer the Charrs; but
that the latter have not been developed directly from
the former, appears from other relations, in which they
stand nearer the Grdlax. One of these relations is
afforded by the character which, to the best of our
knowledge, gives the only tenable distinction between
the Salmons and the Charrs, namely the size of the

SALMONS.

835

scales, expressed in their number. The scales are gene-
rally larger in Salmo salar than in St trutta, but larger
even in the latter than in S. umbla. In a longitudinal
row extending along Vio °f the length of the body, we
find 1) on the anterior part of the belly above the la-
teral line in S. salar on an average 18 scales, in S.
trutta 18 — 21, and in S. umbla 30, 2) on the tail in
S. salar 15 or 16, in S. trutta 16 or 17, in S- umbla
26. Thus the Charrs have probably been developed
from a race nearer to the Gralax than to the Blanklax ;
but the development has followed the same course and
produced the same results in the differentiation of form.

The natural relation between the Charrs is indeed
more complex, but all the more instructive. In Scan-
dinavia we have two forms, the Northern Charr ( Salmo
alpinus ) and the Saddling ( S . salvelinus), whose right to
a specific rank is almost as certain as the difference
between the Gralax and the Blanklax , and which occupy
the same developmental relation to each other, 8. alpi-
nus representing the younger stages of growth, S. sal-
velinus the older ones. Besides these forms however
— in order to gain a correct understanding of the re-
lation between them — we must take into consideration
a third form which strictly belongs to Greenland and
the northern regions of North America, and for which
we may employ the specific name given it by Fabri-
ciusrt, Salmo stagnalis1'.

On closer examination of these three Charr forms,
we see that in many respects0 the averages run in an
uninterrupted series from alpinus through stagnalis to
salvelinus, i. e. alpinus represents the youngest stages,
salvelinus the oldest. Here, for the sake of brevity,
we can only give one example.

The least depth of the tail in proportion to the
length of the head diminishes as a rule in the Charrs
with increasing age. In order more carefully to test
the particulars of this relation and others, I have di-
vided the material possessed by the Royal Museum
into six groups according to age. As we have no cor-
responding terms in popular phraseology, I have denoted
these groups by Roman numbers. The first group (I)
contains the specimens under 228 mm. in length.
Among all these specimens (of all forms and interme-
diate forms) the average length of the body is 188 mm.

The second group (II) contains the Charr between 228
and about 355 mm. long. The average length of body
in this group is 303 mm. In the third group (III)
the maximum length of body is about 458 mm. and
the average length 387 mm. In the fourth group (IV)
these measurements are respectively about 538 mm.
and 494 mm. The fifth group (V) contains only three
specimens, all about 650 mm. long. The sixth group
(VI) consists of only one specimen, a male from Spitz-
bergen, measuring 757 mm.

The alterations caused by growth in the relation
between the least depth of the tail and the length ot
the head, are most regular in our specimens of stag-
nalis, being in this form:

Average in

ii.

III.

IV.

V.

C?

34.0

32.9

30.4

9

35.9

33.6

—

32.6

The averages in the

second group (II)

are:

Average

n

alpinus.

stagnalis.

salvelinus.

d"-~

36.5

34.o

29.4

9 -

37.4

35.9

29.9

The form

series, expressed in

averages

for all oui

specimens, is as follows:

Average

alpinus.

stagnalis.

salvelinus.

Least depth of
of the head.

the tail in % of the length

36.7

33.2

29.2

AH these averages follow a regular course, con-
stantly decreasing. Such is the case where the form
series has retained its original type, and in the rela-
tions where the males evidently represent the highest
stage of development. In several relations, however,
the females have a special representative in stagnalis,
arid the original form series has thus been modified
in a singular manner. We have an example of this
in the proportion between the length of the maxillaries

“ Fauna Groenlandica , p. 175.

b In order to simplify the question, we here omit some intermediate forms, the significance of which we may disregard for the
time being.

c In the work quoted above ( RiJcsmus . Salmon.) I have adduced 18 such points.

836

SCANDINAVIAN FISHES.

and that of the body. The general form series in this
relation is the following:

Average

alpinus.

stagnalis.

salvelinus.

Length of the maxillaries in % of that of

the body

7.9

7.6

9.8

Average distance of the ventral fins from
the tip of the snout in % of the
length of the body.

I.

11.

111.

IV.

V.

(cf

Salmo salvelinus <

51.7

51.4

52.6

54.3

—

1.9

51.4

51.5

51.9

52.2

—

I d*

Salmo stagnalis <

—

48.7

49.i

50.3

—

1 9

—

46.i

47.9

49.4

47.o

I’c?

Salmo alpinus <

19

47.5

48.2

48.i

—

—

40.4

47.2

47.7

—

—

Here the series is irregular: alpinus occupies an
intermediate place in the averages, stagnalis having a
lower average, and salvelinus the highest of all. We
arrive at the same result, if we examine the several
stages of growth; e. g. the third (III):

Average

stagnalis.

alpinus.

salvelinus.

Length of the maxillaries in % of that

C?

7.8

9.9

9.9

of the body <

1 9

7.3

8.1

9.6

Ranged in this manner the form series ascends
with fair regularity, at least in the females, towards
salvelinus; and the changes of growth are also in di-
rect proportion to age. Thus, for example, we find

In this relation, as in many others — we will
not give more examples here — the influence of a
marked sexual difference on the development of form
has ranged stagnalis, with the female characters, on
one side of alpinus, and salvelinus , with the male cha-
racters, on the other. If alpinus were omitted from
the comparison, or if we were ignorant of its existence,
the relation between salvelinus and stagnalis would
seem almost exactly similar to that between trutta
and salar.

The character, derived from the situation of the
ventral fins, which as a rule — not. entirely free from
exceptions — separates alpinus from salvelinus, is mere-
ly an expression of the extremes in an evolution of
form still proceeding. Its signification appears without
difficulty from the following table of averages:

The changes of growth here follow a regular
course with increasing averages, at least in the females
of salvelinus and alpinus and in the males of stag-
nalis. Nearly all the males have greater averages than
females of the same form and age. The form series,
expressed in averages for all our specimens, is as
follows:

Average

alpinus.

stagnalis.

salvelinus.

Distance of
the snout

the ventral fins from the tip of
in % of the length of the body

47.4

48.5

52.o

Not only the form of the Salmones, but also their
manner of life indicates their common descent. It
seems most probable that the Salmons were originally
marine fishes of an arctic or subarctic (boreal) region,
that for procreative purposes they made their way
into arms of the sea and the mouths of rivers, and
that by a stronger and stronger development of this
roving disposition they were transformed to fresh-water
fishes for a great part of their life, or finally changed
their abodes for all time. Among the Scandinavian
Salmons the Blanklax ( Salmo salar ) represents the most
migratory type, the Saddling ( S . salvelinus ) is most
completely a fresh-water fish. As there is nothing in
the form characters to indicate an evolution of true
Salmon from Charr, but everything seems to favour
the converse opinion, it is most natural to assume that
the primitive forms were marine fishes.

During their life in fresh water the Salmons ac-
quire a coloration quite different from their sea dress:
the epidermis becomes tumid, conceals the scales in a
greater or less degree, and assumes a darker colour,
gray, grayish or greenish brown, partly black and red
or yellowish, developing at the same time spots of black
and red. But this coloration appears, in general cases,
only as the generative organs approach maturity; and
at times the Salmons remain long enough in fresh
water before these organs are fully ripe for spawning.

SALMONS.

837

In great rivers, such as the Rhine" and Loire6, at least-
some Salmon ha-ve been observed to pass a whole year
in fresh water under these circumstances. Neither the
spawning dress nor any of the other external characters
of sex — the most remarkable of them being a carti-
laginous projection, turned upwards and recurved, which
•appears at the point of the lower jaw in the males
ready to spawn, and fits into a cavity in the upper
jaw — is developed in such specimens. The Salmon
which observers have traced in this condition for two
spawning-seasons are consequently incapable of spawn-
ing until two, or perhaps three years have intervened.
In Sweden, to the best of our knowledge, no direct
observations on this head have been recorded; but in
the Dal Elf, especially at Elfkarleby, specimens called
OJclor, with bright (blank) dress and undeveloped gene-
rative organs, have long been known. These specimens
are bright Gralax, resembling in coloration the Blank -
lax, and in other parts of Norrland, where they are
also found in the Gulf of Bothnia, they bear the names
of Borting and Tajmen. In Lake Wetter they are re-
presented by the so-called Silfverlax. From Lake We-
ner Widegren c records the occurrence of “adult indi-
viduals of Salmo solar which in autumn had neither
matured roe nor ripe milt”. Thus we possess trust-
worthy evidence from several parts of Sweden as well,
to the effect that the Salmons do not invariably spawn
every year, even if they remain in fresh water the
whole time. Widegren remarks the same circumstance^
in the reproductive operations of the Charrs.

Quite natural is the consequence that the Salmons
require a long time to recover the losses which their
frame has suffered in propagating their kind. These
losses are so great that many, perhaps most, of the
older specimens, especially the males, die of exhaustion.
All their fat and the greater portion of their flesh have
been consumed, partly as a source of nourishment dur-
ing the period when the fish entirely abstains from
food, partly in the ripening of the sexual organs. In
England these fish are known as Kelts , on the River
Nissa in Ha-lland they are called pejsor. Miesciier-

Ruescii has shown8 how the great lateral muscles of
the body and the dermal muscles undergo a fatty de-
generation and yield the greater part of their volume
to develop the ovaries; and a comparison (fig. 209)
between the conditions of the intestinal canal of a
male salmon before and after the spawning, is a strik-
ing illustration of the great changes which take place
in the body, while the spawning-dress is in course of
development.

a b

Fig. 209. Intestinal canal and testes of a Salmo salar before (a)
and after ( h ) the latter have swollen to maturity. After Buckland.
Both figures reduced, a from a clean run fish weighing 35 lbs. and
taken in the Avon on the 14th of March, 1871; b from an unclean
(breeding) Salmon weighing 13 lbs. and taken on the 19th of Janu-
ary, 1871. os, oesophagus; i, intestine; p, pyloric appendages; m,
testes. In a both the pyloric appendages and the intestine are em-
bedded in fat, and the testes thin; in b the testes are distended, and
both the pyloric appendages and the intestine are without adipose layers.

Statements have been made, it is true, which appa-
rently show that in the sea, at least under favourable
circumstances, Salmon may gain in strength and fatness
with astonishing rapidity. From experiments made in
1855 at Stormontfield (a Salmon-breeding establishment
on the Tay in Scotland) — as well as from still earlier
experiments (1795Q in the River Berridale (Caithness)
— - it was concluded that Salmon-fry which left the
Tay at the end of May, weighing at most 57 grammes,

a See Miescher-Ruesch, Statistische und biologische Beitrcige zur Kentniss vom Leben des Rheinlachses im Susswasser , Intern. Fischerei-
Ausstellung in Berlin 1880: Catal. Schweiz, p. 170.

b See Kunstler, Rech. s. la reproduction clu Saumon de la Dordogne, Congr. Intern, de Zoologie, Paris 1889, Compte-Rendu des
Seances, p. 83.

c Ofvers. Vet.-Akad. Forh. 1864, p. 294.
d ibid., p. 295.
e 1. c., pp. 186 etc.

f Day, British and Irish Salmonidce, p. 93.

838

SCANDINAVIAN FISHES.

returned in the beginning of July, about six weeks
after their departure, having attained in the sea a weight
of IV3 kilo; and at the end of July others came back
which had grown to a weight of nearly 4V3 kiloa.
These experiments, however, were not carried out with
sufficient precautions to render the results convincing6.
The growth of older Salmon is better known, and,
though not so rapid, is still considerable. In 1859 the
Duke of Athol marked three Salmon, weighing respec-
tively 10, ll1/,, and 12Va lbs. (4 7, — 5 2/3 kilo), then
on their way to the sea, and took them again six months
afterwards, as they were returning to fresh water, when
their weights were respectively 17, 18, and 19 lbs.
(7 3/4 — 87s kilo.)c. Even if these Salmon could have
attained a breeding condition the same year, we have
other observations and circumstances which indicate that,
as a rule, their stay in the sea is longer. In February
and March Scrope marked several Kelts, weighing 4
lbs., from the River Shin in Sutherland, and on taking
them again in June and July of the following year,
found them to weigh 9 — 14 lbs/ The Salmon which
descend into the Gulf of Bothnia from the rivers of
Norrland and Finland, rove from the said gulf down
to the south of the Baltic. As Gisler observed in
I752d, as Stecksen remarked a century later6, and as
Malmgren quite recently ascertained7, in the rivers of
Norrland and Finland Salmon are often caught which
have hooks in the jaws or stomach that they have torn
loose from long-lines in the south of the Baltic, even
on the Pomeranian coast. Journeys of such length pre-
sumably demand too long a time for a Salmon to de-
scend early in spring into the Baltic, exhausted by the
exertions of the spawning, and to return the same year,
even at so late a season as June — August, and ascend
one of the rivers in breeding- condition. To this we
should add that in the south of the Baltic line-fishing
for Salmon is practised chiefly, if not exclusively, in
autumn and winter, so that we have all reason to be-
lieve that these Salmon must have wintered in the
south. It is also quite possible that a sojourn in the

brackish water of the Gulf of Bothnia — as in the fresh
water of Fake Wener — can replace the year’s stay in
fresh water previous to the spawning, required by the
so-called Winter Salmon of the Rhine. This is all the
more probable now that Mr. Andersson’s experiments
have shown (see Fonnberg, Bill. Vet.-Akad. Hand].,
Bd. 18 (1892), Afd. IV, No. 2, p. 10) that the gene-
rative organs can ripen, even if the fish be detained
in the Gulf of Bothnia while its fellows are preparing
to spawn in the rivers.

The Blanklaxar thus pass the greater part of their
marine life and, at least in certain cases, a considerable
portion of their life in fresh water, without the deve-
lopment necessary for breeding and without the cha-
racters that mark their spawning-dress. This is also
true of the Gralaxar, with the single exception that,
for the most part, they live between the spawning pe-
riods in lakes. Yet it is no very simple task to ex-
plain all the names which these fishes have received
in different dresses and at different stages of sexual
maturity.

Siebold advanced the opinion g that the “Silver
Salmon” ( Schwebforelle , Maiforelle ) which are found in
the alpine regions of Central Europe, are persistently
sterile Gralaxar ( Grundforelle , Lacks f or elle); and that
such individuals do occur, is also maintained by Fatio
in Switzerland. But Widegren showed that the cha-
racters of the former are inconstant, being more and
more approximated to the Grdlax type as their sexual
organs are developed, and that they probably do attain
maturity, “though6 several years may possibly elapse
before a sterile individual becomes fertile and acquires
the characters typical of the generative power.” On
closer examination of this so-called sterility, however,
we find4 that it is attended with characters which are
a combination of Blanklax and Grdlax characters — the
alleged difference between “fertile” and “sterile” Gra-
laxar is really the same as that between Gralaxar and
Blanklaxar — and the same combination of characters
occurs, as is natural, in hybrids between Salmo solar

a Brown, The Natural History of the Salmon etc., pp. 49 and 52.
b Russel, The Salmon , p. 54.
c Day, 1. c., p. 95.
d Vet.-Akad. Hand], 1752, p. 100.
e In Nilsson, Skand. Fn., Fisk., p. 381.

f Bohuslansk Fiskeritidskrift, H. 1 — 3, p. 50. In the Great Belt Fiedler (Nordisk Aarsskrift for Fiskeri, 1884, p. 24) found a
similar brass hook in the mouth of a Salmon caught in April south of Corsoer.
g Sussivasserfische Mittel.europas, p. 301.
h Ofvers. Vet.-Akad. Forh. 1864, p. 292.

1 Smitt, Riksm. Salmon, 1. c., pp. 83 etc.

SALMONS AND CHARRS.

839

and S. trutta. Now hybridism does not- necessarily in-
volve sterility; and with regard to the Silver Salmon
and similar forms, it may well be difficult to determine
in each case whether it is an altered manner of life —
a longer stay in the sea or large lakes — or hybridism
that has brought about the said changes of form. But
it appears to be demonstrated that these changes attend
either or both of the above conditions.

In Sweden and, generally speaking, in all temperate
countries, the Charrs are true fresh-water fishes; but
such is not the case in the arctic parts of their geo-
graphical range. There they live like the Salmon; Es-
mark a and Hetting6 found them in the Arctic Ocean
off Norwegian Finnmark, and off Spitsbergen they are
the favourite food of the Beluga (Delphinapterus lencas),
which follows them to the head of the gulfs and bays.
In the sea they wear the colours of Silver Salmon,
and do not acquire their proper Charr dress before
entering the rivers.

Characters both of form and colour indicate in
their variations a close relationship between the Sal-
mons. Their differentiation seems to be of late origin,
and they should perhaps be regarded as species which
are still developing constancy of characters c.

The skeleton of the Salmons is feebly ossified in
sharp contrast to that of the preceding family. The
cranium is for the most part persistently cartilaginous,
and most of its bones lie as separate covering-bones
on the almost continuous capsule of cartilage. The
occipital ridge is merely a short terete process, extend-
ing about as far back as the ordinary processes from
the mastoid (epiotic) and squamose (pterotic) bones.
The posterior oculo-inuscular canal is large, and pe-
netrates the occipital region behind, being covered
below by the parasphenoid bone, which is strongly
bent in the sphenoidal region, and centrally divided
in front by the Y-shaped ba.sisphenoid bone, which is
wanting in the preceding family. Here, on the other
hand, as in the following genera, there is no pha-
ryngeal process. The orbits are separated internally
by a thick wall, composed of cartilage and the orbito-
sphenoid bones. The osseous framework of the upper
jaw, as we have mentioned above, is quite different
from the corresponding structure in the Cyprinoids.

The maxillaries articulate in front, like the intermaxil-
laries and the palatine bones, and close between these
two pairs, with the cartilaginous rostral part of the
cranium (the rostro-ethmoidal cartilage). But they
have undergone considerable elongation — the inter-
maxillaries, on the other hand, are comparatively short
— and are furnished with teeth, as well as the inter-
maxillaries and the palatine bones. Their anterior part
is, however, covered below by the intermaxillaries, so
that their externally visible length — as it is given in
the above definition of the genus — is measured from
the hind extremity of the latter bones. Above their
posterior part lies the covering-bone (os supplement, are )
which answers to the cheek-bone (os jugate) of the
higher vertebrates. The vomer is set as a covering-
bone on the middle of the under surface of the rostro-
ethmoidal cartilage and on the anterior extremity of
the parasphenoid bone. In the Salmons its shaft (body)
is fairly long and broad, earinated on the under (outer)
surface, but thin; in the Scandinavian Charrs, on the
other hand, ossified hardly at all or only on the sides.
The dentition of the vomer is also confined, as a rule,
in the latter to the anterior part, the so-called head;
while in the Salmons it extends, at least during youth,
to the shaft as well, but is generally more and more
reduced with age. The other bones of the palate, ex-
cept the palatine bones proper, are toothless in this
genus. The pterygoid bones are slender and curved in
the ordinary manner to unite the palatine and quadrate
bones. Within them lie the mesopterygoid bones, which
are thin, but broad behind, and form the greater part
of the roof of the mouth. According to Lilljeborg
these two bones afford a constant distinction between
Trout and Salmon. In the former, he says, their breadth
decreases from their middle point even in a backward
direction, whereas in the latter they grow broader all
the way from the anterior to the posterior extremity.
A great difference from the preceding family appears
in the branchiostegal membranes, which are fur-
nished with a far greater number of rays (9 — 12, in
exceptional cases 8 or as many as 14, in Oncorhyn-
chus sometimes 16). These rays are, however, of the
same broad, blunt, sabre-like form as in the Cy-
prinoids.

a Skand. Naturf. Mode, Chrnia 1868, Forh., p. 527.
b Smitt, 1. c., tab. metr. VI, No. 399.

c “In numerous cases one is much tempted to ask whether we have not to deal with a family which, being one of the most recent
creation, is composed of forms not yet specifically differentiated”: Gunther, Cat. Brit. Mus., Fish., vol. VI, p. V (preface).

106

Scandinavian Fishes.

840

SCANDINAVIAN FISHES.

The spinal column is also marked by its weak
ossification. The neural arches proper (Owen’s neurapo-
pliyses) are persistently cartilaginous in most of the
abdominal vertebrae, and the bases of their neural
spines are united to the bodies of the vertebrae only by
a suture or, in young specimens and in the anterior
part of the column, by a mobile articulation. In the
forepart of the skeleton these spines are also but loosely
united to each other, the right to the left in each ver-
tebra, and with the hind surface of each base articulates
a rib-like muscular bone, directed outwards, backwards,
and upwards (Owen’s diapophysis). In the posterior
part of the abdominal region the cartilaginous neural
arches disappear, the bases of the neural spines take
their place and develop a firmer osseous connexion with
the bodies of the vertebras, the neural arches thus
formed being strengthened at the same time by the
coalescence of the spines on each vertebra. The diapo-
physes decrease in length, and disappear near the caudal
region; but simultaneously there appear, though with
only slight development, the knob-shaped articular pro-
cesses {zyg apophyses) that in the Cyprinoids are gene-
rally so prominent on the anterior abdominal vertebrae,
one pair in front of and one pair behind the neural
arches, which are here most constant in the caudal
region, though even there they may be wanting. The
development of the haemal arches proper (Owen’s hcem-
apophyses) takes the reverse direction. It advances from
the front of the body to the posterior part thereof: on
the first vertebra they are scarcely distinguishable ex-
ternally from the body of the vertebra; on the second
vertebra they are distinct protuberances, one on each
side of the lower part; on the third each of them is
furnished with a rib; and they thus continue to in-
crease until, near the caudal region, a succession of
haemal spines appears, each bearing a pair of ribs on
the hind surface of its top, while the haemal spines of
each vertebra approach each other at the top, forming
(usually on the last six abdominal, as well as on the
caudal vertebrae) a closed haemal arch. On the same
vertebrae as the haemal spines, there also appear lower

articular processes similar to the upper. The ribs are
slender, in sharp contrast to those of the Cyprinoids.

In the shoulder-girdle, the structure of which is
else the same as in the preceding family, the clavicle
is thin and broad, with the upper arm comparatively
short, and the postclavicle consists of three, sometimes
four parts, namely two (sometimes three) upper, thin
and flat, and one lower, pointed and rib-like. This
division of the postclavicle recurs in the Herrings.

The pelvic bones are simple and triangular, with-
out the indentation in front, and with only a rudiment
of the process behind, which we have seen in the Cy-
prinoids. But remnants of the older piscine types’
radialia (mobile basal bones of the ventral rays) and also
of the true pelvis of the said types, have been detected
in the Forell stage by Davidoff".

The intestinal canal is rather simple and short, as
in most predatory fishes, its entire length, when ex-
tended, being about equal to the length of the body or
somewhat less than the latter. The stomach is only
faintly marked off from the rest of the canal; but a
remarkable point is the thickening of its walls which
attends a diet of mollusks or other testaceans* 6. This
thickening lias been most frequently observed in the
so-called Gillaroo Trout of Ireland, a form which has
hence been named by Gunther Salmo stomachicus ; but
it may also be observed in common Lake Trout0. The
pyloric part is directed forwards, as well as the be-
ginning of the true intestine (duodenum), which is ex-
ceedingly well furnished with csecal diverticula. These
appendages, however, vary considerably both in size
and number, from about 30 to nearly 100. They are
generally most numerous in the Salmon proper, where
Krdyer6* has counted 96, though the usual number in
this form is between 50 and 70. In the Trout that
migrate to salt water, the usual number is about the
same; but in those that live exclusively in fresh water,
it is generally less, about 30 — 50. Thus in different
specimens of Trout 29 — 69 pyloric appendages have
been counted0. After the duodenum has advanced al-
most to the diaphragm, the intestine abruptly bends

a Morph. Jahrb., XVI (1880), p. 464, taf. XXI, fig. 5.

6 According to Day a seagull at Hunter’s Museum was fed for some time on corn, and the muscular layers of its stomach were
thereby thickened as in a crop; and from Holmgren’s experiments on pigeons at Upsala it appeared that the muscular layers in their crop
were reduced by a flesh diet.

c It is further known that a molluscous diet gives the body of the Salmons a singular, orange ground-colour.
d Damn. Fislce, Bd. 2, p. 558.
e Day, British Salmonidce , p. 188.

SALMONS AND CHARRS.

841

straight back to the vent; and in the posterior part of
the intestine or the rectum, which is usually somewhat
wider than the anterior part thereof, the mucous mem-
brane is raised on the inner surface, at least in old
specimens, in the shape of transverse rings, here and
there united like the thread of a screw, instead of the
longitudinal folds which it forms in the anterior part
of the intestinal canal from the oesophagus. The liver
is almost one-lobed, the right lobe being extremely
little developed. The place of the latter is occupied
by a rather large gall-bladder, the discharging duct of
which opens into the duodenum at the beginning of
the pyloric appendages. Before the spawning the spleen
is rather large and sometimes extends from the bottom
of the stomach beyond the insertions of the ventral fins;
but after the maturation of the generative organs it
shrinks considerably, and does not begin again to in-
crease, until its functions are laid under contribution
at the commencement of the above-mentioned trans-
formation of fat and flesh into material for the deve-
lopment of the sexual organs (cf. Miesci-ier-Ruesch,
1. c.). The ovaries, as mentioned above, show the pe-
culiarity that they are without special oviducts, the ripe

eggs falling into the abdominal cavity, whence the)' are
expressed through the genital aperture behind the vent.
The testes, on the contrary, are furnished as usual
with vasa deferentia. The air-bladder, which is simple
and long, extends throughout the length of the ab-
dominal cavity, and the pneumatic duct opens on the
dorsal side of the oesophagus. Still more elongated are
the kidneys, which penetrate behind into the hamial
canal of the tail.

The ancient Greeks have not bequeathed to us any
name for the Salmon, which does not occur as a ma-
rine form in Mediterranean regions; but according to
Apostolides" the Trout inhabits most of the rivers in
Greece, and bears among the modern Greeks the name
of 7iioTQO(f>ab. 8almoc is of classical Latin origin and
occurs in Pliny L Among the fishes of the Moselle
Ausonius enumerates Salar (the Parr), Saimo , and
“the form that ranks between them”, Farioe (the Trout).
Such is the explanation given by Figulus (1540) of
these names. Trutta belongs to mediaeval Latin7. Umbla
is a Latinized form of the French Ombre or JJmble.
Salvelinus has arisen in the same manner from the
German Salmling or Sdlbling.

THE CHARR (sw. rodingen).
SALMO UMBLA.

Plate XXXVII, fig. 1 (c/1) and 2 ($).

Scales exceedingly small, their number in a longitudinal row above the lateral line just in front of the perpendi-
cular from the beginning of the dorsal fin and extending along Vio °f the length of the body at least 25 (as a
rule 27 — 3d9), and in a row of similar length above the lateral line behind the perpendicular from the beginning
of the anal fin at least 21 (as a rule 23 — 30, sometimes 33). Sides of the body marked with more or less
distinct and light, red spots on the darker ground-colour , but no black spots behind the head.

R. hr. (9)10-12(13—14); D.

3—4

8—10(11—12)

; A.

3—4

7—9(10)’

P.

1

V.

1(2)

11 — 14 7—8(9)

ca 190—240; Vert. 62— 631.

; C. oe + 1 + 17 + 1+ *; Ser. squ. lat,.1

Syn. Timbre et Umble , Belon, Nat., Div. Poiss ., pp. 280 et 281;

Saimo Lemani lacus sive Umbla, Rondel., Pise. Lacustr.,
p. 160. Saimo lineis lateralibus sursum recurvis, cauda
bifurca, Art., Ichthyol., Gen., p. 18; Syn.. p. 25; Saimo
vix pedalis, pinnis ventris rubris, maxilla inferiore paullo

a La Peclie en Grece, p. 33.

b TCeTQO(pa(l) — phps. answering to the Scand. stenbit and ror, which denote the preference of the Trout and Charr for stony bottoms.
c From salire, to leap.
d Hist. Nat., lib. IX, cap. 18.

e Another, perhaps better reading is Sario, a derivative from the “same root as salar and saimo.

■ Derived from the Latin trudere , to thrust, force; a fish which forces its way against the stream.

g 18 — 20 scales on an extent answering to I/4 of the length of the head, as Nilsson expresses this character: 16 — 20, according
to Lilljeborg.

h Counted above the lateral line; in the lateral line itself about 125 — 145 (Day).

* Sometimes 65, according to Fatio, sometimes 59 (in the Windermere Charr), according to Day, or 58 (in var. fluviatilis ), accord-

842

SCANDINAVIAN FISHES.

longiore, Id., ibid., et Spec., p. 52. Salmo dorso nigro,
lateribus casruleis, ventre fulvo, Lin., Fn. Suec., ed. I, p.
117. Roding , Lin., It. Wgoth., p. 257.

Salmo umbla, Lin., Syst. Nat., ed. X, tom. I, p. 310; Penn.,
Brit. Zool. (1776), vol. Ill, p. 267; Agass., Rep. Brit.
Assoc. Edinb. 1834, p. 617; Yarr, Brit. Fish., ed. 2, vol.
II, p. 121; Agass., Poiss. d'eau douce, tab. IX — XI; Thomps.,
Nat. Hist. Irel., vol. IV, p. 160; White, Cat. Brit. Fish.,
p. 78; Smitt, Riksm. Salmon., Vet.-Akad. Handl., Bd. 21
(1885), No. 8, p. 163; Fatio (Salvelinus), Fn. Vert. Suisse,
vol. V, p. 395.

Salmo erythrceus , Pall., Zoogr. Ross, Asiat., tom. Ill, p. 349.
Salmones salvelini, (—Salmo ventrieosus + S. carbonarius (ex
StrSm) + S. alpinus (ex Lin.) 4- S. pallidus + S. salvelinus
(ex Lin.) + S. rutilus ), Nilss., Prodr. Ichtliyol. Scand., p.
7 ; — unam speoiem ovnnes has varietates censuit in Skand.
Fn., Fisk., p. 422.

Salmo alpinus, Mgrn, Finl. Fiskfn., (disp. Helsingf. 1863),
p. 56; Widegr., Landbr. Akad. Tidskr. 1863, pp. 201 et
209; Mgrn, Ofvers. Vet.-Akad. Forli. 1864, p. 534; Coll.,
Forh. Vid. Selsk., Chrnia 1874, Tillsegsli., p. 160; ibid.
1879, No. 1, p. 86; Malm, Gbgs, Boh. Fn., p. 540; Smitt,
(Jfvers. Vet.-Akad. Fork. 1882, No. 8, p. 33; Mela, Vert.
Fenn., p. 343; Day, Fish. Gt. Brit., Irel., vol. II, p. 112;
Brit. Salm., p. 237; Trybom, Iaktt. Fisk. Ume-Lappm.,
Nord. Aarsskr. Fisk. 1883, p. 300; Reuter, Sundm., Finl.
Fisk., tab. V; Lillj., Sv ., Norg. Fn., Fisk., vol. II, p. 598.
a: Var. Salmo salvelinus ( Vetterns Roding, Nilss. — Storrodingen,
Lillj.), cujus pinnae ventrales pone mediam longitudinem
corporis sitae sunt.

Syn. Salvelin Germanis, Ray, Syn. Metli. Pise., p. 64; Salmo
pedalis maxilla superiore longiore, Art., Ichtliyol., Gen., p.
13; Syn., p. 26.

Salmo Salvelinus, Lin., Syst. Nat., ed. X, tom. I, p. 309;
(+ S. Salmarinus, p. 310); Bl., Fisch. Deutschl., pt. Ill,
p. 149, tab. XCIX ( 4- S. umbla, p. 154, tab. Cl); Cuv.,
Val., Hist. Nat. Poiss. tom. XXI, p. 246 (+*!?. umbla,
p. 233); Nilss., Skand. Fn., Fisk., p. 422; IIckl, Kr,
Siisswasserf. Ostr. Mon., p. 280 ( 4- S. umbla, p. 285);
Sieb., Siisswasserf. Mitteleur., p. 280; Canestr., Fna D'ltal.,
pt. Ill, Pesci, p. 23; Lillj., 1. c., p. 599.

Salmo umbla, Jur., Hist. Poiss. L. Lem., Mem. Soc. Phys.,
D’Hist. Nat. Geneve, tom. Ill, pt. 1, p. 179, tab. V; Rapp,
Fisch. Bodens., p. 32, tab. V; Mor., Hist. Nat. Poiss. Fr.,
tom. Ill, p. 530.

b: Var. Salmo alpinus (Vermlands och Lapplands Roding, Nilss.
— Smarodingen, Lillj.), cujus pinna? ventrales ante mediam
longitudinem corporis sitae sunt.

Syn. Salmo alpinus, Lin., Syst., 1. c.; Ascan., Icon. Rer.
Nat., cah. 2, p. 7, tab. XVIII; Laistadius, Journ. Lappm .,
Forts., p. 75; Cuv., Val., 1. c., p. 249; Nilss., Skand.
Fn., Fisk., p. 426 (4- S. carbonarius, p. 429 4- S. rutilus,
p. 430); NystrSm, Iaktt. Fn. Jemtl. Vattendr. (disp. Up-
sala 1863), p. 11; Olsson, Ofvers. Vet.-Akad. Forh. 1876,
No. 3, p. 14; ibid. 1882, No. 10, p. 50; Lillj., 1. c,
p. 609.

The Charr, which is, generally speaking, the small-
est Scandinavian Salmo , still attains a fair size even in
our fauna. It sometimes reaches, at least in Lake
Wetter, a length of nearly 7xj2 dm. and a weight of
about 8V3 kilo“. Trybom was told at Lake Stor-Uman
in Ume Lappmark that in 1878 a Charr had been
taken there which weighed 7V2 kilo., and that it often
runs to 4 kilo, in the same lake, though its usual
weight is lV3 — 2 kilo. In many parts of Sweden a
Charr 1 foot long (3 dm.) or 1 1/2 lbs. in weight (0*7
kilo.) is considered quite an average-sized fish, and in
many lakes it does not attain even these dimensions.

The body shows in young and middle-sized Charr
the most handsome and best proportioned piscine type,
a terete and regular fusiform shape, so little compressed
that the greatest thickness is about half the greatest
height, which lies a little in front of the dorsal fin
proper, and is contained about 5 times, or in young
specimens even more than 6 times, in the length of
the body. With age, however, both the relative depth
and the lateral compression as a rule increase — the
former most, as usual, in gravid females, the latter in
spent fish — and in our largest Charr the depth some-
times rises to 29 % of the entire length or 31 % of
the length to the base of the caudal fin* * 6, while the
greatest thickness of such specimens is only 2/5 of the
said depth0. During these modifications the sides of
the body become almost flat and parallel, converging
gradually and in a very elongated curve towards the
base of the caudal fin. The differences, however, seem
sometimes to depend on local circumstances: “We find
the species,” says Thompson (Yarrell, 1. c., p. 123),
“to be in one lake herring-like, and in another approx-
imating the roundness of an eel”. But both forms
sometimes occur in the same lake (see Nystrom, 1. c.).
The least depth measures about 7 or 8 % — in the Sib-
ling sometimes nearly 6 %, in the Northern Charr
sometimes nearly 9 % — of the length of the body, or
in the former about 28 — 30 %, in the latter about
34 — 39 %, of the length of the head.

The head shares in the above alteration of the
body, growing more compressed in old specimens, and
acquiring a longer, more pointed snout; but above it

a In the East States of North America there lives a Charr, Salmo Namaycush, which according to Brown-Goode ( Fisheries and

Fishery-Industries of the Un. States, sect. 1, p. 486) attains a weight of 120 lbs. (54‘43 kilo.).

6 According to Fatio the greatest depth may rise to 461/), % of the length to the base of the caudal fin.
c According to Fatio the greatest thickness may sink in very old specimens to fq of the greatest depth.

CIIAKR.

843

is always convex, never flat as in the Huch. Its length,
which is generally greater in the males than in the
females, varies in the Northern Charr between about

20 and 22 % of that of the body, in the Sadbling be-
tween about 23 and 24 % of the same. The eyes are
middle-sized or rather, in old specimens, small; they
are always set in the anterior half of the head and
fairly low, near the maxillary bones. They are nearly
round, the longitudinal diameter being only slightly
greater than the vertical, and varying between about

21 and 12 % (exceptionally 83/4 %) of the length of
the head; but the outer orbital margin is prolongated
in front to a point, and the anterior canthus formed
in this manner is filled by an adipose membrane. The
breadth of the convex forehead above the centre of the
eyes is about i/3 (30 — 35 %a) of the length of the head.
The nostrils of each side are set close to each other, on
about a level with the upper orbital margin and half as
far from the eyes as from the tip of the snout. The
anterior nostril on each side is round, with the margin
raised in a funnel; the posterior is obliquely placed and
oblong. The length of the snout, which in young spe-
cimens is shorter and blunter than in old, and in adult
males longer and more pointed than in females of the
same size, varies between 26 and 36 % of that of the
head. The mouth is large, owing to the elongation of
the snout even relatively larger in old specimens than
in young. The cleft of the mouth ascends somewhat,
though only a little. The teeth, the extent of which
we have above considered, are rather small, but strong
and pointed, with the tip recurved. The small teeth
on the copular parts of the hyoid bone are indeed ir-
regular in occurrence, but rarely wanting. The maxil-
laries, which are generally straight, but in old males
curved downwards behind, extend in the young hardly
behind the perpendicular from the posterior orbital
margin, but in old specimens some way beyond it.
Their length is greatest in the males, and varies in
Scandinavian Charr from about 35 to 46 % of that of
the head; and their greatest breadth, which thus be-
comes comparatively less as a rule in the males than
in the females and in old specimens than in young,
varies between about 25 and 18 % (exceptionally 15 %)

of their length. The lower jaw, which in most old
specimens projects a little beyond the tip of the upper
jaw, and has a small protuberance at the end that fits
into an indentation in the latter, measures in the males
about 60 — 75 in the females about 57 — 65 %, of

the length of the head. The preoperculum is crescent-
shaped. The operculum and suboperculum together
form a parallelogram, usually somewhat wider below,
and with the lower posterior angle rounded. The suture
between them, the length of which may serve as an
expression for the breadth of the opercular apparatus,
is comparatively shorter in old Charr than in young
and also as a rule, in the males than in the females,
its length varying between about 26 and 20 or even
17 % of that of the head. The branchiostegal mem-
branes coalesce with the isthmus in a line with the
anterior part of the eyes, or even in front of the per-
pendicular therefrom. The gill-rakers are fairly nu-
merous in comparison with those of the Iluch and
Salmon, numbering 23 (exceptionally 20) — 27 in the
outer row on the front of the first branchial arch, about
15 (13 — 16) of these being situated on the lower, pro-
jecting part of the arch. The inner row of spines on
this arch, of which spines at least rudiments are pre-
sent in the Salmons, is entirely wanting in the Scan-
dinavian Charrs. The pseud obranchise are distinct in
young specimens, concealed in old.

The true dorsal fin is obliquely quadrilateral (tra-
pezoidal), the length of the last ray being as a rule
less than half, but sometimes (in a form which has
been named alipesb) as much as 2/3, of that of the first
branched ray. The latter is as a rule the longest ray
in the fin, but in young Charr is sometimes a little
shorter than the second branched ray. The distance
between the fin and the tip of the snout increases even
relatively with age, this depending principally on the
prolongation of the snout, and is as a rule greater in
the males than in the females, varying between about
42 and 46 % of the length of the body. Its base mea-
sures about 10 — 11 %, and its height about 11 (ex-

ceptionally 10) — 15 (exceptionally 17) %, of the length
of the body. The adipose (second dorsal) fin, which
lies above the posterior part of the anal, is rather small

a In the Charr reared at the hatchery of Ostanback this percentage was lower, sinking even to 27. In Arctic Charr (var. stagnalis)
it may rise even to 39.

b Salmo alipes , Richardson, Fn. Bor. Amer., pt. Ill, p. 169, tab. 81 et 86 fig. 1. Cf. Smitt, Riksm. Salmonid ., tab. Ill, fig. 50,
a similar form, a male with strongly developed fins, taken in Tornea Trsesk.

844

SCANDINAVIAN FISHES.

in comparison with that of the Huch — though not so
small as in the Arctic stagnalis — being usually of the
same curved form and about the same size as in the
Scandinavian Salmons. The anal tin is similar in form
to the true dorsal, but always shorter and lower. The
distance between it and the tip of the snout is about
67 (exceptionally 65) — -71 % of the length of the body,
and as a rule this percentage is less than 70 in the
Northern Charr, more than 70 in the Saddling. The
caudal fin consists, here as in the preceding family, of

17 branched rays and an inconstant number of sup-
porting rays, of which the hindmost at each margin
extends cpiite or nearly to the end of the fin-lobe. This
fin is as a rule more or less deeply forked in the young;
but in old specimens, especially when in breeding con-
dition, the hind margin is truncate or slightly concave
when the fin is expanded. The middle caudal rays,
which occupy about 9 — 6 % of the length of the body,
generally measure less than half the length of the
outermost rays, except in old Charr, especially males,
where their length may rise to at least 56 % of that
of the latter.

The pectoral fins are pointed in old specimens,
even approaching to the scythe-shape of the Mackerel
type; but in the young they are blunter at the tip.
Their length varies considerably, from about 14 — 19 %
(in alijoes , cf, up to 2l72 %) of that of the body, and
is greater in the males than in the females. The pre-
abdoininal length (the distance between the foremost
points in the insertions of the pectoral and ventral fins),
on the contrary, is as a rule less in the males than in
the females, varying between about 27 (exceptionally
25) and 32 %a of the length of the body. The ventral
fins are obliquely triangular, with the truncate top as
base. They are also longer as a rule in the males
than in the females, their length varying between about
14 and 11 % (exceptionally 10 or, in alines, cf, as
much as 16 %) of that of the body. The postabdomi-
nal length (the distance between the beginning of the
anal fin and the foremost (outermost) point in the in-
sertions of the ventral fins), which is less as a rule in
the males than in the females, varies between about

18 (exceptionally 17) and 22 (exceptionally 26) % of
the length of the body.

The scales are both small and thin, without radi-
ating grooves, but with dense concentric striae. The

dorsal scales are oblong, elliptical or oval; those of the
lateral line of a more quadrangular shape but with
rounded corners; the ventral scales broader (deeper).
From the beginning of the dorsal fin obliquely back-
wards and downwards to the lateral line 34 scales, if
not more, may be counted in a transverse row, and
from the beginning of the adipose fin a similar row
contains about 21 — 24 scales. The lateral line runs
straight, or with a faint downward curve in front,
from the upper angle of the gill-opening to the middle
of the base of the caudal fin.

In its festal dress, just before the spawning, the
Charr is one of our most beautiful fishes. The back
is bluish or greenish black — the former predominant
in the Northern Charr, the latter in the Saddling — ,
the belly flame red or paler — the latter in the fe-
males— , and the sides of the body bluish gray or green,
with scattered, red or paler spots, varying in dimen-
sions but of about the same size as the pupils. The
dorsal and caudal fins are of the same colour as the
back, but distally they grow paler. In male Northern
Charr wearing this dress, the caudal fin is commonly
edged with red, and the lower margin (sometimes the
upper as well, at least in part) generally has the same
white or yellowish white hue as the anterior margin
of all the inferior fins. The rest of these fins gene-
rally partake, in the spawning-dress, of the same red
tint as the belly, but sometimes the posterior (the anal
and ventral) have a more or less predominant, grayish
blue colour, which occurs on the pectoral fins only in
a fainter tone, but has there a still more handsome,
ash-gray shade. The forepart of the belly and the
throat, as well as the branchiostegal membranes and
the lower jaw, are generally of a light, yellow or
whitish yellow ground-colour, more or less spotted with
bluish black. The head is commonly coloured above
like the back, below like the belly, with cheeks and
opercula of the same ground-colour as the sides of the
body; but it is often more or less spotted with bluish
or sooty black, and sometimes entirely, though faintly,
coated with one of these tints, which in the Northern
Charr generally extend to the mouth and pharynx as
well. The iris is brassy yellow, with an irregular,
ring-shaped shading of black.

These sharply defined hues of the spawning-dress
are exchanged during the intervening periods for a

When the belly is abnormally distended, this percentage may rise to 35.

CHARR.

845

lighter coloration, pale gray on the sides, light orange
on the belly; and when the Charr go out to sea, as
mentioned above, their sides adopt the silvery lustre
of the Salmon, on which ground the red spots are
hardly discernible, and sometimes, at least in the great
lakes, the ventral side is almost white.

These forms, as well as the Salmons, wear a Parr
dress during youth, most nearly resembling the pale
coloration just mentioned, but distinguished on the
sides of the body by about 13 — 15 transverse bands
of the darker dorsal colour, rounded below.

In coloration European Charr generally differ from
American, which are most often, but by no means
constantly, marked on the back and the dorsal and
caudal fins with vermiculate transverse spots and stripes.

In Scandinavia and the rest of Europe to the south,
the Charr strictly belong to the mountain lakes, and
hardly ever take up their abode in running water0.
In the Arctic regions — south to Northern Helgeland
in Norway* * * 6 — they are marine fishes which, like the
Salmons, ascend the rivers to spawn. In Lapland they
rank among the most common and most important
fishes, and the form which occurs there goes south in
the interior to Dalsland, Bohuslan, and Wester Goth-
land0'. But their range does not extend so high in the
mountain tracts as that of the Troutd, with which they
are often confounded, and they are probably not found
above the birch -region®. Whether the above-mentioned
large Charr that are met with in the greatest lakes of
Lapland, preserve the form-characters of the Northern
Charr, is unknown to us. The other large Scandina-
vian form has its true and best known haunts in Lake
Wetter, but is also met with to the north at least in
Lakes Storsjo and Hemsjo (Jemtland). East of Lake
Wetter Boheman found the Charr in 1836 in Lake
Oren. According to Nilsson and Widegren the Charr
inhabits Lake Sommen in Oster Gothland, but further
south in Sweden it is unknown. Strange to say, it is
wanting in Lake Wener, as well as in Lakes Malar

and Hjelmar. It evidently prefers clear lakes with
water of a low temperature — though as yet we have
not sufficient observations on this head to state a fixed
number of degrees — , and the singular features in its
geographical range — for instance, its occurrence in
certain lakes, and absence in others situated near them
and apparently of the same nature — cannot be ex-
plained until we have trustworthy information as to the
temperature of our lakes and the food which they offer
the Charr. That its aversion to turbid water is not
unconquerable, we see on the coast of Spitsbergen,
where Charr are most plentiful off the mouths of clayey
glacial rivers. But it is undeniable that water of a
high temperature, whether it be in the sea into which
the river inhabited by the Charr flows, or in lakes on
the lower course of the stream, stops the progress of
the Charr like a dam7.

According to the reports sent in to the Swedish
Fisheries Commission of 1881 — 83 the Charr is want-
ing in the Governments of Stockholm, Upsala, Soder-
manland, Kronoberg, Kalmar, Gothland, Blekinge, Kri-
stianstad, Malmohus, and Halland. In addition to Lake
Wener, the lower and southern districts of Sweden,
generally speaking, are thus without its geographical
range. It does not enter the Baltic, and is at least
rare in the lowlands on the Gulf of Bothnia. In the
valley of the Tornea Elf, for example, it does not de-
scend below Juckasjarvi.

In Norway the Charr is common, according to
Collett, in the lakes within the Government of Bergen,
less common in the Government of Trondhjem, and
occurs only in a few lakes within the Government of
Christiania. Lilljeborg found it often6' in the moun-
tain lakes of Central Norway, where it is commonly
called Iioe. The pale-bellied individuals are known as
Blekroe. Its manner of life in the north of Norway
and off the coast has already been noticed. Its habits
are similar on the Murman Coast and in Finnish Lapp-
mark; but its strict range in Finland is confined, ac-

a In this respect the European Charr is somewhat different from the true Salmons; but in North America a distinction is drawn be-

tween two varieties of Charr, the Lake Trout ( Salmo namaycusK), which attains a greater size than our Charr, but has the same habits, and

the Brook or Speckled Trout ( S . fontinalis'), which does indeed occur in lakes, but commonly spawns in streams.

6 Collett, 1879, 1. c.

c Lake Nedsjo in Bollebygd Hundred, see Malm, 1. c.
d Olsson, 1. c.
e Nilsson, Skand. Fn., 1. c.

f “South of New York they are effectually land-locked by the prevailing high temperature of the lowland streams, and are never able
to gain access to salt or brackish water”. Brown-Goode, The Fisheries and Fishery Industries of the United States , Sect. I, p. 502.
g dfvers. Vet.-Akad. Forh. 1844, p. 213.

846

SCANDINAVIAN FISHES.

cording to Mela, to the inland (eastern) tracts and
Lake Ladoga, while it is also found in the basin of
Lake Onega.

In Denmark the Charr is wanting; but in the
alpine regions of Southern Europe and in Great Britain
and Ireland it leads the same lacustrine life as in Swe-
den. In Switzerland it has been found, according to
Tschudi, at a height of 1,900 m.a, in Bavaria, accord-
ing to Siebold, about 600 — 800 in., and in Austria
about 400 — 700 m.* * 6, above the level of the sea. It
has there maintained its existence since the period when
a great part of Europe was covered with ice, and when
it could make its way in the cold seas to its present
abodes. In the lowlands between the Alpine Regions
and the North it is now wanting.

The Charr is a powerful and voracious fish-of-
prey, in proportion to its size not inferior to the true
Salmons. Its favourite hunting-grounds lie in deep
water, where it leads a sociable life; but it also ascends
to the surface to secure the flies and gnats that come
within its reach. Its food consists mainly, however,
of crustaceans, mollusks, and fishes, both large and
small; and it sometimes gorges itself on fish-roe, even
that of its own species. In the mountain lakes it has
generally to content itself with Entomostraca , which
are, however, plentiful enough, as a rule, to yield it
an abundant supply of food. Lynceus ( Eurycercus ) la-
mellatiis, a small crustacean of the group Cladocera, is
of particular importance in this respect, according to
Nystrom. In the lowland lakes and those of greater
dimensions the Charr finds more ample store of Gam-
maroids, mollusks, and fish, chiefly Gwyniads and Cy-
prinoids. But not even fishes armed with spines escape
its maw: in the stomach of a female Charr from Lake
Wetter no less than five Bullheads ( Coitus quadricor-
nis ) were found. In Lake Wetter, according to Wide-
gren, the Charr passes the greater part of the year in

the depths, and during spring and early summer is
seldom met with in less than 30 fathoms of water, but
after midsummer ascends towards evening into shal-
lower places, where its lustrous body strikes the eye as
it swims at the surface. During this season it also
makes its way into various inlets with a sandy bottom.
Towards autumn it retires to the shallows in the middle
of the lake, or visits some of the reefs and rocky pools
near shore.

The Charr spawns late in autumn and in winter",
in Sweden most commonly in October, the month dur-
ing which almost all the breeding Charr received by
the Royal Museum have been taken. In the fish-ponds
at Ostanb&ck, however, Mr. Lundberg caught a male
with running milt on the 10th of September; and in
the Jemtland lakes, according to Nystrom, the general
spawning begins during that month. At Stor-Uman
in Lycksele Lappmark Trybom was told that there the
Light Charr (the Greater Charr) commences to spawn
at the beginning of October, the breeding-season last-
ing 1 — 1 V2 weeks; but that the Black Charr (the Lesser
Charr) does not commence before the end of November,
and takes three weeks to spawn.

The Scandinavian Charr always select their spawn-
ing-places in a lake — according to all authenticated
observations d — by preference on a gravelly or stony
bottom in 2 — 4 m. of water, or sometimes, according
to Norback, in water so shallow that the dorsal fin of
the breeding fish projects above the surface. But they
frequently resort for this purpose to places near the
mouth of a stream. Nor do the Swiss Charr ascend
the rivers to spawn, according to Ratio; but they choose
water 20 — 60 m. (sometimes 80 m.) deep in the great
lakes e. In England, however, according to Varrell',
the Charr of Lake Winandermere generally make their
way up one of its affluents to breed, only a few of
them spawning in the lake itself.

a Fatio believes, however, that in Switzerland the Charr never ascends voluntarily to lakes more than 800 in. above the level of the

sea; but that it has, no doubt, been introduced by pisciculturists into lakes of greater elevation.

6 According to Heckel and Knee up to 1,900 m.

c In Bavaria breeding Charr are said to have been found at the end of June, and in Scotland the spawning-season is said to extend
from November to February or March (Day, Brit. Salmon., p. 244). Fatio gives instances from Switzerland (1. c., p. 409) of the Charr’s

spawning in the Lakes of Geneva and Neuch&tel both as early as June and as late as April, the usual months being November and December.

d According to Nystkom “it seemed in some places (for example Lake Holder) as if the ror (Charr) also resorted to running water
for the purpose of spawning”.

e The Deepwater Charr which the Royal Museum received from Lake Wetter though Sergeant-major Hall (1830, without specified
date) were in breeding condition, a circumstance which perhaps indicates that in Lake Wetter too the Charr may spawn at a considerable

depth. Nilsson also states that the so-called blanhroding (Bright Charr) spawns in 30 — 40 fathoms of water on a clayey or muddy bottom.

/ L. c., p. 125.

CHARI!.

847

From Sweden we have no observations of the
manner in which the spawning is performed; but in
Land and Water for the 14th of August, 1884, Jack-
son described the actions of a, female specimen in South-
port Aquarium when constructing a nest for her eggs.
“We obtained a fine shoal of char from Windermere at
the opening of the season this year. Soon after they
came, I saw one of them had not spawned, and was
busy making its nest. Its modus operandi was exactly
as described by Mr. Buckland. It swam slowly down
towards the selected place as though concentrating its
energies; when it arrived over the spot, it threw itself
partially on its side, and dropping the hind part of
the body, it gave several violent blows (three or four)
with its tail, scattering the gravel right and left. The
impetus of the blows not only scattered the gravel,
but drove the fish upwards in a slanting direction.
Quietly allowing the force to expend itself, it then
turned round, swam slowly, and repeated the process
time after time, until it had made quite a large hole.”
This observation inclines us to the opinion that in
essential respects the spawning of the Charr resembles
that of the Salmon.

The ova are comparatively large — from 4 to 572
mm. in diameter when ripe. Davy counted 1,230 eggs
in a female weighing 1/i kilo. ", and Lunel 4,108 in a
female weighing 4 kilo* 6. Their hatching requires a
longer or shorter time according to the temperature of
the water. In a temperature of 472° C. (40° Fahr.)
Reynolds found the period of incubation to vary be-
tween 60 and 70 days for most of his Charr ova, 75
days for some, and 90 days in a few instances; and
in a temperature of 123/4° C. (55° Fahr.) most of the
eggs were hatched in 41 days. From the eggs which
Malm impregnated by artificial means at Baldersnas in
Dalsland at the end of October, the fry commenced to
appear in the beginning of the following March, being
then about 17 mm. long. After living 19 days in an
aquarium in his room, they were 27 mm. long, and
had begun to develop the characteristic Parr markings.
At Ostanback, according to Bystrom0, the eggs taken
in September from the neighbouring spawning-places
were hatched in January, but the fry at first grew
slowly in the cold spring-water of the fish-ponds, and

" See Day, Fish . Gt. Bril, and Irel., vol. II, p. 117.

6 See Fatio, 1. c., p. 410.
c See NystrOm, 1. c., p. 15.

were only a little over an inch long in August. The
broods of Charr fry sent from Ostanback to the Royal
Museum show that at the end of the first year the
length is 97 mm., at the end of the second year 123
mm., at the end of the third year 137 — 216 mm., and
in Charr five years old 250 mm. The growth is de-
pendent, however, in the highest degree on the more
or less plentiful supply of food.

The Charr is not only one of the most beautiful
fresh- water fishes in Scandinavia, but also one of the
best — for good digestions the best of all. In summer,
however, it keeps out of the fisherman’s way, living,
as we have mentioned, in the depths of the lakes, and
being taken almost exclusively by angling, as a rule
the least destructive fishery. The Charr is in good
condition at this time of year, best towards the end
of summer, if cooked and eaten at once; but its flavour
rapidly deteriorates, especially in summer. The fisher-
man changes his tactics when the Charr begin to (ap-
proach their spawning-places. Where the bottom is
not too rough, the seine is used; else the place is
surrounded with nets. In either case an abundant
take may reward the fisherman; but the spawning-
season is never the best time of year, either in the
condition of the fish or with a view to the maintenance
of the fishery.

In Lake Wetter Charr are taken during summer on
long-lines baited with Smelt or bits of Gwyniad or Yen-
dace, and where the shores are steep, the line may be
set close to land in a semicircle, so that both ends touch
the shore. In other places the line must be taken
farther out into the lake, and set in at least 20 — 30
fathoms of water. The spawning fishery for Charr in
the shallows of Lake Wetter begins, according to \\ i-
degren, about the 10th of October and does not close
in some parts until the 20th of November. The annual
take, according to his computation, is about 3,000 kilo.
In the south of Lake Wetter distinctions Avere draAvn
about 1830, according to Sergeant-major Hall, between
three kinds of Charr: the Shore Charr ( Landroding ),
which attained a length of 9 dm. and a weight of
57i0 kilo., back green, belly flame-red, fins of the latter
colour Avith Avhite margin; the Deepwater Charr (Djirp-
rddinrj), which attained a length of only 6 dm. and a

Scandinavian Fishes.

107

848

SCANDINAVIAN FISHES.

weight of at most o2/5 kilo., body shorter, head smaller,
coloration lighter, than in the former variety; and the
Salmon Charr ( Laxroding ), of the same size as the Deep-
water Charr and also similar in other respects to the
latter, but with white belly. At the same time, accord-
ing to Dr. Marin, in the north part of Lake Wetter
the following kinds of Charr were named: the Bed ,
with red back, belly and sides of lighter red, yellow
fins, and red flesh; the Spotted , sides and back spotted
with white on a gray ground, belly whitish, flesh white,
whitish yellow, or gray; and the Gray , with grayish
green back and sides, belly and flesh as in the Spotted
Charr. All these names simply indicate the great va-
riableness of the Charr in different haunts, even within
the limits of a single lake. Especially worthy of re-
mark from a culinary point of view is the great dif-
ference in the colour of the flesh, which varies from
bright red to whitish. The pronounced red tint is due
to a diet consisting chiefly of crustaceans.

In Lakes Stor-Uman and Malgomaj (Lycksele Lapp-
mark) a distinction is drawn between Light and Black
Cliarr. There the Charr fishery is best, according to
Trybom, during the spawning-season and immediately
afterwards. Stationary nets are used, and they must
be set, at least towards the close of the fishing, under
the ice. After the breaking up of the ice, and until
the Charr retires to deep water, a productive fishery
is carried on with long-lines. In a few Lapland lakes
the bottom is smooth enough for the employment of
seines, but this is seldom the case at the spawning-
places. In the Lajsan lakes, towards the end of sum-
mer and in autumn, the take includes not only breed-
ing Charr, but also numerous fish that are “ gaW (bar-
ren, i. e. not ready to spawn the same year) and very fat.

In many places the Charr is caught with hand-
lines, and voracious feeder as it is, it takes any kind
of bright bait, if only this be kept moving. Day* 6 * 8
describes a sort of whiffing for Charr practised on Lake
Winandermere: “By far the most important and in-
teresting means of taking char there is by means of
the plumb-line. This line is made of strong cord, and

varies in length according to the number of baits which
are to be put on it; but it is usually between forty
and fifty yards long, and this is sufficient to carry five
baits. The baits usually used are artificial, pieces of
metal silvered on one side, copper, red, green, or brown
on the other, spinning from either the head or tail.
Minnows can be used in the same way, spoon baits too,
and both the blue Phantom and Garnet-quill minnows
have been tried successfully. Still the natives prefer
the metal baits, and the sizes used for ordinary trout
are of course the correct ones. Usually two such lines
as the above are worked by each boat, and the fisher-
man shows considerable skill in his manipulation of
them and rowing his craft along at a proper speed at
the same time — the latter is just sufficient to keep
the baits spinning and the tackle taut. The boatman
knows the ground char frequent, and the nature of the
bottom too, for should he come upon rocks and weeds
his tackle gets entangled, and a big smash must almost
inevitably result. This fishing usually commences about
the beginning of March, and at that time the fish are
got about thirty yards from the surface and in the
deepest parts of the lake. As the weather gets warmer
they gradually approach the top.” The Charr is also
taken at the surface of the water during summer with
the otter-lineb and the casting-rod.

“The Charr is very easy to breed”, says Max von
dem Borne0, “and best adapted of all the Salmon-fishes
for fattening. But it must without fail be supplied
with pure spring- water of a fairly uniform tempera-
ture, which should never exceed + 15 to 1 l1jf C. (59
to 6372° Fahr.). It is especially to be recommended
to the pisciculturists who have a plentiful diet of in-
sects to offer it, as well as to those who desire to keep
their fish in small ponds, and to fatten them for mar-
ket from their earliest youth on a diet consisting
mainly of meat and fish. It is very sociable and
tame, being perfectly at its ease among fishes of dif-
ferent genera and sizes, while the Trout, on the con-
trary, is always shy and especially intolerant of smaller
companions.”

“ L. c., p. Ill; Brit., Ir. Salmonidce, pp. 235 and 243.

6 A line with flies or spinning baits, natural or artificial, and constructed, for surface-fishing, partly on the same principle as the

otter-trawl for ground-fishing. An otter-board at the end of the line is arranged vertically floating at the surface, but obliquely to the di-
rection of the rowboat, thus leaping out from this and stretching the line, trolling the baits on the water.

c Handbuch der Fischzucht und Fischerei , p. 284.

SALM0NID7E.

849

THE SALMON (sw. laxen).

SALMO SALAR.

Plate XXXVII, figs. 3 and 4; plates XXXVIII— XL.

Scales small, but their number in a longitudinal row above the lateral line just in front of the perpendicular from
the beginning of the dorsal fin and extending along Vio °f ^ie length of the body at most 24a (as a rule 16 —
20b), and in a similar row above the lateral line behind the perpendicular from the beginning of the anal fin at
most 19° (as a rule 14 — 18). Sides of the body more or less distinctly spotted with black.

R. hr. (9)10

12(13)^; D.

(2)3-4

(7—8)9—12’

A.

2—4

7—9(10)’

p . y

(10—11)12 — 14 ’

Ser. squ. lat.e ca 120 — 140

1

(6)7— 8(9 ); C'

; Vert. 57 — 60 A

x 4- 1 + 17 + 1 + x;

S/jn. Trutta 1. Salmo proprie, Nilss., Skand. Fa., Fisk., p. 368.

Salmo salar, Sundev., v. Wr., Skand. Fisk., ed. I, tab. 58 et
59; Smitt, Vet.-Akad. Haridl., Bd. 21 (1885), No. 8,

p. 162.

Var. Salmo salar sens. str. 1. nobilis: Spinae branchiales lateri
anteriori arcus pritni paris affixse numeri sunt minimi 18 —
vulgo 19 — 21(22). Longitude ossis maxillaris maximam

partem

longitudinis corporis vel longitudinis capitis

1000 ° * 1 100 b F

ajquat. Distantia inter pinnain adiposam et pinnam caudalem
longitudine superat altitudinem pinnae analis. Altitude minima

caudee maximam partem

27

1000

longitudinis praeabdominis

baud aequat. Maculie nigrae sub linea laterali nullae vel paucae
solum adsunt.

Fig. 210. Two young Salmon, the upper figure a female 2 years old from the Hatchery of Hofsmolla on the R. Laga in Halland, the
lower a mature male, hooked in the R. Nissa off Halmstad in April, 1863. Natural size.

a Extremely seldom 25, in which case the number of scales on the tail may be employed as a character.

1 10 — 12 scales on an extent = 1/4 of the length of the head, as Nilsson expressed this character; 9 — 13, according to Lilljeborg.

c Extremely seldom 21, in which case the number of scales on the forepart of the body may be employed as a character.

(1 Extremely seldom 8 or 14.

e Counted above the lateral line; in the lateral line itself 115 — 130, according to Day.
f Sometimes 61, according to Nilsson.

850

SCANDINAVIAN FISHES.

Syn. Salmo. nobilis, Schonev, Ichthyol. Slesv. Holst., p. 64
(nomen praelinneanum nec modo systeinatico propositum);
Olafsen, Reise gm Island , vol. I, p. 65; Pall., Zoogr.
Ross. Asiat., tom. Ill, p. 342.

Salmulus , Willughby, Hist. Pise., p. 192, tab. N, 2, fig. 2;
Ray, Syn. Meth. Pise., p. 63; Salmo salmulus, Turt., Brit.
Fna, p. 104; Jen., Man. Brit. Vert., p. 426; Fr., Vet.-
Akad. Handl. 1837, p. 3, tab. I.

Salmo rostro ultra inferioretu maxillam ssepe prominente, Art.,
Ichthyol., Gen. Pise., p. 11 ; Syn., p. 22 (ubi Eriox ex
Alberto Magno et e Cuba citatur); Descr. Spec., p. 48;
Lin., Fn. Suec., ed. I, p. 115.

Salmo Salar , Lin., Syst. Nat., ed. X, tom. I, p. 308 ( -f S.
Eriox, p.p.); Nilss., Prodr. Ichthyol. Scand., p. 2; Yare.,
Brit. Fish., ed. I, vol. 2, p. 1 ; Suppl. vol. 2, p. 1 ;
Agass., Ilist. Nat. Poiss. d'eaa douce, tab. 1 et 2 ; Jard.,
Brit. Salmon., tab. 1, 2, 7, 8; Kb., Danin. Fish., vol. 2,
p. 540; Nilss., Ofvers. Vet.-Akad. Forh. 1848, p. 62; Lillj.,
ibid. 1849, p. 36; Nilss., Stand. Fn., Fist., p. 370; Hckl,
Kn., Siisswasserf. Oestr. Mon., p. 273; Storer, Mem. Amer.
Acad. Arts., Sc., vol. VI, pt. II, p. 320, tab. XXV, fig. 2;
Wgrn, Ofvcrs. Vet.-Akad. Forh. 1862, p. 541 , tab. IV, figg.
3 et 4, tab. XXII; Sieb., ( Trutta ), Siisswasserf. Mitteleur.,
p. 292; Mgrn, Finl. Fishfii., (disp. Helsingf.), p. 58;
Gthr (Salmo), Cat. Brit. Mus., Fish., vol. VI, p. 11;
Steind. (Trutta), Stzber. Akad. Wiss. Wien., LIV, I (1866),
p. 23; Coll. (Salmo), Forh. Vid. Selsk. Chrnia 1874, Til-
laegsh. p. 155; ibid. 1879, No. 1, p. 85; N. Mag. Nalur-
vid. Chrnia, I3d. 29, p. 106; Malm, Gbgs, Boh. Fn., p.
534; Feddersen, Naturh. Tidskr. Kbhvn, ser. 3, vol. XII,
p. 76; Mor., Hist. Nat. Poiss. Fr., tom. Ill, p. 525;
Smitt, Ofvers. Vet.-Akad. Forh. 1882, No 8, p. 32; Mob.,
Hcke, Fisch. Osts., p. 124; Reiit., Sunlm., Finl. Fistfn.,
tab. XVIII; Day, Brit. Salmon ., p. 51, tab. Ill et IV;

Lillj., So., Norg. Fist., vol. II, p. 511; Fatio, Fn. Vert.
Suisse; vol. V, p. 298.

Salmo hamatus, Cuv., Regne Anim., ed. II, tom. II, p. 303;

Cuv., Val., Hist. Nat. Poiss., tom. XXI, p. 212, tab. 615.
Salmo Salmo, Cuv., Val., 1. c. p. 169, tab. 614.

Fornue intermedia:

Salmo cauda bifurca; maculis solum nigris, sulco longitudinali
in ventre, Art., Gen. Pise., p. 12; Syn. Pise., p. 25.

Salmo lacustris, Lin., Syst. Nat., ed. X, tom. I, p. 309;
Agass., 1. c., tab. XIV et XV; Sieb. (Trutta), 1. c., p. 301;
Nilss., Stand. Fn., Fist., p. 404.

Salmo Schiefermulleri, Bl., Fisch. Deutschl., pt. Ill, p. 157,
tab. CIII; Retz., Fn. Suec., Lin., p. 348.

Salmo sebago, Gir., Proc. Acad. Nat. Sc. Pliilad. 1853, p. 380
-f- S. gloveri, ibid. 1854, p. 85.

Salmo ocla, Nilss., Stand. Fn., Fisk., p. 397.

Salmo salar, var. lacustris, Hardin, Ofvers. Vet.-Akad. Forh.
1861, p. 382 = Salmo Ilardinii, Gthr, Cat. Brit. Mus.,
Fish., vol. VI, p. 107.

Var. Salmo trutta 1. fario: Spinas branchiales lateri anteriori
arcus primi paris affixie numeri sunt maximi 17 (vulgo
14 — 16, rarius usque ad 18 unius lateris). Longitudo ossis

86 , ,37

maxillaris mimmam partem longitudims corporis 1.

1 1000 r 100

longitudinis capitis superat. Altitudo pinnaa analis distantiam inter
pinnam adiposam et pinnam caudalem mensura superat. Altitudo

27

minima caudas minimam partem longitudinis prasabdominis

F 100 ° P

superat. Color corporis maculas nigras etiam sub linea laterali
crebras priebet.

Syn. Trutta salmonata, Belon, Nat., Divers. Poiss., p. 274;
Trutta fluviatilis , Rondel., De pise, fluviat., p. 169. Salmo
mis. 2 (p.p.), 3, 5, Art., Gen. Pise., p. 12.

Fig. 211. Two young Trout, immature females, hooked in the R. Nissa off Halmstad in April, 1863. Natural size.

SALMON.

851

Salmo Eriox (p.p.), Lin., Syst. Nat., ed. X, tom. I, p. 308;
Turt., Brit. Fna, p. 103; Jen., Man. Brit. Vert. Anim.,
p. 422; Kr., Damn. Fifth., vol. II, p. 602; Nilss., Ofvers.
Vet.-Akad. Forh. 1848, p. 63; Lillj., ibid., 1849, pp. 36 —
37: Nilss., Shand. Fn., Fish., p. 395; Coll., Forh. Vid.
Selsk. Chinia 1874, Tillaegsh., p. 157; ibid. 1879, No. 1,
p. 85; Olss., (Jfvers. Vet.-Akad. Forli. 1876, No. 3, p. 132;
ibid. 1882, No. 10, p. 48; Fedders., 1. c., p. 77; Reut.,
Sundm., Fml. Fisk., tab. XVI et XVII.

Salmo Trutta, Lin., 1. c.; Bl., Fisch. Deutschl., pt. I, p. 143,
tab. XXI ( + N. Goedenii , pt. Ill, p. 155, tab. CII); Nilss.,
Prodr. Iclithyol. Scand., p. 5 ( + S. Truttula, ibid.) ; Agass.,
1. e., tab. VI— VIII; Kr., 1. c., p. 582; Nilss., Ofvers.,
1. c. ; Shand. Fn., Fish., p. 406; Widgn, Ofvers. Vet.-Akad.
Forh. 1862, p. 560, tab. IV, figg. 1 et 2; tab. V; tab. VI,
fig. 1; tab. VII; tab. VIII; ibid. 1864, p. 279, tab. VIII —
XIV; Nystrom, Ialctt. Jerntl., (disp. Ups. 1863), p. 7; Sieb.
(Trutta), 1. c., p. 314; Mgrx, 1. c., p. 61; Steind., 1. c.,
p. 22; Gthr (Salrno), 1. c., p. 22 (+ S. brachypoma , p. 87
+ S. gallivensis, p. 88 + S. mistops, p. 105 + S. vener-
nensis, p. 110 + S. polyosteus, p. Ill + . . .); Malm, 1. c.,
p. 538; Smitt, Ofvers., 1. c., p. 31; Mob., Hcke, Fisch.
Osts., p. 126; Lillj., Sc., Norg. Fish., vol. II, p. 565.

Salrno Fario, Lin., 1. c., p. 309 ( + S. carpio , ibid.); Bl.,
1. c., pt. I, p. 148, tab. XXII et XXIII; Nilss., Prodr.,
p. 6; Agass., 1. c., tab. Ill — V; Kr., 1. c., p. 625; Nilss.,
Shand. Fn., Fish., p. 415; Sieb. (Trutta), 1. c., p. 319;
Steind., 1. c., p. 24; Gthr (Salrno), 1. c., p. 59 ( + S.
orcaclensis, p. 91+ S. fero.v , p. 92 (ex Jard.) + N. sto-
machicus, p. 95 + S. nigripinnis, p. 96 + S. levenensis, p.
101 (ex Walker) + . . .); Canestrini (Trutta), Fna d'ltal.,
pt. Ill ( Pesci ), p. 24; Apostol., Peche en Grice, p. 33;
MSb., Hcke, 1. c., p. 127.

Salmo Illanca, Waktmann, Schr. Berl. Ges. Naturf. Fr. 1783,
p. 55.

Salmo alpinus, Bl., 1. c., pt. Ill, p. 158 (uec synon.),
tab. CIV.

Salmo cambricus, Donov., Brit. Fish., tab. XCI; Gthr, 1. c.,
p. 34.

Salmo spurius, Pall., Zoogr., I. c., p. 343 (= S. Eriox
Linn^ei) + S. hucho, p. 344 (nec. Lin.).

Salmo lemanus, Cuv., R. Anim., ed. 2, tom. II, p. 303 ( + S.
trutta + S. fario + S. punctatus + S. marmoratus, p. 304
+ Saimlet, p. 305).

Salmo ferox, Jard., N. Phil. Jotirn. Edinb., vol. XVIII, p.
55; Brit. Salmon., tab. IV; Nilss., Shand. Fn., Fish.,
p . 412.

Fario argenteus , Cuv., Val., ITist. Nat. Poiss., tom. XXI, p.
300, tab. 616 (+ F. lemanus, p. 300, tab. 617 + Salar
Ausonii, p. 319, tab. 618 + S. Bailloni, p. 342, tab.
619).

Genera Salar et Fario apud Hckl., Kn., Siisswasserf. Oestr.
Mon., p. 247 et cett.

Salmo microps , Hardin, 1. c., p. 383.

Trutta variubilis, Llnel, Hist. Nat. Poiss. du bassin du Leman,
p. 146, tab. XVI— XVIII.

Salmo lacustris, Fatio, Fne Vert. Suisse, vol. V, p. 323.

The Salmon, with its multitude of names, occurs
within the Scandinavian fauna in numerous varieties,
both of form and colour, the extremes being very easy
to distinguish by well-marked characters, but connected
by intermediate forms in a manner that compels us,
after a comparative study, to comprehend them under
one specific designation. The most clearly distinct
varieties have long been known by different names in
popular parlance, and are now recognised even in
scientific nomenclature, the first as the Trout (Sw.
Grdlax, Salmo trutta), the second as the Salmon (Sw.
Blanklax, Salmo salar). The Salmon is generally the
larger: it attains, according to Buckland", a length of
14 dm. and a weight of 313/4 kilo.6, and Fatio c men-
tions specimens as much as 16 dm. long; but even the
Trout (if the determination be correct) sometimes mea-
sures nearly 14 dm.d

The chief external distinction between these two
varieties consists in the more elongated form, the short-
er maxillaries, and the more scattered spots of the
Salmon. In 1848 a fisherman from Elfkarleby showed
Lilljeborg a practical method of telling a Trout from
a Salmon merely by taking it in the hand. If you
grasp a Salmon round the tail, you have no difficulty
in holding it fast; but a Trout held in this manner
easily slips out of the hand. This is due to the greater
contraction of the peduncle of the tail in the Salmon,
a character which also affords the safest expression of
the greater elongation of the body in this form. The
least depth of the tail in the Salmon is less than 28 %
(as a rule less than 25 %), in the Trout about 30 %,
of the preabdominal length (the distance between the
foremost points in the insertions of the pectoral and
ventral fins). But this character holds good only in
adult and typical specimens. The case is the same
with another character expressing the same difference
in the form of the body: the length of the dorsal
margin of the peduncle of the tail (behind the adipose
fin) is greater in the Salmon, less in the Trout, than

a Nat. Hist. Brit. Fish., p. 292.

h According to information received by the Swedish Fisheries Commission of 1881 — 83, the Salmon in Sweden attains a length of
at least 1 1 3 m. and a weight of at least 251/., kilo.
c Fne Vert. Suisse, vol. V, p. 305.

d Day, Brit., Ir. Salm ., p. 181. The above-mentioned Commission was informed that in Sweden the Trout attains a length of 9
dm. and a weight of 7 fy kilo.

852

SCANDINAVIAN FISHES.

the height of the anal tin. But in the young this cha-
racter loses its validity, as well as in very old male
Salmon during the spawning-season. The length of
the maxillaries is, as a rule, less in the Salmon, greater
in the Trout, than 8 % of that of the body or 36 %
of that of the head; but the character, which was ori-
ginally sexual, applies only to typical specimens. Sal-
mon, as mentioned above, in general have comparatively
larger scales than Trout. Gunther and Day have ex-
pressed this by the statement that, in an oblique trans-
verse row forward from the posterior extremity of the

A B

Fig. 212. Vomer of a Salmon (H) and a Trout ( B ), seen from
below (a) and from the left ( b ). Natural size. After v. Siebold.

adipose tin to the lateral line, the Salmon has If or
12 scales, the Trout 13 — 15. But even this character
holds good only in typical specimens0, and sometimes
the two sides of the body differ in this respect*.

Among the internal characters that may be used
to distinguish Trout from Salmon, we have above
mentioned the number of the gill-rakers, which in the
outer row on the front of the first branchial arch is,
as a rule, under 17 in the Trout, over 17 in the Sal-
mon, or, if it lie greater in the former, made up in
front and at the anterior upper corner of verrueose,
spiniferous protuberances. Another character, derived
from the shape and dentition of the vomer, has received

special attention of v. Siebold. In typical Salmon the
head of the vomer has a more or less distinct pent-
agonal or hexagonal shape, in Trout it is triangular.
In the former the vomerine teeth disappear earlier, and
in old Salmon are sometimes almost entirely wanting;
whereas even the oldest Trout commonly have several
teeth not only in a transverse row on the base of the
head of the vomer, but also in a double or single row
along the carina on its shaft.

But these internal characters are no more trust-
worthy than the external: the naturalist who examines
hundreds of specimens will find them too often merged
in each other.

Note. One of the best examples to show the relation of the
two varieties to each other, as regards the last-mentioned characters,
seems to be afforded by the attempts to introduce the typical Salmon
into the Lake of Geneva0. These experiments were commenced by
the hatching of Salmon eggs from the Rhine and from the hatchery
of Hiiningen; but the fry excluded from these ova escaped into
the Rhone, and none of them was ever regained and identified with
certainty. From 1857 to 1860, according to Chavannes, about 7,000
fry were placed in the feeders of the lake, and during 1863 about
4,600; and in 1882 a pisciculturist, Lugrin by name, is said to
have deposited nearly 100,000 Salmon fry in the Rhone near Geneva.
Hardly any results, according to both Lunel and Fatio, are known
to have been attained by these experiments: the ‘Salmon’ which have
been caught from time to time in the Lake of Geneva, have proved
on closer examination to be Trout. The latter author, however, is
still dubious with regard to at least three finds. In these specimens
Trout and Salmon characters were blended, and especially in two of
them the vomer was most nearly approximated to its Salmon type.
Though it is indeed possible that all the Salmon fry have been de-
stroyed, it seems more probable that during growth the fry have
passed to the Trout form, as this appears in its normal development
where the water offers to the fish no practicable means of commu-
nication with the sea.

The Salmon is so well known that a minute de-
scription would be here superfluous, especially as it
differs but slightly from the Charr, which we have
just described. We may instead refer the reader to
our figures and to the following comparative table of
the most important external characters, expressed in
averages which have been computed 1) in the first
two columns, from our measurements of Trout in the
For ell, Oring, and Salmon stages and of typical Sal-
mon at the same periods of growth, 2) in the third
column from all these measurements combined for each
character, and 3) in the fourth column from the cor-
responding measurements of Scandinavian Charr.

° To Salmo trutta fluviatilis (the Freshwater Trout) Day assigns in his last work (Brit., Ir. Salmonidce, p. 199) 12 — 15 scales
in the said row.

6 See, for example, specimens Nos. 284, 285 and 306 in Tabula metriea IV in Smitt's Riksmuseets Salmonider.
c See Lunel, 1. c., pp. 128 — 130, and Fatio, 1. c., pp. 308 — 313.

SALMON.

853

Length of the body expressed in millimetres -

Length of the head.. in % of the length of the body

„ „ „ „ reduced (from the hind extremity of the intermnxillaries to the hind mar-
gin of the preoperculum).. „ „ ,, „ „ „ ,, „

Breadth of the interorbital space „ „ ,, „ „ ,, „ „

Length of the snout „ „ „ ,, „ „ ,, „

„ „ „ maxillaries - „ „ „ „ „ „ „ „

„ „ ,1 lower jaw „ „ „ „ „ „ „ „

Longitudinal diameter of the eyes „ „ „ ,, „ ,, „ „

Vertical . . ,, n n n n n ,, ,, ,, n „

Length of the suboperculum „ „ „ „ „ „ „ „

Distance between the dorsal fin and the tip of the snout „ „ „ „ ,, „ „ „

Base of the dorsal fin „ „ „ „ „ „ „

Height ,, ,, » i, 11 H 11 11 n 11 11 11

Length of the pectoral fins „ „ „ „ „ „ „ „

Preabdominal length „ „ ,, ,, „ „ ,, „

Distance between the ventral fins and the tip of the snout ,, „ „ „ „ „ „ „

Length of the ventral fins „ „ „ ,, „ „ „ „

Postabdominal length... „ „ „ „ „ „ „ „

Base of the anal fin.. „ „ „ „ „ „ „ „

Height ,, ,, ?J ,, ,, ,, ,, ,, ,, ,, ,, ,,

Dorsal margin of the peduncle of the tail „ „ „ „ „ „ „ „

Ventral ,, ,, . . ■ • ^ >>

Least depth of the tail ,, „ „ „ „ „ „ „

Length of the middle caudal rays „ „ „ „ „ ,, „

i? ;) longest ,, ,, ,, ,, ,, „ ,, ,,

Breadth of the maxillaries in % of the length of the head reduced..

n 5? y> ii ii ii ii ii ii ii maxillaries

Number of gill-rakers in the outer row on the first branchial arch of the right side.

1 pff

55 55 55 5 5 55 55 55 55 55 55 55 55 55 55 AC-LU 55 --

Number of scales in a longitudinal row = Vio of the length of the body just above the lateral line and in front of the dorsal fin

55 55 5’ 55 55 55 55 55 55 55 55 55 55 55 5 5 55 55 55 55 55 5 5 abOVC the UUal fin

Trout.

True

Salmon.

Salmons.

Charr.

361

391

364

333

22.2

20.8

21.4

21.5

14.9

13.5

14.6

14.4

7.2

6.9

7.2

6.9

6.9

6.7

6.9

6.6

8.9

7.2

8.6

8.5

13.6

11.7

13.3

13.4

3.4

3.2

3.4

3.3

3.i

2.9

3.1

3.o

5.4

5.2

5.4

4.9

42.3

41.3

42.i

43.7

11.8

11.5

11.8

10.3

12.5

11.0

12.3

11.9

15.i

14.6

15.o

14.9

29.o

29.7

29.i

28.9

49.2

49.i

49.i

49.2

11.8

10.8

11.7

11.6

20.4

20.6

20.4

20.9

8.5

7.9

8.4

8.4

12.2

9.6

11.8

10.5

11.3

12.i

11.3

12.o

12.1

13.3

12.4

13.o

8.9

7.4

8.6

7.2

7.8

6.o

7.5

6.9

14.4

14.5

14.4

15.i

14.8

13.4

14.5

12.4

24.7

25.i

24.7

21.o

15

18

—

23

15

18

—

24

20

18

—

30

17

15

—

26

The first two columns of this table show 1) the
differences between typical Trout and Salmon, 2) the
developmental alterations of Salmons during the more
advanced stages of growth, the latter partly because
the Trout, as we have proved above, in most of their
characters represent the earlier stages of growth, partly
because the specimens measured were on an average
shorter than the typical Salmon included in the table.
The last two columns, on the other hand, show the
relation of the Salmons to the Charr; and although the
Charr measured were on an average shorter than the
Salmons, still we see that in the great majority of
cases the percentages of the former betoken a more
advanced development, in the direction indicated by
the ' changes of growth in the first columns of the
table. Where the relation is the contrary, and cannot

be explained by the difference of age, as in the case
of the distance between the dorsal fin and the tip of
the snout in proportion to the length of the body, or
the breadth of the maxillaries in proportion to their
length, another factor, the difference of sex, has as-
serted itself. In the males the dorsal tin is, as a rule,
situated further back, and the maxillaries are longer,
than in the females; and here the Charr represents the
male characters.

The Salmon, like other fishes, is subject to va-
rious malformations. These are especially prevalent
and have received most attention at hatcheries", where
the fry live under more or less unnatural conditions,
and where there is better opportunity for observation
than in nature. One of these deformities, that may
often be seen, is the so-called pug-nose (reduction of

° See, for example, Day, British and Irish Salmonidce, pi. XII.

854

SCANDINAVIAN FISHES.

the rostral region), which may be persistent in Salmon
normal in all other respects, and admits of quite a
considerable development (tig. 213).

The Salmon leads a life full of changes, and it
adapts itself to them both in form and colouring. The
form undergoes the same variations as in the Charr,
but is never so terete as it may sometimes be in the
latter. Most terete, most nearly fusiform, though always
laterally compressed, is the true Salmon in its sea-
dress, with broad, convex back and sometimes with
the greatest thickness of the body equal to two-thirds
of its greatest depth, which is then about one-fifth of
the length to the middle of the base of the caudal fin.
The difference from the Trout in the same dress is,
however, inconstant and sometimes absent; and in the

a

Fig. 2 13. Monstrous head (simous malformation) of a Salmo salar

a, from the left; b, from in front. T

spawning-dress they both assume a deeper and thinner
form of body. This is especially prominent in old
males; which during the spawning- season develop, be-
sides the singular jaw-hook, to which we shall return
below, a high and more or less sharp dorsal projection
in front, supported internally by an indurated connec-
tive tissue. This hump, however, in no instance attains
the same development in the Salmons as in Charr of
the Oncliorliynchus group". Where the true Salmon
lives confined in fresh water, as in Lake Wener, it
also acquires as a general rule a deeper, more com-
pressed form; and that the Kelts (spent Salmon) have
a thinner body, we have already remarked. During
the earlier stages the distinction between Salmon and

a See Smitt, EiJcsmuseets Salmonider , tail. Ill, fig. 28.

6 In Parrs.

c In the largest breeding males of the typical Salmon which

Trout is, as a rule, more evident in the above respect:
it is generally easy to distinguish by the form young
Salmon (blanklax-foreller) from young Trout ( foreller ),
the body of the former being shallower, with the
greatest depth less than one-fifth of the length. At
this stage, however, the difference is most marked in
the more or less forked shape of the caudal fin: in
young Salmon the length of the middle caudal rays is
less, in young Trout more, than half that of the long-
est rays in the fin. As appears from the table of
averages on the preceding page, this character is in-
deed persistent, in typical specimens of the two varie-
ties, even in later life; but it disappears in old Sal-
mon wearing the spawning-dress, and sterile Trout
have a caudal fin most like that of typical Salmon.

b

nernensis, taken by Mr. Datjlgrun in 1880. 1/2 of the natural size;

specimen probably measured 7 — 8 dm.

Among the changes of growth that most contri-
bute to modify the appearance of the Salmons, the
most striking are those which affect the form of the
snout. In young Salmon the snout is comparatively
short and blunt; but its length sometimes increases
with age from about 5 %h to about 13 %c of that of
the body, or from about 21 %h to 47 %c of that of
the head reduced. This prolongation mainly affects
the anterior, pramasal part, the nostrils remaining at
a distance from the anterior orbital margin which in
young specimens is less, in old only slightly greater,
than the diameter of the eyes. The elongation pro-
ceeds uniformly in the females and the middle-sized
males, where the snout acquires a more or less point-

ve have been enabled to examine.

SALMON.

855

ed, conical shape; but in old males this process is
accompanied by a depression in front, the intermaxil-
lary bones being parted from each other, and the me-
dian region of the pramasal part being hollowed under-
neath or even perforated, to receive the tip of the
hook simultaneously formed by an upward growth of
indurated connective tissue from the point of the lower
jaw. These old males are known in popular language
by the name of kroklaxar (Hook Salmon, the French
becard, Salmo hamatus, see above).

Not only have these males with their singular
snout — the function of which is not yet known" —
been regarded as a distinct species, but the same rank
has also been conferred, for their short snout and pe-
culiar coloration, on the two stages of growth follow-
ing next after the fry, the Parrs ( Salmo salmulus ) and
the Foreller or so-called Common Trout ( Salmo fario ).

Even in early youth the ground-colour of the
Salmons varies according to the colour and light of
their environments. v. Wright’s figure (Plate XL,
fig. 1) represents a young specimen some months old
as it appears during the assumption of the dress which
is more fully developed in the Parr stage.

The dress of the Parrs is described at length by
Fries (1. c.), from the coloration of the Salmon fry in
Norrkoping River: “The top of the head and the back
are olive-green, with dark, round, stellate spots and
patches along the back. These small, dark spots ex-
tend in front down to the lateral line; but from the
neighbourhood of the dorsal fin to the caudal they
terminate half-way between the dorsal edge and the
lateral line. The belly is white with a dash of yellow.
The sides are of a handsome light yellow shading
into red. Along the lateral line lie 8 or 9 small,
round orange spots and the same number* of large
and oval, bluish patches, set transversely and crossed
half-way up by the lateral line. These patches (the

so-called Parr markings) alternate with the orange
spots'’. Above each eye is an arch of 4 round, dark
spots, and further back, between these arches, are 3
similar spots, set in a triangle; but all these markings
may be more or less distinct, and they show some
variation. Very constant, on the other hand, are the
two round, black spots that adorn the gill-covers,
though their size and position vary. In most cases
one of them is situated at the centre of the operculum,
the other in front of the preoperculum, just behind
the eye. Sometimes traces of a third spot may be
detected at the very margin of the operculum. The
dorsal fin is of a light olive green, with one distinct
and one less prominent and irregular row of dark spots,
the former at the bottom, quite near the base and
parallel to the back, the latter along the middle of the
fin. The anterior corner shades into flame-yellow,
marked off by a dark gray band, which starts from
the tip of the second simple ray and runs in a straight
line to that of the fourth branched rayd. The adipose
fin wears the olive-green colour of the back, with yel-
lowish tip. The caudal fin is olive-green, tinged with
yellow, edged with flame-yellow above, below, and,
less distinctly, behind. The anal fin is of a dirty yel-
low, with the outer part of the anterior margin lighter,
and with an indistinct gray band obliquely across the
anterior corner. The pectoral fins are olive-green with
a darker band across the middle. The ventrals are
similar in colour and markings to the anal fin. The
pupil is rounded and very large in the living fish, buf
after some exposure to the atmosphere it assumes a
triangular shape. The flesh is white, without red
tinge.”

During the Forell stage the two varieties differ
more widely from each other, for the young Salmon
make their way to the sea as early as possible, and
when this instinct begins to awaken, they gradually

° Many conjectures have been made on this head. Some have supposed that the hook is a kind of spade or thrusting apparatus to
he used in working holes to receive the eggs. But, so far as is known, the female is at least more active than the male in this operation.
Others have thought it to be a hook with which the male holds the female fast while spawning. But the most trustworthy observers have
seen the spawning performed without its assistance. Others, again, have suggested that it is a protective organ, to prevent the males from
inflicting too severe injuries on their rivals in the breeding-season. But such means of protection seem far from natural. The least violent
explanation appears to be that the hook is an essentially pathological appearance, characteristic of age, but produced by the irritation caused
by blows on the snout, both during combats and in leaping over obstacles.

b Sometimes, however, ns many as 15.

c The ground-colour of the sides generally, grows lighter round the Parr markings, at least along their lower margin, thus rendering
them still more prominent.

d Cf. the analogous marking (the corner pure white and the stripe black) on the dorsal fin of Trout Parrs, according to Jardink,
British Salinonidce, pi. XI.

10S

Scandinavian Fishes.

856

SCANDINAVIAN FISHES.

adopt, their bright dress. The Parr markings and the
red spots slowly disappear, and the sides of the body
at the same time exchange their orange ground-colour
for a brighter and brighter silvery lustre, caused by
the depositing of a silvery pigment on the inner sur-
face of the scales and opercula. Now, and not until
now, do the scales begin to be quite distinct. Young
Salmon of this age are known in England as Smolts.
Great irregularity, however, may be observed in the
growth of these Parrs and the consequent alteration of
colour. At Stor month eld, among a brood excluded from
eggs impregnated at the end of December, 1861, and
hatched in April, 1862, were found on the 1st of April,
1868“, some specimens 54 mm. long and \l grammes
in weight, others 92 mm. long and 33/4 grin, in weight,
and others 165 mm. long and 37 1/4t grm. in weight.
Only the last were Smolts, and these would probably
have migrated to the sea the same season, the others
not until the following year or even the year after
that.

The alteration in the colouring of young Trout
after the Parr stage is generally not great, provided
that they belong to a race which is either unable or
has no inclination to enter salt water. They assume
a dress in which they are known as Foreller (Eng.
River Trout, Common Trout), and which they may
retain in small, land-locked waters almost unchanged
throughout their life. Under these conditions their
growth too is inconsiderable, v. Weight’s figure (Plate
XL, fig. 2) represents a common Scandinavian Forell.
The orange ground-colour of the sides and the red
spots on the lateral line — each surrounded here by a
more distinct, light ring — are persistent; and both
above and below the lateral line there appear similar
spots, scattered over the sides of the body6. At the
same time the number and extent of the dark (black)
spots on the back and head are generally increased,
these spots now occurring even down the sides, while
the Parr markings disappear. The dorsal fin also shows
an increased number of black spots, and in this fin
— but more commonly in the ventral and anal fins
alone — the anterior margin sometimes becomes red or
white, which colour is usually marked off sharply be-
hind, as in the Charrs, by a black streak. The young
Trout which are to repair to the sea or the great lakes

assume, like the Salmon Parrs, a lighter, more and
more silvery dress.

In salt water, as well as in the great lakes, the
main alteration in the colouring of the Salmons after
the Forell (Smolt) stage affects the black spots, which
pass from a round or irregularly angular shape to a
stellate form (X-spots). As a rule, however, though
by no means always, the two varieties may be distin-
guished even in the sea by their coloration, the Sea
Trout being usually more thickly spotted than the true
Salmon, and bearing numerous black spots even below
the lateral line and on the greater part of the body,
even behind. Such is the case with the Taimen from
Tornea figured by v. Weight (Plate XXXVIII, fig. 3).
Under certain conditions, however, especially in the
sterile individuals mentioned above and in the speci-
mens which should probably be explained as hybrids
(mongrels), this difference vanishes. Nor is there any
constant distinction in colouring between the marine
form known in England as the Sewin ( Salmo cambri-
cus ) and a true Salmon. How this distinction asserts
itself in Scandinavian waters, may be gathered from
a comparison between v. Weight’s figures of a Salmon
(Plate XXXVIII, fig. 1) and a Silver Salmon (Plate
XXXVIII, fig. 2).

The Salmon which have no access to the sea or
to any large, deep, and clear lake — whose Oring and
Lax stages are thus more immediate progressions from
the Forell stage — are remarkable for the exceeding
inconstancy of their colours, whence Lunel proposed
to unite them under a distinct specific name, Trutta
variabilis. In Scandinavia, as elsewhere, they occur in
brooks, rivers, and lakes. They are usually small, 1/2
— 1 lb. in weight, but frequently weigh 10 lbs., some-
times up to 18 lbs. Their coloration — lighter (redder)
or darker (sometimes nearly black), with denser or
more scattered red and black spots — is determined,
as in most fishes, primarily by the colour and light
of their surroundings, but also, in a high degree, by
their food. Of this as clear evidence as one could
desire was afforded by a visit in 1883 to Sir James
Maitland’s hatcheries at Howietown (near Stirling)..
Two of the ponds contained, among other fishes, a
variety of Trout from Loch Leven ( Salmo levenensis ).
In one pond, where they were fed on horseflesh, the

“ See The Field for the 25th of April, 1863.

b Red spots above the lateral line are not uncommon in young Salmon.

SALMON.

857

silvery colour prevailed on the sides of the body, while
in the other, where their diet consisted of crushed
scallops ( Pecten ), the ground-colour was orange, not
unlike that of an Oring from Bohuslan figured by v.
Wright (Plate XL, fig. 3). One of fhese Trout varie-
ties, the darkest and most thickly covered with black
spots, sometimes spotted with black even on the pale
ventral side, but often — where it has attained a suffi-
cient size — without red spots, has been named by
Jardine Salmo ferox. The same variety is commonly
the largest fresh-water Trout in Scandinavia, and the
males are furnished during the spawning-season with
a hook almost as large as in old males of the true
Salmon.

The Salmon in all its stages of growth is a pre-
datory fish of great voracity. Where it roves in the
sea — as along the greater part of the west coast of
Europe, in the White Sea, and in the Baltic — it lives
on all kinds of animals, principally crustaceans (shrimps),
Herrings (Baltic Herring) and Sand-Eels. To the last
it seems to be especially partial, and in quest of them
it resorts in large shoals to the sandy shallows off the
coasts of Scania and Pomerania. In these Avaters a
seine-fishery has been instituted, the seine being shot
in the sea, we may almost say, at hazard, and drawn
ashore frequently with a good haul of Salmon. This
fishery is carried on during spring, in March, April,
and May. Another fishery, based on the well-known
voracity of the Salmon, is pursued, as mentioned above,
with long-lines (Salmon-lines) in the south of the Bal-
tic by Swedish, German, and Danish fishermen". During
the roving excursions of the Salmon in the south of
the Baltic at the season specified above, drift-nets are
also used. As the spring advances, however, and the
North Baltic with the Gulf of Bothnia becomes open,
the marine Salmon begin their journey thither. They
are taken here and there on the way with gill-nets
and long-lines, at least up to the island-belt of Stock-
holm. Farther north they come to the shore-traps
(so-called stake-nets and mockor), to which have been
added in recent times the so-called Finnish stor-ryssjor,
where many of the Salmon making for the rivers meet
their fate. In the rivers the marine Salmon eats hardly
any food, at all events after it has been there some

a See Benecke, Fische , Fischerei und Fischzuclit in Ost-
Fisherier , l:sta haftet, p. 78, tab., fig. 18.

b Thierleben, Grosse Ausg., Abth. Ill, Bd. II, p. 227 ; De

time, though the eagerness with which it takes the
fly-fisher’s bait in the lower courses of the Scandina-
vian rivers, shows that it has not yet laid aside its
voracity. Different are the habits of the River Trout
and the large Salmon of the lakes. Hardly any other
fish can lie said to be more voracious or a more greedy
eater in proportion to its size. Breiim has described
the life of the River Trout/'. “In quickness and dex-
terity of movement,” he says, “hardly any river fish
surpasses the Trout. It is probably to be included
among the fishes that are nocturnal in their habits;
all observations indicate, at least, that the Trout does
not display its full activity until evening, and prefers
to perform its principal task, the procuring of food,
by night. In the daytime they mostly take shelter
under projecting stones in the bank or in any hollow
or cranny of the rocks to be found in the water which
they inhabit. But when everything around them is
perfectly still, they move about in the open even by
day, always with the head pointing up the stream.
Here they either remain apparently motionless at the
same spot for a quarter of an hour or more, though
their fins really move with sufficient rapidity to coun-
teract the current, or dart like arrows through the
water, following the main channel with surprising
adroitness, and in this manner finding their way along
shallow brooks, where you would think it impossible
for them to advance. When disturbed, they retreat,
if possible, to some hiding-place, for they are among
the most, timid and cautious of all fishes. Streams
with a swift current they descend by two different
devices, either by turning their heads up the river and
letting themselves slowly drift with the stream, or by
summoning up all their powers and shooting so rapidly
through the wafer as far to exceed the rate of the
current. While the Trout is at rest; it carefully sur-
veys its preserves, the water beside and before it and
the surface or the air over its head. If an insect,
whether great or small, approaches the post occupied
by the Trout, it keeps anxious watch until the insect
has come within reach, and then, with a powerful
blow or two of the caudal fin, rushes upon its victim,
or leaps out of the water to secure its prey. While
the Trout is young, it feeds mostly on insects, worms,

Westpreussen , p. 401, and Lundberg, Meddelanden rorande Sveriges
lodiga ryggradsdjurens lif (Sw. transl. 2nd ed.), p. 358.

858

SCANDINAVIAN FISHES.

leeches, mollusks, small fishes, and frogs. But as soon
as it has attained a weight of 1 — V/2 kilo., it rivals
in voracity all other predatory fishes of the same size,
being hardly surpassed by the Pike, and boldly attacks
any living thing which it believes it can overcome,
not excepting its own progeny, though the peaceful
Bleaks and Gwyniads are perhaps its most frequent
victims. Even in later life, however, its diet is com-
posed mainly of all insects that live in the water in a
larval or a perfect state, and small crustaceans. Its
partiality to the former is so great that it seems to
suffer from want of food, if other insectivorous fishes
— even those which it readily preys upon itself —
multiply too extensively in the same waters.” When
the lemmings migrate from the mountain tracts of
Lappmark, and endeavour, as they often do, to swim
the rivers in their path, they are devoured in numbers
by the large Trout ( Gralaxar ).

Such is the life led by the true fresh-water Sal-
mons from the extreme north of Europe to the south-
ernmost parts of Spain, in Algiers, Asia Minor, and
probably, to the north of the Hindu Kush. In those
southern regions they are reminiscences of the time
when their present abodes were connected with seas
cold enough for the Salmon to thrive and rove about
in their waters. Their occurrence in Africa is no so-
litary phenomenon; the range of the Sticklebacks shows
the same memories of primeval times. Since the Me-
diterranean has become too warm and perhaps too salt
for the Salmons, they have succeeded in adapting them-
selves to their environments in some scattered rivers
and lakes, and in there maintaining their existence at
the Forell stage. In Scandinavia these are the fishes
that ascend to the greatest, heights among the moun-
tains, as high as a, fish can well advance in the moun-
tain lakes and brooks. Clear and oxygenated waters,
freshened by rushing falls, are the favourite haunts of
the River Trout. On the plains and to sluggish, clayey
rivers it is a stranger. In Switzerland, according to
Patio", it ascends to a height of 2,630 in. above the
sea-level. In salt, water, on the other hand, the range
of the Salmon is terminated to the south by Cape

Finisterre, in about 43° N. lat., and it thus does not
enter the Mediterranean. On the west side of the
Atlantic its southward range extends to about 41° N.
lat., but there its northward extension is not so great.
It occurs, it is true, up to the middle of Labrador,
but is probably wanting in Greenland and on the west
coast, of North America. It has, however, been intro-
duced, after several unsuccessful attempts, both about
1860 and in more recent, years, into Australia* 6 and
New Zealand, where Trout, originally hatched from
ova of the English River Trout, are said to have
adopted the habits and dress of the migratory Sal-
mons1'. From these regions Day'6 received, among
other specimens, a male and a female, the former mea-
suring 825 mm., the latter 800 nnn., whose characters
he found most closely to resemble those of the English
Salmo ferox.

The Salmon spawns, like the Charr, in autumn
and winter. As a general rule it is, no doubt, true
that the Salmon is bound by its love of home to re-
turn from the sea to spawn in the watercourse and
the place where it. was itself born and bred. Herein
it is guided by instinct so unerring that in many lo-
calities the fishermen declare they can distinguish with-
out. fail between Salmon belonging to different rivers,
even if the mouths of two or more of these rivers lie
close besides each other. Marked fish have often been
retaken in the river where they had been marked and
set at liberty. But exceptions also occur; according
to I )ayc, for example, Salmon have been seen making
their way up the Thames, a river which had long
been deserted by the true Salmon. From the results of
the fishery in the rivers which fall into the Gulf of
Bothnia, it is also known that the run of the Salmon
in these rivers — more plentiful one year in one stream,
less plentiful in another — is greatly affected by diffe-
rent states of wind and weather, which would probably
exert, a less appreciable influence if the same Salmon
always repaired to the same river.

Early in spring, soon after the breaking up of
the ice, the Salmon begin to appear at the mouths of
the rivers which they are to ascend. The Salmon is

a Fne Suisse , vol. V, p. 373.

6 See Nicols, The Acclimatisation of the Salmonidce at the Antipodes, London 1882.
c Day, British Salmonidce, p. 145.
d L. c., p. 198.
e L. c., p. 66.

SALMON.

859

then in its best condition, though infested with a num-
ber of parasites, which it gets rid of in fresh water,
where they die. For a time it roves to and fro in
the brackish water; but when the spring floods come,
it commences the ascent in earnest. But not all the
Salmon arrive at the same time; the companies seem
generally to contain some 30 — 100 fish, and the ascent
may last the whole summer. Throughout the season
for the upward journey, the Salmon may be seen
thronging at the surface and leaping into the air both
in the sea off the mouths of the rivers and in the
lower part of their course; but higher up the stream
they muster in an angular formation, like that observed
by birds of passage, with one of the strongest fish at
their head, and the others tailing off gradually on each
wing. In this array they sometimes advance “with
such vigour,” says Gisler, “that the din thereof has
been heard on shore like a storm or subdued thunder,
the fish now and then swimming with the back half
above water or appearing like waves on the surface.”
In stormy weather or oppressive heat, hoAvever, they
keep nearer to the bottom. The females usually lead
the way, the smallest males bringing up the rear; and
when the fisherman takes one of the latter, he may
conclude that the main body of the shoal has passed
by. When obstacles bar their progress, they disperse;
but the ranks are again closed when by strength or
cunning the impediment has been surmounted. If they
come to a fall, they pass it by leaping, perhaps after
many unsuccessful attempts and only by springing
from one resting-place to another, found behind some
stone or jutting rock with slack water under its lee.
In this manner the Salmon leap to a height of 1 —
1 a/2 m. and a distance of 2 — 3 m., if necessary; and
from the brink of the fall they advance seemingly
none the worse for their exertions. But many of them
fail in the endeavour, and pay the penalty with their
lives. Natural obstacles are, however, less destructive
to the Salmon than the contrivances of man. In the
sea it is intercepted, as we have mentioned above, by
apparatus of ordinary design, nets hung on stakes
driven into the bottom, strandsdtt in Halland, stakandt
( mockor ) in Norrland (fig. 214), and kilnotar (mostly
in Norway), with a long arm straight out from the
shore and a bend (kil) at the outer end, or Finnish
storryssjor, in which the bend is replaced by a huge
ryssja (p. 33, fig. 7) and its short arm. In the lower
parts of the rivers too similar nets are used, ior in

\

Fig. 214. Plan of a stake-net, with the so-called inocka or jut a
(the angle at the outer end). After Lundberg.

the R. Laga. Salmon-pens ( laxgdrclar ) are also con-
structed, these consisting, according to law, of upright
stakes at a fixed distance from each other; within these
the larger Salmon are detained for a time also ap-
pointed by law and taken in seines, into which they
are driven with ‘beaters’, or enticed into so-called kar,
constructed in the same manner, where they may be
caught with greater ease. Baskets are also hung in
the falls, for the Salmon to drop into if they miss the
leap. The rivers a, re lined too with more expensive
engines, pcitor, minor , tinbyggnader, and vrakhus. The
first are built on the same principle as the Salmon-
pens, of upright stakes, but the shore arm and the
interior of the court itself are covered with nets, and

Fig. 215. Plan of a pata in the Ume Elf, built out from the right
bank. The arrows indicate the direction of the current.

After Lundberg.

within the court, the entrance of which is first closed
with a net, seines are hauled. The minor (fig. 216)

Fig. 216. Plan of a Salmon mina at Baggbole (Ume Elf). The
arrows indicate the direction of the current. After Lundberg.

860

SCANDINAVIAN FISHES.

are regular lock-weirs, with sluices above and below, the
former in two rows. The uppermost row (a) is merely
a grating, which continually lets the water through,
as well as the gates ( c ) at the lower end. The latter are
let down only when the watertight gates in the second
row at the upper end (b) are shut, the water now running
out, but the fish being left behind. In the tinbyggnader
large Salmon-traps ( tinor ) are set, which are lifted with
a hoist when they are to be examined". The vralclius
are, strictly speaking, large Jcar (see above, Salmon-
pens), constructed beside a race. So manifold are these

Fig. 217. A Salmon stair. After Day.

contrivances that Sweden, to her cost, can boast of
her wealth of devices for the taking of Salmon.

In order to counterbalance the destruction wrought
by these engines, and to extend the run of the Sal-
mon by enabling them to surmount falls otherwise
impassable, Salmon stairs are built. In these the force
of the water is diminished by transverse walls pro-
jecting alternately from the sides, and extending across
more than half the width of the staircase but not
reaching to the opposite side. The stairs may be erected
beside a fall in many different modes, according to

the nature of the locality; but care should be taken
to avoid placing the lower opening of the stair in the
same direction as the fall, and to turn it obliquely
across the stream or, still better, straight against the
current.

The River Trout too — even those which pass
their whole life in brooks and small streams — rove
to the spawning-place. “In a valley not far from my
native place,” says Brehm*, “there rise copious springs,
which together form a brook that has power enough
to drive a mill-wheel. This brook falls into the Roda
and clears the water of the latter, which is sometimes
rather thick. As long as can be remembered, Trout
have lived here, but only for an extent of at most
eight kilometres. Above and below this part of the
brook they do not occur, as a general rule; and only
during the spawning-season does it happen that they
abandon their true home and rove to rocky parts of
the Roda in quest of breeding-places, though they have
equally good spots in their usual haunts.” The spawn-
ing-dress of the River Trout sometimes includes a black
coating on the forepart of the body, such as that we
have above remarked in the Charrs.

The longer marine Salmon remain in the rivers,
the more tumid are their generative organs, and the
darker their dress. Our figures show the appearance
both of the true Salmon (Plate XXXVII, figs. 3 and 4)
and the Sea Trout (Plate XXXIX, figs. 1 and 2) at the
beginning of these alterations; but at the end of then1
wanderings, when the spawning-season commences, they
have assumed quite a different colour, of much darker
tone, as depicted by v. Wright in his figure (Plate
XXXIX, fig. 3) of a female Sea Trout ( Oring ) in fresh-
water dress. The belly of the males becomes entirely
red. In old males of the true Salmon the margin of
the caudal fin is also straightened, without a trace of
indentation, or even, like the margin of the anal fin,
convex. By a retrogression to the characters of youth,
they are besides approximated to the Sea Trout in the
characters relating to the least depth of the tail and
the height of the anal fin, otherwise the most constant
distinctions between the two varieties.

In September the spawning-season begins. The
old fish commence first. The young sometimes wait
even till the following March. And among the latter

a See Gisler, Vet.-Akad. Handl. 1752, tab. I.
b Thierleben, 1. c., p. 227.

SALMON.

861

the males sometimes attain a breeding state in the
Parr stage, but this is probably never the case with
the females. The accounts of the spawning differ some-
what in certain particulars; but in general terms we
may say that the eggs are deposited in a hole on a
sandy or gravelly, loose bottom and covered, after im-
pregnation, with sand or gravel. Each female is at-
tended by one or more large males and usually by
several smaller ones. When she has found a suitable
spot in shallow water — often so shallow that she can
only just keep her body submerged — she turns to right
and left and lashes with her tail — - according to others
she also roots with the snout — until a hole one or
more decimetres deep is formed. She then sheds a
portion of her roe and quits the place, the male at
once coming up to impregnate the eggs, unless he be
engaged for the moment in combat with some rival.
These conflicts are sanguinary enough, often one of the
antagonists leaves the other to die on the field, or at
least tears away great pieces of his flesh. But the
female continues to deposit her roe, portion after por-
tion, returning from her excursions to the spawning-
bed, and lashes up showers of sand, gravel or pebbles,
so long as she has a male to hasten to her side when
the eggs are laid. Young observed how a single fe-
male Salmon conducted nine males, which were all
taken, one after the other to her spawning-bed; and
when even the last male was caught, she returned
with a large Trout in her train. Among the smaller
Trout, several of which usually keep at a respectful
distance below the spawning-place, the greater number
are most probably on the look-out for the eggs that
are carried away by the current; but now and then a
male seizes his opportunity of joining in the spawning,
when the female Salmon’s partner is battling with a
rival. In this manner the spawning proceeds, with in-
cessant pauses, a fullgrown female taking 3 or 4 days,
according to some observers up to 8 or 10 days, to
empty her ovaries. We have mentioned above the con-
dition of the Salmon after the spawning: some die of
exhaustion, the greater number repair to some calm
haven for rest, afterwards prepared to rove or drift
with the current down the path along which they have
ascended. When they let themselves drift, the descent

is made backwards, tail first; and they may then be
speared with a leister, as they stand in a race before
a stone, resting the caudal fin against the latter.

The eggs are of an orange colour and vary in size
according to the dimensions of the mother fish; when
ripe, their diameter is about 5 — 7V2 mm. in the Salmon
and Sea Trout, 4 V3 — 6 mm. in mature River Trout.
Their number varies in the same manner. In England
it is customary to reckon 800 — 900 ova to each pound
in the living fish; but Day" computed the number of
eggs in a female Salmon weighing '20 lbs. at 27,850.
According to Norback* 6 a Brook Trout 1 lb. in weight
contains only about 150 — 200 eggs. The period of
incubation varies greatly, but as a rule depends on the
temperature of the water. According to Dayc it proved
in a constant temperature of 45° Fahr. (+ 7 ‘2° C.) to
last 90 days, in 43° Fahr. (+ 6'1° C.) 101 days, in
41° Fahr. (+ 5° C.) 97 days, and in 36° Fahr. (+ 2'2°
C.) 114 days. By keeping them all the time in alow
temperature, their hatching has been delayed until the
148th day after impregnation; but if frozen hard, they
generally die.

The new-hatched fry are about 15 — 18 mm. in
length, with a vitelline sac about half as long hanging
from the forepart of the body, and lie still at the
bottom, keeping the pectoral fins, however, in continual
motion to renew the water in the gills, or flounder to
and fro in the water, soon to sink again to their shelter
behind a stone or in some other hiding-place- In about
4 — 6 weeks, at a length of 25 or 26 mm., the }rolk is
absorbed, and the fry -grow more lively. They now
begin to eat, seeking their food chiefly in small crus-
taceans ( Entomostraca ), but not disdaining other small
portions of animal food that come in their way. At
the age of 2 months Salmon-fry are about 32 mm.
long, at the age of 4 months on an average 63 mm.,
and when 6 months old, on an average 95 mm. But
the growth is extremely variable, of which we have
above adduced evidence.

The Salmon has always been one of the most
highly esteemed fishes for table; and to the sportsman
the River Trout has given most amusement of all
fishes, even if his passion — the enjoyment of over-
coming a powerful resistance, of mastering a strong,

° British Salmonida. p. 78.

6 Handhole i Fiskevdrd och Fiskafvel, p. 284.
c 1. c., p. 82.

862

SCANDINAVIAN FISHES.

a stubborn, and a cunning lord of the waters — be
more tickled when a huge Salmon has gorged the fly
on his line, and been brought ashore after desperate
struggles to escape. In delicacy of flavour too, the
River Trout is preferred to the Salmon; but its smaller
size renders it of less economical importance.

The annual value of the Swedish Salmon fisheries
may be estimated at a minimum of between 600,000
and 700,000 crowns (<£66,000 — 77,000)“. In Norway
the corresponding figures are half as great, or even
more*. Young c estimated the value of the Salmon
fisheries for 1877 in England at £100,000, in Ireland
at £400,000, and in Scotland at £250,000. On the
Lower Rhine, including the whole of its course in
Holland, the average annual take for 1878 — 79 was
44,302 fishd of an average weight of 8'28 kilo.; and
the average weight per annum of the whole catch was
thus 364,320 kilo., each kilogramme of Salmon fetching
in the Rhine countries, according to Brehm6, between
3 and 9 reichsmarks. The value of the Salmon is in-
fluenced, however, in an essential degree by the depth
of red colour shown by the flesh, a tint which seems
most, though not entirely, to depend on the proportion
in the Salmon’s diet of crustaceans. The flesh is also
paler as a rule in the Sea Trout than in the true
Salmon. The spent fish (Kelts) are of but very little
alimentary value, and should never appear in the mar-
ket, from which Salmon in the spawning-dress should
also be excluded. By the legislation of most coun-
tries all Salmon fishing during the Spawning-season
is, as a rule, forbidden. The fry and the Parrs too
must be protected by law if the fisheries are to retain
their value.

The great importance everywhere possessed by
the Salmon fishery has induced efforts not only to
preserve, but also to extend its bounds, and a special
means to this end has been discovered in Salmon cul-
ture. Hardly any other European fish — the Carp
perhaps excepted — has lent itself more readily to cul-
tivation; and the Salmon was also the first European

fish to attract the attention of pisciculturists. In the
fifteenth century a monk, Dom Pinciion by name, is
said to have successfully hatched ova in a trough filled
with running water; but not until long afterwards were
more elaborate experiments instituted. In 1763/ S.
L. Jacobi, a Westphalian farmer from Hohenhausen,
published in a letter to the editor of the Hannover.
Magas. (No. 23) his observations on the artificial
breeding of Salmon. The English Government found
the question so important that they rewarded Jacobi
with a pension. About 1850 the French Government
established at Htlningen in Alsace, not far from Basel
and near the Rhine and Rhone Canal, an extensive
hatchery, which in 1871 passed into the hands of the
Germans. During the present century many persons
have gained a reputation by efforts in this direction.
First we may mention the Englishmen Shaw, Buck-
land, and Sir James Maitland, and the German Max
von dem Borne; but all Europeans were eclipsed by
the American Spencer Baird ( d . 1887), who prevailed
upon the Government and Congress of the United States
to make grants at that time unprecedented, for the
advancement of pisciculture. Doubts have indeed been
raised, whether the results, especially as regards Sal-
mon breeding, have really repaid this lavishing of
money and exertion. But America, with practical dis-
cernment, has not been disheartened by these more
or less well-meant warnings from foreign quarters^.
Salmon culture has also gained an undeniable victory
in the above-mentioned introduction of the genus Salmo
into Australia and New Zealand.

The cause of the success won by Salmon culture
and of its importance, is to be found simply in two cir-
cumstances. First the mass of ova can be more com-
pletely impregnated than in nature, the discovery having
been made that the fecundation can be accomplished
by the so-called dry method , by pouring the undiluted
milt on the eggs in a dry vessel, in which manner its
fertilising properties are not dispersed or weakened
by water previous to the impregnation. Second it is

Fischer.

a Underdanigt Betcinkande med Forslag till Ng Fiskeristadga , Stockholm 1883, p. 148.

b Average annual value 1877 — 81 386,000 kr. (£42,460), according to Norges officiella statistik, Ny Rmkke, Udgiv. 1884, C. No. 9.
c See D. Milne Home, Salmon and Salmon-Fisheries , Gt. Intern. Fish. Exhib. London 1883, p. 55.

d According to Miescher-Ruesoh, Statistische und biologisclie Beitrdge zur Kentniss vom Leben des Rheinlaches im Siisswasser, Intern.
Ausst. Berlin 1880, Cat. Schweiz, p. 157.
e Thierleben , 1. c., p. 200.

f Four years before the publication of Alderman Lund’s method of cultivating fishes that spawn in spring (Vet.-Akad. Hand!. 1767).
9 See Nordisk Aarskrift for Fiskeri 1883, p. 324, and Brown-Goode and Wilmot in D. Milne Home, 1. c., p. 27.

SMELTS.

863

easier in the receptacles of a hatchery and in well-
constructed ponds than in nature to protect the eggs
and the defenceless fry from their numerous enemies.
As soon as the voracity characteristic of the genus
begins to awaken in the young brood, it is not diffi-
cult to supply them with suitable food, either Ento-
mostraca, where these can be procured in sufficient
quantity — no doubt a rare occurrence — or lean beef
or horseflesh, mixed with hard-boiled eggs and chopped
fine. But important conditions of success are to have
a constant supply of pure running water at a proper
temperature, and to remove every diseased egg. The
latter end is most conveniently attained by admitting

the current of water at the bottom of the hatching-
vessel and allowing it to run over at the top, the dis-
eased eggs, which become white and rise to the sur-
face, being thus carried away with the overflow.

Here we have no space, however, for a full de-
scription of Salmon culture in its details. The reader
who desires more information on this head may con-
sult either the Handbuch der Fischzucht und Fischerei
by Max yon dem Borne, a work which we have often
quoted, or Sir James Maitland’s History of Ilowie-
town. In Swedish these questions are handled with
ability by 0. G. Norback in his Handledning i FisJce-
vdrd och Fiskafvel.

Genus OSMERUS".

Canine teeth on the vomer and tongue; two rows of small teeth in the anterior part of the lower jaw, one row
of larger teeth in its posterior part, and, on the palatine hones; small teeth of uniform size, set in one row, on
the inter maxillaries, the maxillaries, and the mesopterygoid hones. Length of the maxillaries as a rule 60 — 66
%b, of the lower jaw as a rule 85 — 100 %c, of that of the head reduced. Number of rays in the anal fin greater

than in the dorsal, hut at most 17, 12 — 14 of which are
of the dorsal. Pyloric appendages few (at most about
row along the sides of the body, only the anter

According to the above limitation of the genus,
based on the most generally adopted opinions, it con-
tains only three species, very nearly related to each
other, one from the basin of the Atlantic, the second
from the Arctic Ocean, and the third from the basin
of the Pacific. Its most prominent character consists
in the dentition of the mouth and palate. The vomer
is especially remarkable, its development having been
arrested so that it is made up of the anterior part
(the head) alone, and officiates merely as a point of
attachment for twod large and widely separated, canine
teeth. Another dental character, shared, however, by
the following genus, is the presence of inner palatine
teeth. On opening the mouth of a Smelt, we see a
deep and narrow, longitudinal groove at the middle
of the palate, furnished at the sharp edges with a row

branched; and the base of the anal fin longer than that
7) and short. Scales middle-sized, about 60 — 70 in a
ior (10 — 22) being pierced by the lateral line.

of small but keen teeth, which are set on the inner
margin of the broad mesopterygoid bone, the largest
bone on each side of the palatine arch. The pterygoid
bone proper is toothless, consisting simply of a narrow
osseous disk, which lies close to the outer margin of
the mesopterygoid bone and unites the palatine and
quadrate bones. All the palatine teeth are canines,
but their development as such differs considerably,
bein'? most advanced in the foremost and hindmost
teeth. The largest teeth in the whole mouth, however,
are as a rule the outermost in the transverse row
belonging to the head of the vomer. The teeth in the
lower jaw are also canines, the posterior larger and
set in one row, the anterior set in two rows, with the
smallest teeth in the outer row. The intermaxillary
teeth are about equal in size to the last, and show at

a From the Greek oo/.n]Qrig or ooyriQog, odorous.

6 ’59'4 — G6'7 %, according to onr measurements of specimens in the possession of the Royal Museum.

85:— 102-4 %.

Orlrhree, or two pairs — one in front of and within the other — or five, in which case the inner teeth are smaller than the outer.

Scandinavian Fishes.

109

864

SCANDINAVIAN FISHES.

least a trace of canine formation; but the maxillary
teeth are still smaller, set closer together, at least in
places, and rather blunt. The tongue is boat-shaped,
with fleshy lateral margins and bears at the tip one or
two large canines and on each side of the true hyoid
bone (os linguale), within the said margins, a row of
somewhat smaller, similar teeth or only one or two,
in the latter case large canines. The copular part of
the hyoid bone is armed with smaller teeth, of about
the same size as the intermaxillary teeth, the largest
in a row along each of the edges, the others (the inner
ones) irregularly distributed in two rows. The pharyn-
geal teeth are about equal in size to those on the co-
pular part. Among the upper pharyngeals of the Scan-
dinavian Smelt only the two hind pairs are furnished
with teeth, the hindmost with several (3 — 4) rows,
the penultimate with one row. On the elongated lower
pharyngeals the teeth are set in two rows. These
numerous teeth of different form show that the Smelt,
in proportion to its size, is a voracious fish-of-prey.

The intestinal canal is short. The oesophagus
with its longitudinal inner folds passes uniformly into
the thick-walled stomach, which resembles a caacum,
and the bottom of which sometimes does not even
extend to a line with the tips of the pectoral tins, but
sometimes reaches to the anterior extremity of the
pelvic bones. On its under surface and in about a
line with the middle of the pectoral fins, the pyloric
part originates, running forwards at an acute angle,
and extending almost to the diaphragm, where it bends
abruptly back at the transition to the intestine, which
is here furnished with a few" comparatively short, but
thick pyloric appendages. The intestine runs straight
back to the vent. The liver is short and lies almost
entirely to the left of the oesophagus and stomach; it
extends back to about a line with the pyloric angle.
On the right, between the oesophagus and the small
lobe which the liver sends out on this side, lie the
gall-bladder and, a little further back, the spleen, which
in a healthy state is very dark, nearly black, and
ellipsoidal, more or less flattened* 6. In the structure
of the generative organs we have above remarked a
peculiarity characteristic of the Smelts: behind the

ovaries or, when they are fully ripe (in the spawning-

season), beside their posterior part, the mesoarium joins
its ventral margin, which in the preceding genus hangs
free in the abdominal cavity, to the lateral wall of the
said cavity, thus forming an oviduct open in front
(fig. 208, p. 828). The left ovary is the larger and
lies more to the front. In the spawning-season it
occupies almost the whole length of the abdominal
cavity, all the way from the diaphragm. The right
ovary lies in the posterior part of the abdominal ca-
vity, and extends forward only a little beyond the in-
sertion of the ventral fins. The testes occupy the same
position with relation to each other, and the left testis
is also the larger; but the difference in their size is
not so great as in the case of the ovaries. The air-
bladder is simple, long, and united throughout its length
to the dorsal side of the abdominal cavity, but does
not extend forward to the diaphragm or back quite to
the end of the cavity. From the lower part of its
anterior extremity the pneumatic duct runs to the
oesophagus.

The skull is weakly ossified, more elongated and
shallower than in the Salmons, and without the cur-
vature characteristic of the latter in the sphenoid re-
gion. Nor does the parasphenoid bone extend so far
back as in the preceding genus: its hind extremity
lies outside (under) the front portion of the basilar
part of the occipital bone, which part behind this point
is tumid on each side, whereas the Salmons here have
only a longitudinal, terete carina. This swelling, the
anterior part of which belongs to the temporal region,
includes the sacculus of the labyrinth, with the flat,
but long, heart-shaped otolith, which in a Smelt mea-
suring 177 mm. is 5 mm. long. The posterior oculo-
muscular canal extends about as far back as the para-
sphenoid bone, and thus belongs to the sphenoid and
temporal regions. The basisphenoid bone is wanting,
and the wall between the orbits is reduced to a car-
tilaginous ridge, ascending in front and belonging to
the hind part of the ethmoidal cartilage. The maxil-
laries with their os supplement are {jugate), the inter-
maxillaries, the palatines, the vomer, and the pterygoid
bones, are arranged as in the Salmons; but the reduc-
tion of the vomer and the dentition of the mesopte-
rygoid bones we have already noticed. The bran-

0 According to Keoyeb 3, to Day sometimes as many as 7 ; we have most often found 5.

6 In a male Smelt 133 mm. long the spleen was G mm. long during the spawning-season, in two females measuring 139 and 177
mm. the spleen was respectively 5 and 7'8 mm. in length.

SMELTS.

865

chiostegal apparatus differs from that of the Salmons in '
its fewer rays, 7 or at most 8, the anterior needle-
shaped, the last three broad and ensiform. The oper-
cular apparatus is distinguished from that of the Sal-
mons principally by the fact that the preoperculum is
attached to the upper part of the hyomandibular bone
(which is here elongated in the form of a, backward
process) only at the very top, in common with the ar-
ticular knob of the operculum. Below the said process

asq p psq

C

Fig. 218. A, Scale from the lateral line of Osmerus eperlanus with
imperfectly closed duct, magn. 8 diam. For the further elucidation
of its structure are added a scale ( B ) from the lateral line of a
Salmo trutta (9 338 mm. long, from Norrkoping), also magnified
8 diameters, and a section (G) of 21/, scales (magn. 10 diam.) from
the lateral line of a large Salmon, t , duct of the lateral line; asq,
anterior part of the scale; psq, posterior part thereof; p, pore in the
scale (answering to the deep incision in the scale from the lateral
line of the Smelt), through which the nerve of the lateral line passes
from one scale to the next, after sending out a ramification with
a sensory organ into the duct on the outer surface of the posterior
part of the scale.

is a large opening between the preoperculum and the
hyomandibular bone, which in front, on the other hand,
expands into a disk, forming a stronger support for
the palatine arch, and is itself strengthened b}T a down-
ward ridge at the middle of its inner surface. The
spinal column is similar to that of the Salmons; but
the haemal arches are not completely closed until they
reach the beginning of the caudal part. The series of
bones which in the Salmons forms the postclavicular

apparatus, is here represented by a small, round and
thin, disk-shaped bone at the hind inferior extremity
of the clavicle, with its upper part attached inside the
lower angle of the posteriorly expanded disk of the
latter. The pelvic bones are weak and resemble those
of the Salmons.

The scales of the Smelts are thin, on the sides of
the body elliptical, with longitudinal axis set crosswise
(up and down), on the belly more rounded or of broad
oval shape. In texture they resemble the scales of the
Salmons, without radiating grooves, but with more or
less numerous and dense concentric stria?. The indi-
stinct nucleus is eccentric and lies in the anterior (in-
serted) part of the scale. The lateral line is distinct
only on the anterior part of the sides. Its scales are
deeply cloven behind, where the canal of the lateral
line has its course, and the duct belonging to each
scale, and surrounding the canal, in front of this cleft,
and on the outside of the scale, is open or at least
imperfectly closed — as in the earlier developmental
stages of more complete lateral lines - — the margins of
the primitive canaliculate duct not coalescing, even
where one of them overlaps the other (fig. 218, A).
The scales of the Smelt are further distinguished by
the absence of the layer of silvery pigment which else-
where lines the scales where this colour appears. The
silvery lustre is indeed present in a longitudinal band
along the sides of the Smelt; but on removing the
scales we distinctly see that the coat of pigment lies
in the skin under the scales.

The relation to the Salmons has already been ex-
pressed in the above generic characters. But this re-
lation, as paleontology seems to indicate, is probably
one of development as well, Osmerus standing nearer
to the common ancestors of the genera. This appears
most clearly on a comparison of the average figures
expressing the corresponding relations of form in the
two genera.

From the above table of averages (p. 853) we
easily see that the position of the dorsal tin, expressed
in the distance from the tip of the snout to the be-
ginning of the base of this tin, becomes more forward
with age — and the averages for the several ages show®
that this alteration of growth is greatest during the
earlier and earliest stages. The said distance decreases
both in proportion to the length of the body and to

Cf. Smitt, Riksmuseets Salmonider, pp. 11 and 25.

866

SCANDINAVIAN FISHES.

the distance between the ventral tins and the tip of
the snout. Accordingly too Osmerus , as a representa-
tive of one of the earlier stages of the family, has
greater averages in these respects than the Salmons.
This is also the case with the length of the base of
the anal tin'*. A general rule for the whole life of the
Salmonoids is that the length of the maxillaries is re-
duced with age in proportion to that of the body6,
and the same rule applies to the length of the sub-
operculum in proportion to that of the head or of the
head reduced c, with a slight exception in the last case.
The breadth of the interorbital space, on the contrary,
increases with age in proportion to the length of the

head or of the head reduced d. The length of the pe-
duncle of the tail at the ventral margin e, and the least
depth of the tail7, in proportion to the length of the
head or of the head reduced, undergo alterations of
growth which at the modification in the life of the
Salmons — from fresh water to the sea — change di-
rection: their averages increase as a rule in the earlier
and earliest stages, but subsequently decrease, after
attaining their maximum at the end of the life in
fresh water. Here too Osmerus represents the earliest
stages.

Arithmetically expressed, this developmental rela-
tion appears as follows:

Average

Distance between the dorsal fin and the tip of the snout in % of the length of the body :

„ „ „ „ ,, „ „ ,, „ „ „ „ „ „ „ distance between the ventral fins and the tip of the snout.

Base of the anal fin in % of the length of the body

„ „ „ ,, „ „ „ ,, „ „ „ „ head reduced - __

Length of the maxillaries in % of the length of the body

„ „ „ suboperculum „ „ „ „ „ •„ head — -

,, ,, „ „ „ „ „ „ „ „ head reduced

Breadth of the interorbital space in % of the length of the head

„ „ „ „ „ „ „ „ „ „ „ „ head reduced

Length of the peduncle of the tail at the ventral margin in % of the length of the body

Least depth of the tail in % of the length of the head _

„ „ ,, „ ,, „ „ „ ,, „ „ „ head reduced

Osmerus.

Snlmo.

47. s

42.i

96.5

86.i

11.7

8.4

79.2

55.b

9.3

8.8

28.5

25.4

41.9

37.7

25.i

31.8

37.7

47.2

8.4

12.5

23.2

39.i

34.8

58.o

Iu other points, again, it appears that the Smelts
and the Salmons have each taken their own path
of evolution, and that the Smelts in several respects
— in some like the Charrs — occupy the higher rank
in the scale of development. On continuing our search

for the most different averages in the Smelts and Sal-
mons, and on comparing the signification thereof with
the alterations of growth to which the Salmons are
subject, we find the following relations most deserving
of notice:

a Cf. Smitt, 1. c., p. 14.

b „ ; v „ P- 10-

c „ „ „ „ pp. 18 and 22.

d „ ,, „ ,, PP- It and 20.

1 ) , 1 , • , 1 1 p • 1 5 .

f Cf. above, p. 833, the lower table, where these proportions may easily be calculated from the averages there given, and Smitt,
Riksmus. Salmonider , pp. 19 and 23.

SMELTS.

867

«■ S

11 11 11

11 11 11

Base of the dorsal fin

Length of the pectoral fins

Postabdominal length

Height of the anal fin

Least depth of the tail

Length of the middle caudal rays.

Length of the inaxillaries

Breadth „ „ ,,

No immediate descent of the Salmons from the
Smelts can thus be assumed.

On examining the relations between the three
forms hitherto distinguished in the genus of the Smelts,
we find that the Atlantic Smelt ( Osmerus eperlanus )
distinctly appears in some respects to have adopted its
own course of development, and that in these respects
it has remained nearer to the presumable origin of the
genus. But in most points the development seems to
have been common to the three forms, and the Pacific
Smelt ( Osm . clentex) is apparently the nearest repre-
sentative of the original type in the modern fauna.
We are besides struck by the analogy in the relations
of development to those between the three above-men-
tioned Charr forms, Salmo alpinus , S. stagnalis, and
S. salvelinus, for we find within the genus of the Smelts
too a defined natural series of three forms,

Osmerus clentex — 0. spirinchus — 0. eperlanus ,
as the most general expression of the course of evolu-
tion. Several deviations, however, meet us, which may
here too be explained as manifestations of the influence
of sex on the development, the most Arctic form —

Osmerus.

Salmo.

Changes of growth in
the true Salmons.

Umbla

i of the body °

14.8

15.2

juv. > sen.

14.4

b

8.0

12.i

„ > „

—

11 11 11

14.5

16.3

„ > „

14.9

(1

22.8

19.9

ii < ii

20.9

11 11 11 6

7.7

12.3

?? ii

—

„ „ „ /

5.i

8.8

ii ^ ii

—

„ „ „ 9

6.4

7.9

„ > „

6.9

head reduced *

62.8

58.i

ii "" ii

59.4

11 11 *

11.3

15.4

ii ??

12.4

maxillaries i ...

18.o

26.7

” > ”

21.o

Osmerus

sp irinchus k, known

from Alaska, Siberia,

the White Sea — being differentiated in most respects
from the other two forms by the predominant in-
fluence of the female characters, while Osmerus eper-
lanus, generally speaking, bears the stamp of the male
characters.

In order to find an expression for the general
alterations of growth within the genus, we have dis-
tributed our specimens among four classes according
to age, three of these classes fixed by comparatively
wide gaps in the decrease or increase of the percen-
tages during growth, the fourth (the oldest) marked by
a reversion to the earliest characters, best explained,
perhaps, as the result of a growth exceeding the most
natural limits of the forms. In certain parts of Swe-
den a distinction is made in popular language' between
nors (the smaller Smelts) and slom (the larger ones),
and at a length of 200 mm. there intervenes in the
Scandinavian Smelt one of the said gaps, for example
in the decrease of the percentage for the relative di-
mensions of the eyes. At a length of about 150 mm.
we observe a similar gap, and at a length of about

° Cf. Smitt, Riksm. Salm., p. 9.

6

1. c.,

P-

12.

c

1. c.,

P-

12.

d

]■ c.,

P-

14,

e

1. 0.,

P-

14.

f

1. c.,

P-

15.

y

1. c.,

P-

16.

h

1. c.

P-

21.

i

1. c.

P-

21.

j

1. e.,

P-

23.

k The name of spirinchus was applied by Lilljeborg (Vet.-Akad. Handl. 1850, p. 304) to the White Sea Smelt, and although Pallas,
who first coined the name, by no means restricted it to that form, we have refrained from giving the said form a new name, as spirinchus
in its old signification cannot be employed for the future as a specific name. In our first classification of the Salmonoids in the Koyal
Museum (Ofvers. Vet.-Akad. Furh. 1882, No. 8, p. 34 and Gt. Intern. Fish. Exhib. Loud. 1883, Swed. Cat., p. 184), we called this species
Osmerus dvinensis.

1 See Artedi, Descr. Spec. Pise., p. 47 and Nilsson, Skand. Fna , Fisk., p. 433.

868

SCANDINAVIAN FISHES.

260 — 270 mm. the above-mentioned retrogression be-
gins to assert itself in the changes of growth. The
four age-classes may therefore be named: nors, stor-
nors, slom, and storslom. Only in the third of these
classes have we been able to examine fully comparable
specimens of all the forms, in order to obtain an ex-
pression for the sexual differences within the genus.

One of the most distinct examples of the general
tendency shown by the development of form within
the genus — among the many that might be adduced” —
we find in the averages for the alterations caused by
growth in the relative length of the suboperculum.
The said length, expressed in percent of that of the
head, and reduced to an average for the whole genus
Osmerus, is 28'5. The several averages, as expressions
of the form-series and the changes of growth, are the
following:

Average in

Osmerus

dentex.

Osmerus

spirinchus.

Osmerus

eperlanus.

29.8

28.1

27.7

At the age of

Nors

—

—

30.7

Stornors ...

29.8

(27.9)

27.6

Slom cP

29.21 .

27.7)

27.7 1 ,

29.8

28.5

>26.6

„ 9

80. 4 j

29.3 )

20.6 |

Storslom _

—

27.9

25.6

Here the averages decrease with increasing age,
and the form-series also shows uniformly sinking aver-
ages. No constant sexual difference can be discovered,
for the male averages in 0. dentex and 0. spirinchus
are less, in 0. eperlanus greater, than the female.

The case is different, when a constant sexual cha-
racter exercises its influence on the development. This
appears, for example, in the relative dimensions of the
eyes. The vertical diameter of the eyes, calculated on
an average for the whole genus Osmerus, is 41‘5 % of
the length of the maxillaries. The several averages
expressing the difference between the forms and the
changes of growth, are the following:

Average in

Osmerus

dentex.

Osmerus

spirinchus.

Osmerus

eperlanus.

46.1

37.5

40.8

At the age of

Nors

48.4

Stornors

48.4

(36.8)

41.6

Slom cf1

45.4] ,

38.0)

39.7)

4».o

138.0

•38.2

„ 9 -

44.7 1

37. o )

36.8 J

Storslom

—

36.3

33.9

a Cf. Smitt, Rilcsmuseets Scilmonider, pp. 168 — 189.

Here too the changes of growth show sinking aver-
ages; but in the form-series Osmerus spirinchus occu-
pies the most advanced rank, as the representative of
a fairly distinct female character. Should tve desire
to see how the relation appears when this form breaks
the series by too high an average, we have a clear
example in the relative length of the lower jaw. This
length, expressed in percent of the length of the head
reduced, and calculated on an average for the whole
genus Osmerus, is 91 ’9. The averages for the form-
series and for the several ages are the following:

Average in

Osmerus

dentex.

Osmerus
spirinchus .

Osmerus

eperlanus.

89.7

96.1

90. o

At the age of

Nors

—

—

86.4

Stornors

89.3

(93. i-)

90.i

Slom <f

89.8)

95.2)

91.8)

89.8

95.7

92.4

„ 9

89.8 1

96.3 (

92.7 1

Storslom

—

98.8

96.7

Here we observe a still more prominent sexual
difference both in Osmerus spirinchus and 0. eperlanus
— and the former together with the females has the
highest averages — but Osmerus dentex , the least dif-
ferentiated form, shows no sign of a sexual character
in this respect.

Again, should we wish to see how specific cha-
racters manifest themselves during the course of de-
velopment, we can hardly find a better example than
among the Smelts. The only characters not infringed
by exceptions, i. e. the only constant specific charac-
ters — so far as we can gather from our specimens —
are the following:

A: 19 — 20 gill-rakers on the lower part of
the first branchial arch, and at most 29
on the whole front surface thereof.

Height (longest ray) of the anal fin at

most 7 ’6 % of the length of the body Osmerus dentex.

B : 23 — 25 gill-rakers on the lower part of
the first branchial arch, and at least
33 on the whole front surface thereof.
a: Height of the anal fin at most 7'6

°/o of the length of the body Osmerus spirinchus.

h: Height of the anal fin at least 8'4

% of the length of the body Osmerus eperlanus.

The manner in which the height of the anal fin
passes through its changes of growth, and affords one

SMELT.

869

of the external sexual characters, may be expressed in
the following table of averages:

Average height of the anal tin in %
of the length of the body, in

Osmerus
dent ex.

Osmerus

spirinchus.

Osmerus

eperlanus.

6.9

7.0

9.2

At the age of

Nors

9.6

Stornors ...

7.1

(7.i)

8.6

Slom cf -

6.81

6.9 |

8.9 I r

> 6.8

, > 7.o

■ 9.2

„ 9

6.9 J

7.2 1

9.3 |

Storslom —

—

6.9

9.3

The average in this relation for the whole genus
Osmerus is 7 '7, so that in the first two forms the
average falls short of this amount — both of them
showing almost the same average — but exceeds it in

the last form. In the first two the percentages sink
uniformly with increasing age. This is also the case
during youth in Osmerus eperlanus; but after the stor-
nors stage its course of development is reversed — the
female characters gain the ascendency in the basin of
the Atlantic. Here, as in several other respects, it
appears that the course of development was originally
common, but has diverged in connexion with a geogra-
phical separation, which in the Smelts, as well as in
the Salmons, probably dates from the Tertiary Period,
from the elevation of the Rocky Mountains in North
America".

The genus Osmerus was established by ArtedP,
who included within its limits, however, the modern
Saurns, a Scopeloid genus.

THE SMELT (sw. norsen).
OSMERUS EPERLANUS c.
Plate XLI, fig. 1.

Number of gill- rakers on the front of the first branchial arch 33 — 37, 23 — 26 of them being situated on the
lower part of the arch. Height of the anal fin ( length of its longest rag) more than 8 % of the length of the
bodg or than 36 % of the length of the head. Length of the middle caudal rags less than s/i of that of the

longest anal rag.

R. hr. 7(8);

D.

(2)3
8 — 9’

P 1 . y -!■

12 — 14’ ‘ 10 — 11(12) ’ ' 7 ’

(J. x + 1 + 17 + 1 + x; L. lat. (58)62 — 64(68); L. tr. (supra
pinn. ventr.) 13; Vert. 60 — 62.

Syn. Epelan de mer et Epelan de riviere , Belon., Nat., Div. Poiss.

p. 282 et 284; Eperlanus , Rondel., Pise. Fluv., p. 196;
Spirinchus et Stincus, Gesn., Paralip. p. 29; Spirinchus ,
Schonev., Iclithyol. Slesv. Hols., p. 70; Osmerus rndiis
pinnae ani septendecim, Art., Iclithyol., Syn. Pise., p. 21,
Descr. Spec. Pise., p. 45; Lin., Fna Suec., ed. I, p. 118.
Salmo Eperlanus, Lin., Syst. Nat., ed. X, tom. I, p. 310;
Fna Suec., ed. II, p. 124; Bl., Fiscli. Deutschl., part. I,
p. 179, tab. XXVIII, fig. 2(4- var. S. eperlano- mar inns,
p. 182, tab. cit., fig. 1); Retz., Fna Suec. Lin., p. 348;
Lacep. (Osmerus), Hist. Nat. Poiss., vol. V, p. 231; Pall.
(Salmo), Zoogp. Ross. Asiat., tom. Ill, p. 386; Mitch.,
Trans. Lit., Phil. Soc. N. Y., vol. I, p. 435; Nilss. (Osme-
rus), Prodr. Iclithyol. Scand., p. 12; Ekstr., Vet.-Akad.
Handl. 1834, p. 7; Yarr., Hist. Brit. Fish., ed. I, vol. II,
p. 75; Kr., Damn. Fisk., vol. Ill, pp. 1 et 1215; Cuv.,
Val., Hist. Nat. Poiss., vol. XXI, p. 371, tab. 620; Lloyd,
Scandin. Adventures, vol. I, p. 121; Nilss., Skand. Fna,
Fisk., p. 433; Widegr., Fiskfn, Fisker. Norrb. L., Landtbr.

Akad. Hand!. 1861, p. 9 (sep.); Sieb., Siisswasserf. Mit-
teleur., p. 271; Mgrn, Finl. Fiskfna (disp. Helsingf. 1863),
p. 65; Gthr, Cat. Brit. Mus., Fish., vol. VI, p. 166; Fed-
ders., Tidskr. Fisk. Kbhvn, IV (1870), p. 100; Coll., Forli.
Vid. Selsk. Chrnia 1874, Tillaegsk. p. 162; Malm, Ghgs,
Bch. Fna, p. 549; Seidl., Fna Balt., p. 95; Winth., Naturh.
Tidskr., Kbhvn, ser. 3, vol. XII, p. 44; Fedders., ibid.,
p. 78; Bncke, Fiscli., Fischer., Fiscliz. O., W. Preuss., p.
155; Mela, Vert. Fenn., p. 344; tab. X; Day, Fish. Gt.
Brit. Irel., vol. II, p. 121, tab. CXXI, fig. 1; Mob., Hoke.
Fiscli. Osts., p. 128; Norback, Handl. Fiskev., Fislcafv., p.
396; Reut., Sundji., Finl. Fisk., tab. XV; Sm. Riksm. Balm.,
1. c. ; Lillj., St)., Norg. Fna, Fisk., p. 630.

Atherina inordax , Mitch., Trans. Lit., Phil. Soc. N. Y., vol. I,
p. 446; Gill. (Osmerus), Smithson. Misc. Coll., No. 283,
p. 32; Jord., Gilb., Bull. U. S. Nat. Mus. No. 16, p. 293;
Br. Goode, Fisher., Fish. Industr. U. S., sect. I, p. 543,
tab. 199.

Salmo (Osmerus) spirinchus, Pall., Zoogr ., vol. Ill, p. 387;

Cuv., Val., 1. c., p. 387. — nec Lillj., nec Sm.

Osmerus viridescens, Lesueuk, Jouru. Acad. Nat. Sc. Philad.,
vol. I, p. 230; Dek., N. Y. Fna, part. IV, Fish., p. 243,
tab. 39, fig. 124; Cuv., Val., Hist. Nat. Poiss., vol. XXI,
p. 388; Stor.. Mem. Amer. Acad. Arts, Sc., vol. VI, p.

a Cf. Smitt, Rik-smuseets Salmonider, p. 171, § (5).
b Ichthyol., Gen. Pise., p. 10.

c A Latinized form of the French eperlan, which according to Rondelet refers to the pearly lustre of the coloration.

870

SCANDINAVIAN FISHES.

327, tab. XXV, fig. 4; Gthr, Cat. Brit. Mus., Fish., vol.

VI, p. 167.

Eperlanus vulgaris, Gaim., Voy. Isl., Groenl., tab. 18, fig. 2.

In the largest Swedish lake, Wener, the Smelt
frequently attains a length of a foot or about 3 dm."
to the extreme tip of the caudal tin. In Lake Malar
too it is frequently of a fair size, about 27 cm. in
length* 6; but in the smaller lakes it seldom exceeds a
length of 10 — 20 cm. As we have mentioned above,
the Scandinavian fishermen distinguish by size and
odour between two kinds of Smelt, calling the larger
kind slom. The line between these two kinds is drawn
at a length of about 2 dm. According to Nokback
the Smelt

at a length of about 6 cm. weighs about 13 — 17 grammes,

„ „ „ „ „ 9—12 „ „ „ 34-43

„ „ „ „ „ 18-20 „ „ „ 85 — 127

„ „ j, „ 30 „ „ ,, 300 „

Pennant’s largest specimen teas 33 cm. long and weighed
227 grammes c.

The body is rather elongated and of a compressed,
fusiform shape, the upper and lower contours being
slightly and similarly arched. According to the ripeness
or emptiness of the sexual organs the greatest depth of
the body, which is situated at the beginning of the
dorsal fin, varies between 14 and 18 % of the length
of the body. The greatest thickness, which lies nearer
to the back than to the belly, and is fairly uniform
from the opercular region to the dorsal fin, varies be-
tween 8 1/2 and 11 % of the length of the body or

between 48 and 70 % of the greatest depth. The least
depth of the body (of the peduncle of the tail) varies
between 6 (or a little more) and 4 lj2 % of the length.
The form of the body is thus subject to considerable
variations, and is as a rule deeper in the females than
in the males'2. The back, including the forehead, is
broadly rounded, as well as the belly, which at the
ventral fins even approaches to flatness.

The head resembles a compressed cone, the point
being formed by the tip of the lower jaw; and its
length from the articular knob of the maxillaries e is

somewhat less than 1ji (21 — 24 %) of the length of the
body. The convex snout resembles an obliquely cut-
cone, and consequently the sharp margin of the upper
jaw forms an ellipse. Its length is about 26 — 30 % of
that of the head7, the percentage rising with fair uni-
formity during growth, and its breadth across the
articulation of the maxillaries is only slightly less.
The longitudinal diameter of the eyes is somewhat-
greater than the vertical, measuring in young Smelts
about 1/i, in old about 1/6 , of the length of the head,
or in the former nearly 2/5 , in the latter about V4, of
the length of the head reduced (from the articular knob
of the maxillaries to the hind margin of the preoper-
culum). In the former their longitudinal diameter
sometimes is more than 1/i greater than their distance
from the articular knob of the maxillaries, in the latter
it sometimes sinks to 3/ 4 of this distance. The least
breadth of the interorbital space increases even rela-
tively with age from about 22 to about 28 % of the
length of the head, or from about 1/3 t-o 2/5 of the
length of the head reduced. The nasal cavities lie just
behind the articular knob of the maxillaries, the di-
stance from the middle of the tip of the snout- to the
front margin of the anterior nostril being, however,
nearly twice that- from the hind margin of the posterior
nostril to the front margin of the eye. The two nostrils
on each side of the snout are small, the posterior
somewhat larger than the anterior, but usually covered
to a great extent by the backward dermal fold con-
taining the thin ridge that- divides the nostrils. The
intermaxillary bones are about equal in length to the
eyes; and the length of the maxillaries, which increases
even relatively with age, varies between about- 9 (8’/2)
and 11 % of that of the body, between 42 (40 V2) and
50 % of that of the head, or between 62 (60) and. 67
% of that of the head reduced. The supplementary
(jugal) bone of the maxillaries is behind lancet-shaped,
in front elongated to a point. Its anterior part lies
loose in the dermal fold uniting the maxillary bone to
the cheek. The lower jaw is of a flat, boat-like shape,
in front curved uniformly, but rather slightly upwards.

“ Tenow gives even 4 dm. (See Ver inlands och Dais Ryggradsdjur , p. 108).

6 Underd. Bet. m. Forsl. t. Fiskeristadga 1883, p. 158; Lilljeborg, 1. c., p. 631.
c Brit. Zool. (London 1776), p. 274.

d In Osmerus dentex and O. spirinchus the sexual difference seems to be reversed. Cf. Smitt, 1. c., p. 176.

e For the sake of uniformity in a comparison with Coregonus we have measured the length of the body etc., here as in Riksmuseets
Salmonider (1. c.), from the articular knob of the maxillaries, and not from the middle of the tip of the snout.

f In Riksmuseets Salmonider the length of the snout in the Smelt is measured from the articular knob of the maxillaries, and there-
fore less.

SMELT.

871

Its length, which shows with fair uniformity even re-
lative increase with age, varies between 12 1/2 and 1 (5 1/2
% of that of the body, between about 60 and 76 % of
that of the head, or between about 86 and 98 % of
that of the head reduced. The mouth and pharynx
with their dentition have already been described. The
gill-rakers are set in one row of 34 — 37 on the first
branchial arch, and are setiform, tine, dense, and long.
The pseudobranchhe compose a narrow, longitudinal
band on each side, made up of about 10 short, trans-
versely set threads, just behind the boundary between
the palatine arch and the branchial cavity, and running
straight back from the upper insertion of the first
branchial arch. The operculum is quadrilateral, trape-
zoidal, with the upper margin shortest, the posterior
slightly concave at the top, and the inferior straight.
The length of the last-mentioned margin, which is di-
rected obliquely backwards and upwards as in the Sal-
mons, varies in different individuals between about 32
and 24 % of that of the head. The suboperculum also
resembles that of the Salmons, being rather narrow but
of uniform breadth, and rounded at the lower posterior
corner. The interoperculum is triangular, most pointed
in front. The preoperculum is rounded at an obtuse
angle. The gill-openings are large, extending up above
the middle of the superior opercular margin and down
to a level with the centre of the eyes. The branchio-
stegal membranes are broad and thick, free both from
each other and the isthmus; but at the extreme front
they cross, the left overlapping the right. The first-
two (three) branch! os tegal rays are slender, setiform; '
the other five sabre-shaped, gradually increasing in
breadth backwards (upwards).

The dorsal tin begins at a distance from the arti-
cular knob of the maxillaries that increases with age
from about 46 to 50 % of the length of the body. Its
base varies between about 7 1/2 and 8V2 %■> and its
greatest height (the length of the 3rd or 4th ray) be-
tween about 14 and 15 %, of the length of the body.
Above it is obliquely truncate, with the upper posterior
margin straight or somewhat convex. The adipose fin
is of the same shape as in the Salmons, but of a di-
stinctly fibrous texture. It is situated above the poste-
rior part of the anal fin, at a distance from the caudal
fin measuring about Vio — Vs (9 Vs — 12 Vs 0/0 ) °f the
length of the body. The anal fin begins at a distance
from the articular knob of the maxillaries equal to about
70 % (691/, — 72 Vs %) of the length of the body, the

percentage being generally greatest in the females. The
length of its base varies between 14 and 11 %, and its
greatest height (the length of the 3rd — 5th ray) between
10 and 81 /s %, of the length of' the body. Its inferior
margin is somewhat concave. The caudal fin is deeply
forked, the length of the middle rays being only slightly
greater than (sometimes equal to) the least depth of the
tail, or about 2/5 (varying between 35 and 43 %) of
the length of the longest caudal rays, which varies be-
tween about 17 and 15 % of that of the body.

The pectoral fins are set as in the Salmons, rather
low. When folded, they are of an obliquely pointed
shape; when expanded, oval. Their length varies be-
tween 16 and 14 % of that of the body. The third
(second branched) ray is commonly the longest. The
ventral fins are of a broader shape, and are always
shorter. Their length varies during growth between
1472 and 12 % of that of the body. Their distance
from the tip of the snout (i. e. from the articular knobs
of the maxillaries) is generally the same as that of the
dorsal fin. The preabdominal length is as a rule more,
the postabdominal length less, than 1/i of the length of
the body, the former varying between about 29 and
26 % , the latter between about 22 and 23 % thereof.

The structure of the scales we have already de-
scribed. Below Ave shall see hoAV the scales of the lateral
line indicate the origin of those of Argentina and Sco-
pelus, as a transition stage in this respect between the
said genera and the Capelin. During the spawning-
season both the males and females are covered with a
granulate dermal eruption, Avhich varies, hoivever, con-
siderably both in prominence and extent.

The internal organs and the most important pecu-
liarities of the skeleton have also been described in
the preceding pages.

The coloration is principally characterized, here as
in the following genus, by the thinness and transpa-
rency of the scales; and this transparency is commu-
nicated in some degree both to the dorsal musculature
and the head, the vertebrae as Avell as the cerebral
parts being visible in the living fish. The back and
the upper portion of the head are pale green or some-
times dashed with bluish gray, and doAvn the sides,
especially in the breeding males, this colour passes into
a lustrous violet. Along the middle of the sides runs
a silvery band, and this silvery lustre extends to the
shoulder-girdle, the opcrcula, and the cheeks. Under
the scales the skin is strewn on the upper parts of the

Scandinavian Fishes.

110

872

SCANDINAVIAN FISHES.

body with tine spots of bluish black pigment, more
scattered but larger in the silvery band. Down the
sides these spots become still more dispersed, until they
disappear on the belly. The snout, which is otherwise
transparent, and the tip of the lower jaw bear the
densest and largest spots of pigment. The silvery

peritoneum gleams through the ventral sides. The lower

*

part of the tail is transparent, of the same colour as
the back, only paler (yellower), and the median line of
the belly is partly transparent, partly more or less
milk-white. The iris is of a silvery lustre, but the
upper part of the eyeball is coated with a thick, bluish
black pigment, which sometimes advances over the top
of the iris. In the tins the membrane is transparent,
and the rays are of a lighter or darker gray, darkest
in the caudal and pectoral tins, and, after these, in the
anterior part of the dorsal tin.

One of the most noticeable characteristics of the
Smelt — which it possesses, however, in common with
the Capelin — is its peculiar odour. This is most
powerful in young specimens, and comes nearest to the
smell of the cucumber, though not without a trace of
violet perfume. It is at all events far from pleasant
to the ordinary individual, and its seat being the dermal
mucus of the tish, it communicates itself to everything
with which the Smelt conies in contact.

The Smelt has an extensive range in the North
Atlantic from east to west, for Osmerus mordax ( viri -
descens), the form belonging to the east coast of North
America, can hardly be regarded as a species distinct
from our common Smelt. On the Atlantic coasts and
in the rivers that fall into the Atlantic Ocean, its oc-
currence is, however, confined, generally speaking, to
the zone bounded by the 40th and 60th degrees of
latitude", though in the Baltic it is found further north,
up to the head of the Gulf of Bothnia. South of France
it is unknown, and it is not common south of the
north-west of that country; but from this region, in-
cluding the British Isles and the Continent, up to the
south-east of Norway, throughout the greater part of
Sweden, in Finland, and in Russia, it occurs, and in
suitable localities is common, within the basins of the
North Sea and the Baltic. In Russia, according to

Grimm* 6, it has also spread to the basin of the Volga.
It is really an anadromous tish, ascending the rivers in
order to spawn, and in the southern part of its range
its occurrence is almost exclusively restricted to tidal
waters. But in many places it has become a stationary
fresh-water tish. In the lakes of Jutland it is fairly
common, according to Kroyer, who had also procured
specimens from Roskilde Fjord, and records a statement
as to its occurrence in Lake Fur (Zealand); but with
these exceptions the Smelt is said to be wanting on the
Danish islands. In Norway, according to Collett, it
is found only in Christiania Fjord and in fresh water
south of the Dovre Fjeld, though not west of Lake Nord
in Telemark; and on the west coast it is said to be
wanting0. According to the reports sent in to the
Fisheries Commission of 1881 — 83 the Smelt is wanting
in Sweden only within the Governments of Jonk5pingd,
Kronoberg, Gothland, Blekinge, and Halland. Through-
out Finland the Smelt is common, up to lat. 66 N.
(Malmgren); but in the White Sea it is replaced, as
we have mentioned, by Osmerus spirinchus. A singular
gap in its geographical range appears in the Baltic
within the Sound, where it is wanting or at least ex-
tremely rare on the Scanian coast, off Bornholm, and
on the coast of Blekinge. The gap reminds us of a
similar break in the range of the Common Sea-Snail0,
though the Smelt is by no means an Arctic species.
It is especially common in Russia, North Germany
— particularly in the Haffs — and the Netherlands;
and the Smelt fisheries of the Seine and Thames are
to Paris and London what the fishery in the Norr-
strom is to Stockholm. In Irish waters the Smelt
seems to be wanting.

“The Smelt is of a stupid and sluggish tempera-
ment”, wrote Ekstrom, and this opinion has afterwards
been reiterated by other writers — “silly as a Smelt,”
is a common Swedish saying. But why it is thus stig-
matized more than other fishes, we cannot say. Gathered
in shoals during the spawning, when it is ruled by
sexual instincts alone, it is easy to catch like many
other fishes; and this is probably the origin of its re-
puted stupidity. It is a voracious fish-of-prey — which
may easily be seen by its teeth — and its form of body

“ According ro Gaimard (1. c.) the Smelt occurs on the coasts of Iceland; but Faber had no information on this head.

6 Fishing and Hunting in Russian Waters, p. 20.

c Olsen ( Piscatorial Atlas) states, however, that the Smelt occurs in Bucke Fjord off Stavanger.
d Yet it is found in Lake Wetter.
e Cf. above, p. 289.

SMELT.

873

indicates at least no lack of agility. But out of the
water it soon dies, no more tenacious of life than the
Herring. It is a migratory fish like the Salmons, though
not in so high a degree, roving at the spawning-season
from salt water to fresh, or, in the lakes, from deep
water to the shallows. As the spawning-season ap-
proaches, it assembles in large and dense shoals; but at
other times it leads a more solitary life, being fre-
quently taken in the Herring-nets used in the Baltic,
but not in any great number. It feeds principally on
fish, small or large — at least up to half of its own
size — and especially on Herring-fry and the young of
its own species. Other kinds of food, however, such
as crustaceans (shrimps and Gammaroids), worms, and
larva?, do not come amiss.

According to Yarrell the marine Smelt of the
English coasts repairs to fresh or brackish water, and
remains there from August to May. In the Norrstrom
off Stockholm a female 196 mm. long and a male 188
mm. long, both quite ready to spawn, were taken on
the 4th of November, 1892; but as a rule the Smelt
does not muster in Sweden for its breeding expeditions
until the end of March or even later. From the island-
belt of Sodermanland Ekstrom wrote, “In March or
April, according to the earlier or later breaking up of
the ice, the Smelt ascends to rivers, straits, or shores
where there is some current, always choosing, however,
water of some depth with a clean, sandy bottom. It
generally -rises' towards evening and continues its jour-
ney the whole night, but at daybreak again retires for
the most part to deep water. A remarkable circum-
stance is that, whereas all other fishes prefer to spawn
in fine weather, in the Smelt the case is just the re-
verse. In squally and snowy weather it is most eager
in its ascent, the violent gusts of wind and snow that
occur during the said months being hence known as
nors-il (Smelt squalls). Males and females swim in
company during the spawning, and are so densely
massed that they seem merely to rub their bodies to-
gether in order to rid themselves of the roe, which is
deposited on the bottom beneath.” The young start
first, but do not ascend so far up the rivers as the
older fish, and often spawn in the lakes on shallow
shores. Each shoal completes its spawning operations
iii a few days; but one shoal follows in the wake of

another, and thus the spawning continues as a rule
from the latter part of March to the first weeks of May.
The greater part of the shoal is composed of females,
and after the spawning the shore and the bottom are
strewn with numbers of dead Smelts which have strug-
gled in vain to disburden themselves of the roe". The
ova are light yellow; their diameter was estimated by
Benecke at 0'6 — 0'8 mm., and their number in a fe-
male 18 — 20 cm. long by Norback at 50,000, by Olsen
at about 36,000. They attach themselves in a singular
manner to the objects on which they fall after impreg-
nation. According to Cunningham b, the outer membrane
of the ovum, the so-called zona radiata , “is differen-
tiated into two layers, the outer of which is somewhat
thinner than the internal. In the zona radiata externa
the pores are larger and farther apart than in the in-
terna. But the important fact, which I believe no one
has previously observed, is that the external zona sepa-
rates very readily from the internal, and, rupturing at
one portion of the ovum, peels off, becoming turned
inside out in the process, and, remaining attached over
a small circular area., forms the suspensory membrane”,
by means of which the ovum is attached to any ex-
ternal object.

The eggs are hatched, according to Blanchere, in
8 — 10 days, according to Feddersen in 12 days, and
according to Sundevall in 18 days, a discrepancy of
observation which in all probability depends on the dif-
ferent temperature of the water during the period of
incubation. On their first exclusion the fry are elongated,
according to Sundevall 5 mm. long, and perfectly trans-
parent; they are characterized by the unusually back-
ward position of the vitelline sac, the distance between
it and the insertion of the pectoral fins being more than
half of that between these fins and the tip of the snout.
Ehrenbaum0 gives some personal observations on the
growth of the larval Smelts. In the first month after
the hatching, according to him, they grew to a length
of 14 — 18 mm.; in the second he found them to be
27 — 34 mm. long, in the third 32 — 37 nun., in the
fourth 35 — 44 mm., in the fifth 44 — 60 mm. Still,
in August, according to Yarrell, they already measure
as much as about 71/2 cm. The Smelt attains maturity
in the following spring, or, according to Norback,
even at a length of 43 — 60 mm.

a Faith in Krdyer (1. c.) and Feddersen in the Tidslcr. f. Fiskeri, vol. IV (1870), p. 102.
b Proc. Zool. Soc. Lond. 1 886, p. 292, tab. XXX.
c Sonderbeil. Mittheil. Sekt. Kiist., Hochs. Fisch., Jalirg. 1892, p. 12.

874

SCANDINAVIAN FISHES.

The rank odour emitted by the Smelt offends the
taste of many; but the flesh is good, and the belly
between the spawning-seasons full of fat. Among epi-
cures it is esteemed as a delicacy, after the disagreeable
smell has been removed by very simple culinary me-
thods. The fish must also be carefully gutted, especi-
ally if in breeding condition, to rid it of the numerous
intestinal worms, which penetrate even into the air-
bladder. It is best fried and served with lemon-juice
or vinegar, but is often boiled or stewed in sauces.
It is also split and dried for future consumption, and
in this state may be eaten without further preparation".
Another method is to soak the dried flsh in lye, and
afterwards dress it for table like other stockfish. In
addition to its utility as human food, the Smelt also
possesses importance as one of the best baits for pre-

readiness. When the Smelt spawns on shores or off
headlands, it is taken with drag-nets, which differ from
an ordinary seine only in the comparative fineness of
the meshes. This fishery is pursued only at night, bon-
fires being not unfrequently lit on shore by the fisher-
men, in the belief that the fish, enticed by the glare,
come nearer land.” The well-known Smelt-fishery in
the Norrstrom off Stockholm is carried on with large
hoop-nets (tig. 219), such as are in general use at many
places among the island-belt of Stockholm, to secure
all kinds of small fishes for bait. These hoop-nets,
usually about 3 — Sl/2 in. in diameter, are let down and
hoisted up from a boat, with the aid of a long pole erected
obliquely upwards in the stern; and a hand-net is em-
ployed to scoop the fish out of the large net. The Smelt
is also caught on the hook with a bait of shrimps,

Fig. 219.

Smelt-fishing with hoop-net from a boat.

datory fishes of a greater size; and in several localities
where it is taken in too great a quantity for imme-
diate use, it is even made into guano.

Of the Smelt-fishery Ekstrom writes, “It is during
the spawning-season that the Smelt is taken in any
quantity, the fishery being commonly conducted in the
following manner. Across the straits or the rivers to
which the Smelt ascends in order to spawn, fences are
built of green spruce branches, arranged so as to leave
gaps at the deepest parts of the channel. At these gaps
the fisherman stations himself with a scoop-net large
enough to fill the opening and breaded of meshes so
fine that the Smelts cannot slip through. This net,
which is distended on staves, he lets down into the
opening, and takes up after a longer or shorter interval
according to the numbers of fish that come up, the take
being then turned out of the net into a coble held in

sand-hoppers (Gammaroids), worms, or bits of fish; but
this method is successful only when used for the large
Smelts, the slorn or nor skiing as they are called in
some parts of Sweden 'when they occur in solitary
specimens among the smaller Smelts.

Man is not the only enemy which the Smelt has
to fear; it often falls a victim to predatory fishes and
waterfowl. Although its great fecundity can, no doubt,
compensate in most cases the losses inflicted by an ac-
tive fishery, still it is advisable to protect the young fish.

In lakes where the Smelt is wanting, it may easily
be introduced, and is very useful, especially as food
for other fishes, in particular for the Pike-perch, as
Nilsson has pointed out. The impregnated eggs may
be transported from one lake to another, or with a little
care the spawning Smelt may be conveyed alive during
the cool season in vessels filled with pure water.

“ “It used to be split and dried and was thus considered to add a peculiar relish to the morning dram of spirits”. Day.

SALMONIDJE.

875

Genus MALLOTUS'.

No canines; small , fine teeth , hardly distinguishable to the naked eye, set in one row in the anterior part of the
lower jaw , on the inter maxillaries, the maxillaries, the head of the vomer, the anterior extremity of the palatines,
the mesopterygoid bones, and the tongue. Length of the maxillaries as a rule 57b — 63 %, and, that of the lower
jaw as a rule 7 5C — 90 %, of the length of the head reduced. Number of rays in the anal fin 20 — 24, 17d — 20
being branched; and the base of the anal fin longer than that of the dorsal. C cecal diverticula at the pylorus

short and few ( about 4 — 6). Scales smalt, about 200 in a row along the sides of the body, and the lateral line

complete, at least to the perpendicular from the end of the anal fin.

Between Mallotus and Osmerus stands the Arctic
American Eulachon (Thaleichthys pact ficus), with scales
only slightly smaller than those of the Smelt and den-
tition feeble as in the Capelin C Thus we cannot ex-
pect to trace the descent of the Capelin immediately
from the Smelt or the reverse. In several respects, it
is true, we find that the Smelts represent an earlier
stage of development, more nearly approximated to the
presumable original form. One of the most prominent
differences between the two genera lies, for example, j
in the shape of the anal fin, the height of which is
always more in the Smelts, less in the Capelins, than
half its base. Now in the former7 we find in the aver-
ages a difference both of age — the young have a rela-
tively higher anal fin than the old — and of sex — the
females have a relatively higher anal fin than the
males — both of which relations point to the transition
from Osmerus to Mallotus. In most other respects,
however, Mallotus comes nearer to the original form,
such as we may assume this form to have been, jud-
ging by the manner in which the characters appear.
Thus, for example, Mallotus has a narrower snout than

Osmerus , the breadth of the snout across the articular
knobs of the maxillaries being less in Mallotus, more
in Osmerus, than 1 /'4 of the length of the head or 1/3
of the length of the head reduced. In Mallotus as in
Osm. eperlanus the percentages for these relations also
rise with increasing age. The sexual distinction is,
however, different here: in the Capelins (as in Osm.
dentex and Osm. spirinchus ) the snout is as a rule
narrowest in the females, in Osm. eperlanus in the
males. Another character wherein Mallotus also repre-
sents a lower grade of development, may be found in
its adipose fin, which is of uniform height, long and
low, and which, like the embryonic fins that we have
remarked in several preceding genera, is furnished
with a countless number of fine supporting fibrils.
The feeble dentition approximates Mallotus more close-
ly to the following subdivision of the Salmonoid
family, although the immediate transition from Osme-
rus to the said subdivision passes through other
genera, as for instance Nelma ( Stenodus ) and Hypo-
mesas.

Only one species of the genus is known C

a

a Greek gaXXcoxoq = Lat.
6 In exceptional cases 55.

villosus,

hairy.

e Eulaclion is besides remarkable for its fatness. In a dried form it is a food of good flavour and, when lit. burns like a lamp.
f Most distinctly, judging by the Royal Museum specimens, in Osmerus dentex and Osm. spirinchus.

rj The difference given in Rilcsmuseets Salmonider between the Capelins of the Old and New Worlds has proved, on examination of
greater number of specimens from Greenland, to be untenable.

876

SCANDINAVIAN FISHES.

THE CAPELIN (sw. loddan).

MALLOTUS VILLOSUS.

Plate XLI, figs. 2 {(p) and 3 (9).

Milters and spawners widely different , the former with the length of the pectoral fins more than 15 % of that
of the body , with the anal base , which is elevated in an arcuate curve beyond the ventral line , more than 16 %
of the length of the body and perceptibly greater than the length of the head reduced , and also with the scales
in a longitudinal row above the lateral line and in another along each side of the belly prolongated and pointed

or at least distinctly larger than the scales on the rest of the body.

B. to. 8 — 10"; D. A. ;

10—13 (16)17—20' 15-19(20)

1 jg 20

V. ; 0. x + 1 + 17 + 1 4- x; L. lat. ca 200; L. tr. ca -

7 14 -16

supra pinn. ventr. ; Vert. 67 — 70J.

Syn. Anchovas et Capelinas , Parkhurst in Hakluyts Voyages 1598
— 1600, vol. Ill, p. 133; Egede, Gronl. N. Perlustr., p.
50; StrSm, Sondm. Beskriv., part. I, p. 293; Leem,
Beskr. Finm. Lapp., p. 323; Olafs., Reis. Isl., part. I,
p. 358, tab. XXVIII. Angmarset, Cranz, Hist. Gronl.
(ed. suec. 1769), part. I, p. 124. Capelan de VAmerique
septentrionale, Duh., Peches, part. II, p. 149, tab. XXVI,
figg. 1—8.

Sahno Eperlanus, Mull., Zool. Dan. Prodr., p. 48 (ex Str5m).

Clupea villosa, Mull., ibid., p. 50 (ex Olafs.); Vahl, Rathke
{Salmo) in MOll., Zool. Dan., part. IV, p. 45, tab. CLX;
Faber, Fisoh. Isl., p. 174; Richards. [ Sahno {Mallotus) ex
Ccv.] Fna Bor. Amer., part. Ill, p. 187; Gaim. {Mallotus),
Voy. Isl., Groenl., tab. 18, fig. 1 ; Cuv., Val., Hist. Nat.
Poiss., vol. XXI, p. 392, tab. 622 et 623; Mgrn, Finl.
Fiskfna (disp. Helsingf.), p. 66; Gthr. Cat. Brit. Mus., Fish.,
vol. VI, p. 170; Coll., Forli. Vid. Selsk. Chrnia 1874, Til-
lsegsh., p. 163; ibid. 1879, No. 1, p. 86; N. Mag. Naturv.
Chrnia, Bd. 29 (1884), p. 106; Mela, Vert. Fenn., p. 345,
tab. X; Joed., Gilb., Bull. U. S. Nat. Mus., No. 16, p.
291; Storm, Norsk. Vid. Selsk. Skr., Trondhj. 1883, p. 28;
Br. Goode, Fischer., Fischer. -Industr. U. S., sect. I, p. 544,
tab. 201; Smitt, Riksm. Salmonid., p. 189; Lillj., Sv.,
Norg. Fna, Fisk., vol. II, p. 646.

Sahno arcticus, Fabr., Fn. Gronl., p. 177; Nilss. {Osmerus),
Prodr. Ichthyol. Stand., p. 11; Kr. {Mallotus) in Gaim.,
Voy. Scand., Lappon., cett., tab. 16, fig. 1; Id., Damn.
Fisk., vol. Ill, p. 23; Nilss. {Osmerus), Skand. Fna, Fisk.,
p. 441.

Sahno groenlandicus, Bl., Naturg. Ausland. Fisch., pt. VIII,
p. 99, tab. CCCLXXXI, fig. 1 ; Cuv. {Mallotus), R'egne
Anim., ed. 2, tom. II, p. 306.

Salmo socialis, Pall., Zoogr. Ross. Asiat., torn. Ill, p. 389,

Osmerus microdon , Cuv., Val., Hist. Nat. Poiss., vol. XXI.
p. 385, tab. 621.

The male Capelin is as a rule not more than 19
cm. in length to the end of the caudal lobes, and the

female not more than 17 cm.; but Lilljeborg mentions
males of a length of 22 cm., and females of a length
of 18 cm. According to the method in which our
measurements are here taken, from the articular knobs
of the maxillaries to the end of the middle caudal
rays, the length of the body in our specimens does
not exceed 17V2 cm. for the males or 16 cm. for -the
females.

The great difference between the sexes, which is
so characteristic of the genus that we have included it
in the above diagnosis — no other character is neces-
sary, only one species of the genus being known —
appears in the very form of the body. The Capelin is
elongated and, in contradistinction to the Smelt, as a
rule of more uniform depth and more compressed. In
the adult males we have found the greatest depth, at
the beginning of the dorsal tin, to vary between about
14 and 15 % of the length of the body, and the great-
est thickness, which is fairly uniform above the lateral
line from the shoulder-girdle to the end of the true
dorsal fin, between 7J/2 and S1/i % of the same. In
the adult females the corresponding percentages have
varied from 11 to 1 3 and from 5 to 8 respectively.
In the males the Greatest thickness has varied between
53 and 60 %, in the females between 43 and 62 %, of
the greatest depth. The least depth of the body shows
on an average the same sexual difference, being in the
females about 5 %, in the males about 57s % of the
length of the body. The dorsal edge is broadly con-
vex, and the dorsal profile almost straight from the
very occiput* 6, not beginning to descend towards the
base of the caudal fin until past the dorsal fin proper.
Below the lateral line the sides gradually converge

8 Sometimes up to 11, according to Kr0YER.

6 Sometimes 65, according to Krgyer.

c In all our fresh, but dead specimens there appeared in the median line of the back, just behind the head, a compressed swelling,
due to a callosity of the central sinew in the musculus gracilis, where the latter broadens towards its point of attachment on the occiput.

CAPELIN.

877

towards the ventral margin, which is narrower, but
terete. The ventral profile in front runs almost pa-
rallel to the dorsal, beginning to rise towards the pe-
duncle of the tail at the origin of the anal fin, in the
females uniformly, but in the males with a break
caused by the arcuate expansion of the base of the
said fin.

The shape of the head reminds us most of a three-
sided pyramid, two sides of the pyramid being formed
by the cheeks, which approach each other below, and
the third by the flat forehead and occiput. Its length,
which is greatest in the females and the young, varies
between 23 and 19 % of that of the body, and the
length of the head reduced, from the articular knobs
of the maxillaries to the hind margin of the preoper-
culum, similarly between 15'/2 and 13 % of the length
of the body. The horizontal profile of the tip of the
snout is broadly rounded, but its breadth at the arti-
culation of the maxillaries is considerably less than in
the Smelt, measuring only 1 fi — 20 % (in exceptional
cases 21 %) of the length of the head. The length of
the snout, on the other hand, is greater than in the
Smelt, being about 33 — 30 % of that of the head. The
eyes are also comparatively larger than in the Smelt:
in adult Capelins their longitudinal diameter, which is
somewhat greater than the vertical, is about 24 — 23 %
of the length of the head. The postorbital length of
the head is consequently always less in the Capelin
(about 44 — 48 %) than half the length of the head,
while in the Smelt it is more than half of the same.
The forehead is flatter, but its breadth, like that of
the snout, is on an average less than in the Smelt,
being only about 24 — 22 % of the length of the head,
and in exception to the general rule in the family, it
is the males that here represent the earlier stages of
development. In the opercular apparatus the operculum
itself is of the same shape as that of the Smelt, but
the sinus at the hind margin is indistinct or wanting.
The suboperculum is somewhat larger than in the Smelt,
and more nearly resembles the quadrant of an ellipse.
The lower posterior angle of the preoperculum is al-
most a right angle. That portion of the margin of the
upper jaw which is formed by the intermaxillaries
measures only about % — 5/7 (63 — 74 %) of the longi-
tudinal diameter of the eyes; but the length of the
maxillaries is about the same as in the Smelt, on an
average about 41 %, varying between 38 and 44 %, of
that of the head. The length of the lower jaw, on an

average less than in the Smelt, is about 57 %, varying
between 55 (exceptionally 53) and 60 (exceptionally
62) %, of that of the head; but the more essential
difference in the shape of the lower jaw between these
two genera is due partly to the downward convergency
of the sides of the head in the Capelin, the halves of
the lower jaw being thus brought nearer to each other,
partly to the less marked curvature in an upward di-
rection of the point of the lower jaw, which, however,
projects, even in the Capelin, beyond the tip of the snout.
As regards the distribution of the teeth in the mouth,
palate, and pharyiix the Capelin in all essential respects
resembles the Smelt; but all the teeth of the Capelin
are of uniform size and small, about equal in size to
the maxillary teeth of the Smelt. On the true hyoid
bone (os linguale or glossohyale ) lies an elliptical row
of teeth within the fleshy margins of the tongue, with
one or two teeth set inside the ellipse, and on each
mesopterygoid bone the row of teeth is not situated,
as in the Smelt, .at the very inner margin, the bone
having grown further inwards to form a more com-
plete bottom for the orbit. The gill-rakers resemble
those of the Smelt, but are as a rule somewhat more
numerous, 35 — 39 on the front of the first branchial
arch, and 25 — 30 of these on the lower part of the
same. The pseudobranchise lie just behind the orbits
and above the dermal fold that bounds the palate be-
hind. They consist of 15 or 16 short lamellse, set in
an oblique transverse row. The gill-openings are large,
extending from about half the upper edge of the oper-
culum to a line with the anterior margin of the eyes,
and the two branchiostegal membranes are entirely free
from each other, crossing only to an extremely small
extent in front, where the left membrane overlaps the
right, but Avhere each is covered by an outer fold of
the skin, the right fold lying outside the left. The first
(lower) four branchiostegal rays are slender and subu-
late, the last (upper) 4 — 6 gradually become flatter
and broader (sword-shaped) behind (upwards).

The dorsal fin begins at a distance from the arti-
cular knobs of the maxillaries that measures about 50
— 55 % of the length of the body, farthest back as a
rule in the females. Its base measures about 1/10 (9
— 12 %) of the length of the body, and is generally
somewhat less (but sometimes, especially in the males,
somewhat greater) than its height as expressed by the
length of the longest (3rd and 4th, i. e. first branch-
ed) rays. When erected, the fin is of an obliquely

878

SCANDINAVIAN FISHES.

quadrangular shape, with somewhat convex upper mar-
gin, and with the hindmost ray measuring about 2/3
of the longest ray. The shape and structure of the
adipose tin have already been noticed. It is situated
above the posterior three-fifths of the base of the anal
fin, but the distance between it and the caudal fin
(about 7 — 11 % of the length of the body) is in ge-
neral perceptibly greater than that between the anal
tin and the latter. The anal fin begins at a distance
from the articular knobs of the maxillaries in the males
equal to about 2/3 (66 — 68 V2 %), in the females nearly
3/4 (71 — 7272 %)-> °f the length of the body. The
sexual difference in the base of this fin we have al-
ready remarked. The length of its base is in the males
about 17 — 19 % , in the females about 13 — 15 %, of
the length of the body. Its lower margin is arcuate,
and its height (the length of the longest, the 5th or
6th ray) is always less than half" (38—47 %, excep-
tionally 49 %) of its length (base). The caudal fin is
as a rule somewhat less deeply forked than in the
Smelt, the length of the middle rays being about 43
— 53 % of that of the longest ones and usually greater
than the least depth of the tail, only exceptionally,
and then in the males, equal to the latter. The length
of the middle rays varies between about 5 72 and 61/2
% (exceptionally 7 %), and the length of the longest
rays between about 12 and 14 %, of that of the body.

The pectoral fins are set low, as in the Smelt,
and the great sexual difference lies not only, as we
have mentioned above, in the length of the fins, but
also in their breadth. In the males, when the fins are
folded, one of them overlaps the other under the belly
to a considerable extent. When expanded, the two
fins together form in the males almost a semicircle.
In the females they are of a broad oval shape. Their
length varies in the males between about 15 h and
16 72 0//°i in the females between about 10 and 137' 2
of that of the body. As a rule even their relative
length increases with age, the females thus representing
the lower grades of development. The ventral fins
resemble the pectoral both in shape and sexual dif-
ference, though the latter is less marked, and the
ventral fins of the males together compose, more cor-
rectly speaking, a half-ellipse. Their length is also on
an average equal to that of the pectoral. They are

inserted almost vertically belotv the beginning of the
true dorsal fin, comparatively further back in young
Capelins than in old and in the females than in the
males, the average preabdominal length being in young
females 31 72 in old females 3272 %, in the males
30 72 %>, of the length of the body, and the average
postabdominal length in the females 1 8 1/2 %, in young
males 1 8 1/3 %, in old males 1 7 1/2 %, of the length of
the body.

The scales are thin and small, as we have men-
tioned above, but in essential respects (fig. 220, a) re-
semble those of the Smelt, both as regards shape and

c ha

Fig. 220. Scales from the left side of a Mallotus villosus, 1 21/,
times the natural size, a, one of the ordinary scales below the lateral

line in a female; b , a scale from the lateral line of a female; c, one

of the larger scales at the base of the anal fin in a male; d. one

of the villiform scales in the lateral band of a male, with the epi-

dermal covering preserved, and with lobes of the torn dermal follicle
(corium) on each side.

texture, only that the concentric strife are few and
scattered (usually no more than 5 or 6 behind the
indistinct nucleus of the scale, 3 in front of the same),
and interrupted at the top and bottom of the scale.
The scales of the lateral line ( b ) have the same open
incision in the hind part, and the duct is simply cana-
liculate, nowhere closed by the elevated margins. The
rule for the growth of the scales in the lateral line
being that the duct is at first canaliculate, these scales
accordingly occupy in the Capelin a lower develop-
mental rank than in the Smelt. In the males, as we
have mentioned above, scales of a special' shape appear

“ In the Smelts always greater, 68 — 84 / in Osmerus eperlanus.

h In exceptional cases we have found the pectoral fins of a male to measure only 14‘7 % of the length of the body.

CAPELIN.

879

in four longitudinal bands®, one on each side of the
body above the lateral line, and one along each side
of the belly, at the boundary between the ventral side
and the side of the body, beginning below the posterior
part of the pectoral fins, when these fins are at rest,
interrupted at the ventral fins, but recommencing at
about the middle of their length and extending along
the base of the anal fin. On the elevated base of this
fin, as well as on the lower margin of the peduncle
of the tail, the remaining scales (c) are larger than on
the rest of the body and more or less oblique, with
one corner more or less elongated. In the above-
mentioned longitudinal bands the scales are elongated
to a villiform shape, but covered to the very tip with
skin, in which lie scattered pigment-cells, most nu-
merous and most constant at the tip of each scale,
rare and most scattered on the proximal part thereof.
Comparatively large scales form a, lobe on each side of the
vent, and this lobe is distinguishable even in the female.

The coloration in essential respects resembles that
of the Smelt. In the Capelin too the upper parts of
the head and body and the hind part of the tail are
transparent. The back is green; the sides are silvery
with black dots, but the operculum and suboperculum
yellowish (with a golden or brassy lustre); the belly
is white. Our figures, however, depict the spawning-
dress, in which the back is darker, and the head some-
times entirely covered with a sooty black pigment.
Below the above-mentioned villous bands of the males
runs a more or less prominent, yellowish streak. The
fins are transparent, of a light grayish colour; the
caudal fin is darkest, especially at the base, the pe-
duncle of the tail being also more or less blackish.

The internal organs too are similar to those of
the Smelt. The liver is small, only the middle and
the left lobes being present. It extends back to about
a line with the middle of the pectoral fins. The ciecum-
like stomach extends almost back to the anterior ex-
tremity of the pelvic bones. The pylorus is furnished
with 6 or 7 rather large appendages, four directed
forwards and three backwards, two of the latter on
the right side of the intestine. The pyloric coil and
the intestine are as in the Smelt. The spleen is nar-

row and ribbon-shaped, lying to the right, at about
the middle of the stomach, between the forepart of the
intestine and the air-bladder, which is comparatively
short and occupies a little more than the middle third
of the dorsal margin of the abdominal cavity. The
testes are double, but the right is considerably smaller
than the left, and only the left ovary is present, its
inward oviduct resembling that of the Smelt. The
peritoneum is sooty black inside, but its outer coat is
silvery white.

The Capelin is a boreo-arctic species from the
northern regions both of the Atlantic and the Pacific.
Off Spitsbergen none of the numerous Swedish expe-
ditions has met with the Capelin; but at certain times
it is extremely common on the coasts of Norwegian
Finmark, Iceland, Newfoundland, Greenland, Alaska,
the Aleutian Islands, and Kamchatka. From the White
Sea the Royal Museum of Stockholm has received the
species through Lieutenant H. Sandeberg. Its occur-
rence in shoals on the Norwegian coast does not really
extend beyond Trondhjem Fjord; farther south it is
more and more scattered, and the Capelins from Chris-
tiania Fjord sent to the Royal Museum in 1842 by
Professor Esmakk must rank as a rare find. On the
other side of the Atlantic the Capelin goes farther
south-, one of the consequences of the Arctic current
which follows the coast of North America. According
to Jordan and Gilbert its range extends to Cape Cod;
but Bean states that it does not occur much south of
Halifax in Nova Scotia, and is never met with on the
coast of the United States. Richardson found the spe-
cies in Coronation Gulf, whence it appears that in this
quarter the range of the Capelin extends between the
Atlantic and the Pacific; but off the extreme north of
Asia it has never yet been seen.

To all the above-mentioned points in which the
Capelin resembles the Smelt, we should add its rank
odour, which calls to mind that of stale cucumbers.
In its manner of life too the Capelin is very like the
Smelt. But it is of a more pelagic nature, and never
enters quite fresh waters. Its haunts and its life be-
tween the spawning-seasons are little known; but ac-
cording to J uel b it is frequently met with in summer

“ According to Fabricius these bands are sometimes wanting, even during the spawning-season, and the Esquimaux of Greenland have
a special name ( Sennersuitsut ) for such males. That the scales in these bands only gradually develop the villiform shape, being at first tri-
angular, appears from the remark on Capelin No. 22 in my Riksmuseets Salmonider.
b Norsk Fiskeritidende, 1892, p. 4.

Scandinavian Fishes.

Ill

880

SCANDINAVIAN FISHES.

by whalers out at sea in the Arctic Ocean. Fabricius
states that at this time it is less rash than during the
spawning-season, and seeks to evade its pursuers by
leaping out of the water. In some fjords it loiters
long after the spawning is over. This has been ob-
served in the upper part of Varanger Fjord", and also
in St. Mary Bay (Newfoundland), where, during the
winter-fishery under the ice, Capelin just devoured
have been found in the stomach of Cod taken in Janu-
ary6. It attracts all the more attention during the
spawning-season, when it gathers in immense shoals
and repairs to shallow water. This takes place gene-
rally late in winter and early in summer, off Norwe-
gian Finmark, according to Sars, in April, May, and
June, on the coast of Greenland, according to Fabri-
cius, in May, June, and July. But this rule is subject
to many irregularities, which are of importance not
only for the fishery, but also to shed light upon similar
variations in the appearance of the Herring. For se-
veral years, seemingly at periodic intervals, the Capelin
deserts its usual spawning-places; in other years it
comes at an unusual season, seemingly, as in the case
of the Herring, earlier at the beginning of a period.
Thus Sheriff Sommerfelt (1799) writes of the Capelin",
“that else they have the experience here that it has
abandoned the coasts of Finmark for many years, up
to 16 or 20, in succession”; and to the quintennial
reports of the sheriff (1830 — 1840), according to which
the ‘Capelin-fishery’ (i. e. the Cod-fishery with a bait
of Capelin) had not been practised on the coast of
Finmark, Juel appends the remark d: “As old fisher-
men state, however, the Capelin did appear off the
coast in one of these periods of five years, but came
early (in December) and departed in March, so that
no fisherman seized the opportunity. Subsequently the
Capelin came later and later in the year.” Similar
irregularities meet us in the Herring’s approach to the
coast. They very probably depend in an eminent
degree, as Sars assumed, on variations in the “physio-
meteorological conditions,” perhaps too on variations in

the supply of food and in the numbers of the Capelin’s
enemies. But as yet we can only record them among
the lessons of experience; their explanation we must
leave to future research. How important it would be
to Norway to ascertain the causes of these irregula-
rities, we may easily gather from the results of the
so-called Capelin-fishery, which depend on the said
irregularities, and which in 1875, according to Sars,
were estimated at nearly twenty million Cod, but in
some of the preceding and the following years at only
five or six million.

During its wanderings the shoal is incessantly
harassed by all kinds of enemies in search of food.
In their efforts to escape the Capelins crowd so close
together that the fish in the middle of the shoal are
even lifted above the surface, where they wildly lash
about on the backs of those beneath them. When the
weather is calm, the Capelin shoal looks like a tum-
bling, glittering wave on the surface. Above it hover
flocks of kittiwakes ( Larus tridactylus), which time
after time swoop down and seize a fish; and round the
shoal blow whales (the common rorqual, Baleen op ter a
musculus, and the lesser rorqual, Balcenoptera rostrated).
But the Capelin’s most eager pursuers are Cod and
Coaltish'. Meanwhile the shoal makes for the coast,
which it follows till it finds a suitable spawning-place.
Often enough it comes so near land that the fish may
be scooped up in a hand-net from the beach, or even
leap ashore to escape their pursuers. As a rule they
swim against the wind, a sea-wind keeping them from
the coast, and a land-breeze alluring them thither.
Enormous are the numbers in which the Capelin ap-
pears during these migrations, and many are the ac-
counts thereof. Thus Collett states that the Capelins
often spread in shoals many miles long, touching the
coast almost simultaneously at the extreme end of
Western Finmark and in Varanger Fjord. “The females
go first in a separate shoal,” writes Fabricius, “and seek
out places suitable for their progeny; the males follow and
seek out the eggs, to impregnate them with their miltb

a Sars, Loddejisket, ved Finmdrken, Indber. Depart, f. d. Indre, 1879, p. 11.

* Hind in Bean, 1. c.

c The quotation is taken from Juel, 1. c., p. 9.
d L. c., p. 11.

e These two according to Sars. According to Collett the Capelin is also eaten by the humpbacked whale ( Megaptera boops).
f Besides these, according to Collett, Anarrhichas minor and lupus, Hippoglossus vulgaris, and several more.

g According to Atwood (Proc. Bost. Soc. XIV (1872), p. 134) the males lead the way, and the females follow them to the spawning-
place, but in comparatively small numbers, one female to ten males.

CAPELIN.

881

When the females have left the place, the bottom is
so entirely covered with their eggs that it gleams with
a yellow colour; but when the males have shed their
milt on the ova, the water is milky white, and the
bottom no longer visible.” Pallas mentions “as a
curious fact” that several fish, two, three, or even ten,
cling so fast together with the aid of the villous lateral
line that, if one of them be taken up, the others fol-
low with it, as if they were glued to each other. He
adds, “Even in the sea they are seen swimming together
in this manner, and this union between the sexes is
perhaps necessary for the fertilization of the spawn.”
As we have seen, the males alone are furnished with
the villous bands; and Pallas’ observation can there-
fore refer only to the emission of the milt by the
mutual pressure of the companion fishes, a supposition
which bears out the statement that the males and fe-
males swim in separate shoals. Lanman, however,
gives an account" that points to a different conclusion.
“The female,” he writes, “on approaching the beach to
deposit its spawn, is attended by two male fishes, who
huddle the female between them, until the whole body
is concealed under the projecting ridges, and her head
only is visible. In this position all three run together,
with great swiftness, upon the sands, when the males,
by some inherent imperceptible power compress the
body of the female, between their own, so as to expel
the spawn from the orifice at the tail. Having thus
accomplished its delivery, the three capelins separate,
and paddling with their whole force through the shal-
low water of the beach, generally succeed in regaining
once more the bosom of the deep; although many fail
to do so, and are cast upon the shore, especially if the
surf be at all heavy.” If the observation be correct,
the spawning may thus be performed in two essentially
different manners.

On the coast of Finmark the spawning, says Saks,
“commonly takes place at a depth of 4 — 20 fathoms,
though it may possibly be performed on rocky bottoms
and in deeper water as well, on which head, however,
we have as yet no trustworthy observations6.” “In
Varanger Fjord,” writes Collett, “there is a good
and sure spawning-place off the mouth of the Jakob
Elf. At the flood the Capelins ascend high up the

river, as far as the water is brackish enough.” After
the breeding numbers of Capelins, especially old males,
float in a- dying state at the surface, or are cast ashore
in great heaps. We have above seen a similar mor-
tality, though not so great, attend the spawning of
the Smelt.

In 1879 Capelin fry were first observed by Sals
on the 17th of June off Vadso, after a fresh east wind
with a strong landward current. They were then, he
says, evidently just hatched, none of the specimens
collected measuring more than 8 — 10 mm.; but the
hatching-place certainly lay further east, no Capelin
roe having been observed during the said year, either
off Vadso or at the other two places in Varanger Fjord
examined by Sars. At this stage the fry were trans-
parent as wafer, with a very thin body, edged with
a transparent vertical fin, and with a shapeless, broad
head, furnished with large, silvery eyes. They were
found in very great numbers, swimming about at the
surface. They were extremely sensitive, and died very
soon after leaving the water, however carefully they
were handled. Afterwards Capelin fry were repeatedly
observed by Sars, even far up Varanger Fjord; and
at the beginning of July they proved to compose the
principal food of the Coalfish. The Cod has also been
found by Collett to have its stomach full of Capelin
fry. When one year old, according to Collett, the
Capelin is about 1 dm. long.

Capelins large and small have thus to serve as
food for the fishes which give rise to the greatest
fisheries of the world, both in Norway and Newfound-
land. The Capelin allures them in its train to the
fishing-grounds; but when if has spawned and again
returns to sea, it entices them away. It is the best
bait that can be used for Codfish in general. As hu-
man food it does not find favour with fastidious pa-
lates, on account of its unpleasant smell; but the Green-
landers dry it, and in this condition it is one of their
most important foods. It is taken with hoop-nets and
seines. In Finmark, according to Juel, the Capelin
seine has 38 meshes to the aln (2 ft.), i. e. a mesh
1 6 Va mm. in diameter.

The food of the Capelin consists mainly of small
crustaceans. It finds a plentiful diet in the Arctic

a U. S. Comm. Fish, and Fisheries, Rep. II (1872 and 1873), p. 225.

b The above-mentioned observations of Pallas, however, point to this conclusion: and in consideration of the great density of the
Capelin shoals even out at sea, Juel remarks (1. c., p. 4), “The greater number of the Capelins thus spawn probably in deep water.”

882

SCANDINAVIAN FISHES.

seas, with their wealth of Schizopoda ; Hyper idee, and
Copepoda, the multitudinous minute animals collectively
known by the Norwegians as Kril. It often seems,
however, as Fabhicius remarked, to live on its own
roe as soon as deposited; even in the females we have
found the stomach crammed with eggs similar to those
dropped from the oviduct and still left in the abdo-
minal cavity".

In the Glacial clay the Capelin occurs'' in the
same way as Gadits saida (see above); but the ma-
trices are usually still more characteristic, following in
their outer contours the shape of the Capelin, and still
commoner. They are found in scattered localities
throughout Norway, from the extreme north to the

neighbourhood of Christiania, sometimes 200 feet, if
not more, above the level of the sea.

A fish of such economical importance has, of course,
received many names from the fishermen, who in their
undiscriminating interest imagine that they increase
our knowledge of the species by distinguishing between
a number of forms. From Finmark Sparre-Schneiderc
enumerates the Lodde, Vaslodde, Havlodde, Fjordlodde,
Blanklodde, etc. From olden times different names have
been given to the sexes; and even in 1882 Sparre-
Schneider found it impossible to convince the fisher-
men that the male and the female belong to the same
species. The male has been known as the Jernlodded and
Fackselodde e, the female most commonly as the Sildelodde.

Genus THYMALLUS.

Teeth in the mouth scattered and small, set in a single row on the inter maxillaries, the maxillaries, and in the

lower jaw, in a small card on the head’ of the vomer, in
with age on the tongue, cardiform on the pharyngeals.
jaw 66 — 75 %f of that of the head reduced. Length
the body and greater than that of the head reduced.

20 — 30). Scales middle-sized, less than 1009

The Grayling genus, Thymallus, occupies, as we
have already mentioned, in many respects a remark-
able intermediate position between the Salmons and
the Gwyniads. But in the transition to the Salmons
there is another connecting-link, the genus Brachy-
mystax'1, which in comparison with Thymallus, has

two rows on the anterior part of the palatines, disappearing
Length of the maxillaries 38 — 48 %, and of the lower
of the base of the dorsal fin more than 15 % of that of
Pyloric appendages well- developed and numerous ( about
(75 — 93) in the lateral line , which is complete.

stronger teeth, persistent on the tongue, smaller (more
numerous) scales, and a shorter dorsal fin (with at
most 14 rays). Apart from this genus, and also ex-
cluding Oncorhynchus from the comparison, the said
intermediate position of the Graylings is best expressed
by the following relations:

a Positively to determine the species of eggs contained in the stomachs of specimens that have lain for years in spirits, is a task
we will not undertake; but the resemblance to the eggs of the Capelin itself is striking.

h M. Saks: Foss. Dyrel. Qvartcerper. Uuiv. Progr. Chrnia 1864, p. 25. Collett, Glac. Mergelb. fra Bejeren, TromsO Mus.
Aarsb. III.

c Zool. Iagttag. fr. Vardo, Thoms0 Mus. Aarsb. 1882, p. 23.

d Really Jarlodde or Jadrlodde , i. e. Edged Capelin, from Jadar, Jar, or Jeer (e. g. in Jsederen), Engl, edder , Sw. gjcirde.
e Old Norwegian Fax, mane, fringe.
f Exceptionally (in young specimens) 76 %.

3 Thymallus grubei, var. baicalensis, according to Dybowski (Verh. Zool. Bot. Ges. Wien, Bd. XXIV (1874), p. 391), has 92 — 108
scales in the lateral line.

h Gthr, Cat. Brit. Mus., Fish., vol. VI, p. 162. To this genus should probably be referred both the Siberian Salmo coregonoides,
Pall, and the Dalmatian Thymallus microlepis , Steind. ( = Salmo obtusirostris, var. oxyrhynchus , Steind.). Both these species, however,
instinctively suggest the possibility of hybridism — the former = Thymallus + trutta, the latter = Thymallus + fluviatihs. Cf. Smitt: Riksm.
Salmon., p. 199, not. 4.

GRAYLINGS.

883

Aver a g e.

Salmon.

Grayling.

Gwvniad.

Length of the maxillaries _ in % of the length of the head

39

31

27

,, - „ „ „ „ ., ., „ reduced

59

42

37

n ?? t, lower juw — — ?? n n n ?? 5? n head

59

52

42

„ ., .. „ - „ •, •• „ „ „ reduced

90

60

56

Interorbital breadth of the forehead ... „ „ „ „ „ ,, „ „ „

47

44

41

The Graylings’ closest approach to the Salmons appears in the following relations:

Average.

Gwyniad.

Salmon.

Grayling.

Number of rays in the anal fin

15

11

12

Base of the anal fin in % of the length of the body

11

8

9

Distance between the adipose fin and the caudal fin ,, ,, „ ., ,, ., „ .,

9

11

11

» „ » anal ,, „ „ „ „ - „ „ „ „

9

12

11

To the Gwy niacls the Graylings come nearest in the following relations:

A v e r

g e.

Salmon.

Gwyniad.

Grayling.

Postabdominal

length

in % of the

length of the body

20

25

26

Height of the

dorsal fin

11 ?? 11 11

13

15

15

Length of the

ventral fins

11 11 11 11

12

14

14

n n

middle caudal rays

11 11 11 11

7

5

5

n n n

suboperculum

'll 11 11 11

„ „ „ head

25

29

29

11 11 .5?

head reduced

- 11 11 11 11

11 11 11 11

67

74

73

In the size and texture of the scales the Graylings
also stand nearer to the Gwyniads, and partake, to-
gether with the latter genus, in the approximation of
the Sal monoid type to the Cyprinoid.

The development of the dorsal tin in the Gray-
lings, their most distinctive character, has the result
that the average distance between this tin and the
snout is only about 35 % of the length of the body.
In this respect Thymallus stands alone in the Sal-
monoid family. The opposite extreme we find in Mal-
lotus, where the said average is more than 50 % of
the length of the body.

The name of Thymallus (Gv/uaA/.og) dates from
.Elian — about 120 A. D. — and has reference to the
thymy smell which some have supposed the fish to
emit. In Linnaeus it was a specific name; Cuvier
raised it to a generic rank".

Excluding the two nominal species Tliym. ontari-
ensis and Tliym. Mertensii, which have been recognised
by Valenciennes* 6 alone, and which have already been
repudiated by Gunther*, and also excluding the ob-
scure Tliym. Grubei, Dye. and Tliym. brevirostris, Ivessl.
from Southern Siberia and Central Asia, there hardly
remain more than two species of the genus Thymallus
that can lay claim to recognition. Of these two the
North Siberian Tliym. arcticus ( Pallasii)d is especially
remarkable as representing in by far the most respects
the characters of youth and of the females, while in
others — even in the most distinctive character of the
genus, the length of the base of the dorsal fin — it
has attained a more advanced development than our
common species,

a Regne Animal , ed. 2, tome 2, p. 306.

6 Cuv., Val., Hist. Nat. Poiss., vol. XXI, pp. 452 and 453.
c Cat. Brit. Mus., Fish., vol. VI, p. 200.

d Thymallus signifer , Richardson, from North America, is probably identical with this species, though for the present we must leave
this question open. See Smitt, Riksm. Salmon., p. 206.

884

SCANDINAVIAN FISHES.

THE GRAYLING (sw. harren).
THYMALLUS VULGARIS.

Plate XLII, fig. 1.

Number of gill- rakers on the whole of the first branchial arch more than 21a, on the lower part of the arch
more than 13h. Length of the head more than 61 % of the preabdominal length. Length of the snout more than
24 % of that of the head or than 32 % of that of the head reduced , and the breadth of the snout more than
74 % of the length of the maxillaries. Least depth of the tail less than 42 % of the length of the head or than

57 % of the length of the head reduced.

R. br. 10 c; D. — —(=17—22); A. -——(=10—14);

13 — 1 6V ' 8— ir

- — - ; V. — ; C . x -f- 1 -f- 1 7 -)- 1 + x ;

(11)13 — 15(16) (9)10—11

Lin. Icit. (65)74 — 93; Lin. tr. — 1 (supra pinn. ventr.); Vert. 61'

Syn. Qvga'/J.og, 7Elian., De nat. anim ., lib. XIV, cap. XXII; Thy-
rnallas, Belon, A 'at., Divers. Poiss., p. 276; + Umbre de
riviere, ibid., p. 280; Thymus , Rondel., De pise, fluv., p. 187
+ Umbra fluviatilis, ibid., p. 172; Gesn., De Aquat., pp.
978 et 1032; Willughb., Hist. Pise., pp. 187 et 188. Core-
yoniis maxilla superiore longiore, pinna dorsi ossiculorutn vi-
ginti trium, Art., Ichthyol., Gen. Pise., p. 10; Syn. Pise.,
p. 20; Descr. Spec. Pise., p. 41; Lin., Fna Suec., ed. I,
p. 119. Trutta, No. 15, Klein, Hist. Pise. Nat., Miss. V,
p. 21, tab. IV, fig. 5.

Sahno ( Coregonns ) Thymallus , Lin., Syst. Nat., ed. X, tom. I,
p. 311; Bl., Fisch. Deutsclil., part. I, p. 158, tab. XXIV;
Retz., Fna Suec. Lin., p. 349; Couch. ( Coregonus ), Hist.
Fish. Brit. Isl., vol. IV, p. 280.

Thymallus vulgaris, Nilss., Prodr. Ichthyol. Scand., p. 13;
Yarr., Brit. Fish., ed. 2, vol. II, p. 136; Kr., Damn.
Fisk., vol. Ill, p. 35; Lloyd, Scandinav. Advent, p. 127;
Nilss., Skand. Fna, Fisk., p. 447 ; Wdgrn, Landtbr.-Akad.
Handl. 1858, pp. 180 et 207; Sieb., Sussivasserf. Mitteleur.,
p. 267 ; Mgrn, Finl. Fisk fna , (disp. Helsingf.), p. 64: Nystr.,
Ialctt. Fnan Jemtl. vattendr. ( disp. Dps. 1863), p. 15; Wdgrn,
Landtbr.-Akad. Tidskr. 1863, pp. 202 et 203; Gthr, Cat.
Brit. Mus., Fish.,, vol. VI, p. 200; Canestr., Fna ltdi.,
Pesci, p. 23; Lunel, Hist. Nat. Poiss. Bass. Lem., p. 120,
tab. XIII; Coll., Vid.-Selsk. Forh., Chrnia 1874, Tilliegsh.,
p. 171; ibid. 1879, No. 1, p. 91; Olss., Vet.-Akad. Ofvers.
1876, No. 3, p. 135; 1882, No. 10, p. 50; Fedders., Nat.
Tidskr., Kbhvn, ser. 3, vol. XII, p. 78; Mor., Hist. Nat.
Poiss. Fr., tom. Ill, p. 543; Bncke, Fisch., Fischer., Fischz.
0., IK. Preuss., p. 153; Mela, Vert. Fenn., p. 345, tab. X;
MOb., Hoke, Fisch. Osts., p. 129; Day, Fish. Gt. Brit., Irel.,
vol. II, p. 131, tab. CXXIV; Norb., Handl. Fiskev., Fiskafv.,
p. 400, fig. 120; Smitt, Riksm. Salmon., Vet.-Akad. Handl.,

Bd. 21, No. 8, p. 198; Lillj., Sv., Norg. Fisk., vol. II,
p. 664; Mela, Sundm., Finl. Fisk., tab. XXV.

Thymallus vexillifer, Agass., Poiss. d'eau douce, tab. XVI,
XVII, XVII* 6; Cuv., Val., Hist. Nat. Poiss., vol. XXI, p.
438; Hckl., Kn., Sussivasserf. Oesterr. Mon., p. 242; Fatio,
Fne Vert. Suisse, vol. V, p. 286.

Thymallus gymnothorax, Cuv., Val., 1. c., p. 445, tab. 625;
Gthr, Jahresb. Ver. Vaterl. Naturk. Wiirteinb., Jahrg. IX
(1853), p. 341; Rapp, ibid., Jahrg. X, p. 161.

Thymallus gymnogaster, Cuv., Val., 1. c., p. 446, tab. 626
+ Thym. FEliani, p. 447.

The Grayling attains in Sweden a length of 5 or
6 din. and a weight, at least in ordinary cases, of at
most 2 kilo/ The form of the body is fairly elongated
and compressed. The greatest depth, which occurs at
the beginning of the true dorsal tin, is about a/5 (18
— 21 %), but sometimes more than 1/4 (26 %), of the
length of the body, and the greatest thickness in old
specimens is about 46 % of the said depth; but during
youth, at a length of 3/4 dm., when the dark transverse
bands are still present, the body is more terete, the
greatest thickness being about. 56 % of the greatest
depth. The least depth, just in front of the caudal fin,
is about 6 — 71/ 2 % (seldom 8 %) of the length of the
body. The dorsal and ventral profiles are as a rule
fairly similar to each other, the back rising from the shal-
low (seen from the side, pointed) snout to the beginning
of the dorsal fin in a curve equal to that in which the
ventral profile descends for the same distance, and be-
hind this point the depth of the body decreases with
fair uniformity. But when the ventral muscles are
powerfully contracted, it often happens that the anterior

a In young specimens 21 or 20.

6 In young specimens 12.
c 9 — 11, according to Lilljeborg.
d 60, according to Kiidyer.

e From the Governments of Westernorrland, Jemtland, and Westerbotten it was indeed reported to the Fisheries Commission of 1881 —
83 that the Grayling there attains a weight of 8 — 10 Sw. lbs. (31 /3- — 4J/4 kilo.); but the greatest length adduced at the same time, 1'7 Sw.
ft. (5 dm.), seems hardly to admit of such a weight, except perhaps in the case of gravid females.

885

GRAY

curve of the back becomes more elevated, while the
ventral profile 'is rendered more or less straight. In
front the dorsal margin is broadly convex or, on the
head, even fiat; but towards the beginning of the dorsal
fin it gradually becomes more or less compressed; be-
hind the dorsal fin it is again flattened. The ventral
margin in front of the anal fin is broadly convex, at
the ventral fins even fiat; behind the anal fin it is flat,
like the dorsal margin of the peduncle of the tail. The
sides of the body are slightly convex.

The length of the head from the articular knobs
of the maxillaries is about equal to the greatest depth
of the body, but varies conversely to the latter, from
about 21 to 20 % of the length of the body in young
Grayling, and from 20 to 18V2 % thereof in old. The
head is wedge-shaped, with compressed or only slightly
terete cheeks. The most characteristic feature in its
appearance consists in the shallow (sometimes even sharp)
and broad tip of the snout, which in a horizontal di-
rection is truncate or slightly curved, but is occasionally
somewhat prolongated like a beak, if only to a short
distance, and which, when the mouth is closed, always
projects beyond the point of the lower jaw. The eyes
show the peculiarity that the pupil, as in the Gwyniads,
is anteriorly somewhat pointed, at an angle. Their size
varies with age, the longitudinal diameter being in
young specimens about 29 %, in old about 16 %, of
the length of the head. The nostrils are set somewhat
nearer to the eyes than to the tip of the snout, the
anterior on each side being raised in a tubular form,
the posterior, which lies just behind, resembling a trans-
verse slit, covered by a dermal fold from in front. The
lips are quite flesh}% especially on the intermaxillaries;
and the form of the broad, but short gape is essentially
determined by the said bones. These bones are reduced
each to a thin, triangular disk, transversely set and
consisting principally of the articular process, only the
lower, toothed margin, which is also the true corpus
of the bone, being somewhat thickened. The maxil-
laries are terete in the anterior, inward part (the ar-
ticular process); but behind this point they are disk-
shaped, curved like a sword, but of fairly uniform
breadth. The teeth extend along about the middle
third of their inferior margin. The oblong jugal bone
is about 2/3 as long and 1/2 as broad as the maxillary.
The length of the maxillaries is less than in any pre-
ceding Salmonoid genus, varying between 35 and 27 %
of that of the head, or between 48 and 38 % of that

f LING.

of the head reduced. Together with the jugal bone
the maxillary bone varies in breadth between about
v4 (in the young) and 2/s (hi old specimens) of its
length. The lower jaw is deep at the middle, but at
the broad anterior margin shallow, almost sharp. Its
length is about 54 — 48 % of that of the head or 76
— 66 % of that of the head reduced. The distribution
of the teeth in the mouth and pharynx we have al-
ready touched upon. The teeth are all alike, of a
blunt conical or cylindrical shape, in contradistinction
to the pointed and more or less curved teeth of the
preceding genera. The characteristic features of the
gill-rakers we have also noticed. In the upper jaw
we find a well-developed transverse fold (palatal cur-
tain), partially concealing the vomerine teeth, behind
the intermaxillary teeth. The lower jaw is not with-
out a similar fold, but this is very low. The pseudo-
branchiae are rather large, the length of their longest
lamella (at the middle) being about equal to the ver-
tical diameter of the pupil. The gill-openings are
large, extending from the middle of the upper oper-
cular margin down to the isthmus in about a line with
the anterior margin of the eye, where they cross each
other for a little distance, the left fitting into the
double fold of the right, as we have observed in Mal-
lotus. All the branchiostegal rays are fiat, the anterior
(lower) curved in an ensiform shape. Behind they
gradually increase both in length and breadth. The
operculum is obliquely quadrangular, with the upper
posterior corner rounded, the posterior margin more
or less concave, and the inferior margin obliquely
ascending in a backward direction along the suboper-
culum, the length of which measures at this margin
about 26 — 31 % of that of the head, and the breadth
(height) of which is about half that of the operculum.
The lower posterior corner of the suboperculum is
sharply rounded. The hind margin of the preoper-
culum is broad, and its lower posterior corner almost
rectangular.

The dorsal fin begins at a distance from the tip
of the snout (the articular knobs of the maxillaries)
that measures on an average 36 — 34 % (exceptionally
38 or 33 %) of the length of the body. The average
length of its base, which increases even relatively with
age and is greater as a rule in the males than in the
females, varies between about 18 and 28 V2 0//° (hi old
males exceptionally 25 %) of that of the body. Its
shape shows great variations. Sometimes (especially

886

SCANDINAVIAN FISHES.

in young specimens and the females) it is of fairly
uniform height or highest in the anterior part (the
6th — 8th ray), the height being about 11 — 14 % of the
length of the body. Sometimes again (at least in old
males) the posterior part of the tin (the 16th — 21st ray)
rises to a height of as much as 7 4 of the length of
the body. The adipose fin is of the same shape as in
the Salmons, forming a curved lobe of fairly uniform
breadth. It lies nearly vertically above the end of the
anal fin, the distance between it and the first upper
supporting ray of the caudal fin being about Vs — V9
of the length of the body. The anal fin also resembles
that of the Salmons. It begins at a distance from the
tip of the snout (the articular knobs of the maxillaries)
which measures on an average about 71 % of the length
of the body. The length of its base is 8 — 9 % (ex-
ceptionally 7 or 11 %\ and its greatest height (the
length of the first or second branched ray) during youth
about 8 — 10 %, in old females about 10—11 %, and in
old males sometimes 1 3 1 / 2 %, of the length of the body.
Its hind inferior margin is convex. The caudal fin is
deeply forked, with pointed lobes. Its middle rays,
the relative length of which decreases with age from
about 7 to 4 % of the length of the body, always
measure less than half, sometimes only one-third, of
the length of its longest rays.

The pectoral fins are set low, with almost hori-
zontal base. They are obliquely pointed, and their
length is about 14 — 17 % of that of the body. The
ventral fins are as a rule somewhat shorter, but some-
times even longer, than the pectoral. They are also
broader and more obtuse. The distance between them
and the tip of the snout (the articular knobs of the
maxillaries) is about 46 — 48 % of the length of the
body. The preabdominal length is on an average 28 %,
the postabdominal length 26 %, of the length of the body.

The scales are most like those of the Cyprinoids,
both in texture and shape. They are not only larger
(fewer), but also thicker than those of the Salmons,
and are furnished with radiating grooves. These are
especially deep in the anterior (inserted) part of the
scale, to which part they give an undulating appearance,
and the truncate margin of which they render sinuate,
forming 3 — 5 indentations. They are very short at the

posterior, rounded margin of the scale, where they leave
only slight notches. The lateral line follows a fairly
median course, in front somewhat nearer to the back
than to the belly. The middle part of its scales pro-
ject. at the inserted margin in a rounded form. The
ordinary scales cover the whole body, but leave most
of the fins, the head, and a greater or less portion of
the breast naked". The base of the caudal fin is, how-
ever, clothed with smaller and thinner scales, and this
covering follows the caudal rays, especially the longest
ones, the scales here growing gradually smaller outwards
and being of a linguiform shape, calling to mind in their
structure the scales we have seen above in Mall of us.

Its varied and shifting colours render the Gray-
ling a handsome fish. Dark and light phases of colour
follow with different environments, and the Grayling,
like so many other fishes, has a particular festal dress.
The ground-colour of the back is brown or grayish,
shading into blue, green, or a purplish lustre in dif-
ferent lights. The sides are lighter, those of the head
with a metallic lustre, those of the body with a silvery
lustre or a tinge of yellow. The belly is of a more
or less pronounced silvery white. Along the sides of
the body, between the back and belly, run a number (up
to 16) of dark, parallel stripes, marking the limits be-
tween the regularly arranged rows of scales; and be-
sides there lie sharply defined, dark spots, reminding
us of the coloration of the Salmons, irregularly scat-
tered on the sides of the head and varying in occur-
rence, but sometimes distributed in longitudinal rows,
especially on the forepart of the sides. The fins of the
ventral side are yellowish or ashy gray at the base
(sometimes throughout), towards the top (especially the
posterior ones) violet. The caudal fin, as well as the
adipose fin, is bluish gray, violet, or even of the same
dark hue as the back. Most prominent is the coloration
of the large dorsal fin, which is of an iridescent violet
or lake, with three or four rows of dark, quadrangular
spots, generally ocellated in lake, on the fin-membrane
between the light rays. In the spawning-dress all these
colours are heightened, and the body gleams with a
golden lustre.

The peritoneum is white. The short oesophagus
passes into a very thick-walled stomach* * * 6, which extends

" At the median line the belly is usually covered with scales (even if these are only small) forward to a line with the insertion of

the pectoral fins; but on each side and in front throughout the whole ventral side the skin is naked. The variations in the extent of this

scaleless part led Valenciennes to establish the species gymnothorax and gymnogaster.

6 In a male 55 cm. long the wall of the stomach is 6 mm. thick.

GRAYLING.

887

to about a line with the anterior extremity of the pel-
vic bones, where it bends abruptly forward, so that the
end of the pylorus lies in about a line with the ter-
mination of the first third of the pectoral fins when at
rest. The anterior bend of the intestine with its nume-
rous pyloric appendages runs upwards and slightly to
the right, but turns with equal abruptness just behind
the diaphragm, whence the intestine proceeds without
further curvature to the vent. Almost exactly in a line
with the insertions of the ventral tins, the intestine
passes into the rectum, which is lined, as in the Sal-
mons, with large, annular, transverse folds. The liver
is comparatively small, and lies only in the left side of
the body. It extends back hardly to a line with the
tips of the pectoral tins when folded. The spleen lies
obliquely to the right of the stomachic bend and behind
this point, directed towards the left. Its size varies, as
in the Salmons, according to the greater or less deve-
lopment of the generative organs". The air-bladder is
thin-walled, but large, extending along the whole dor-
sal margin of the abdominal cavity.

The Grayling is strictly a mountain fish, and pre-
fers clear, rapid streams and their dead water, with a
bottom of sand or gravel. The best place to look for
it is, therefore, in waters where falls alternate with
deep pools or broads. But it also roves into lakes, and
is found even in the sea, where the water is brackish
and of a temperature suited to its requirements. As a
general rule we may say that, where the common Trout
can thrive, the Grayling is also at home; but they are
seldom met with in company. The geographical range
of the Grayling also embraces the greater part of Europe.
Whether it occurs in Asia, and how far its range ex-
tends there, is as yet uncertain, for the Graylings brought
home from the Yenisei by Nordenskiold’s expedition
in 1876, proved to belong to the closely related species
which Pallas once named Salmo arcticus , and which is
probably identical with the North American Grayling.

In Europe the Grayling is most plentiful in Lap-
land, the alpine tracts of Central Europe, and England.
According to Lvestadius’ manuscript (1831) it is fairly
common in all the rivers and lakes of Tornea Lapp-
mark, up to the extreme north of Sweden (Kilpisjarvi,
69° N.). According to Fjellner (MS in the Royal Mu-

seum) it is plentiful in Juckasjarvi. It seems to occur
more sparingly in the lower parts of the Lapland rivers.
In Jemtland, according to Olsson, it goes up to the
Norwegian frontier. In Lake Fla and in Stroms Vattuclal
it is an important food-fish. The Harr Lakes (harr =
Grayling) on the Fulafjall in Northern Dalecarlia derive
their name from its abundance in their waters; and
throughout the Dal Elf it is fairly common. It also
occurs in the Klar Elf, most plentifully in its upper
(northern) parts and in the lakes on its course. Accord-
ing to IIogberg’s notes in 1835 (MS in the Royal Mu-
seum), a certain form is here known as Sandharr, which
appears to be a small Grayling about 22 cm. long.
According to Norback a distinction is drawn between
the Flodharr (River Grayling), which is short and thick,
with convex back and of dark coloration, and which
prefers comparatively still water, and the Stromharr
(Stream Grayling), which is longer and more slender, of
more slender frame and lighter coloration, and which
frequents falls and the water beside them. Widegren as-
signs the Grayling to Lake Wener. From Lake Wetter
the Royal Museum received this fish through Hall in
1832, and Arosenius forwarded specimens in 1834 from
the Motala River off Norrkoping''. Further south the
Grayling is rarer in Sweden; but it occurs in Smaland
and in HallamT’, at least in the Laga River, where it
is known, according to Trybom, as the Haspa.

In Norway its range is no less extensive, but ap-
parently still more sporadic. South of the Dovre Fjeld
it is found, according to Collett, only in the waters
of the Oster Valley and the Gudbrand Valley, but there
is rather plentiful. Along the west coast- it is wanting,
but finds its way through the Lesjeverks- water on the
Rauina Elf almost to the mouth of this river in Runs-
da-1 Fjord. From this point north to Finmark the Gray-
ling is known only from some waters in the parish of
Lierne (North Trondhjem), in the Vefsen Elf (Nordland),
and in the valley of the Maa-1 Elf. It is common, Iioav-
ever, in many, though not in all, of the rivers in Fin-
mark. According to Mela it is common throughout
the greater part of Finland, but Avanting in the south-
ern districts, Avhere it occurs but sparingly in Nyland
alone. ToAva-rds the head of the Gulf of Bothnia it is
common, both inside and outside the island-belt; but

“ Kh0YER found it small and round. In a male 55 cm. long, with testes hardly ripe, it is oblong, rather blunt behind, pointed in
front, and 58 mm. long.

b Smitt, liiJcsm. Sahnonul., tab. metr. VIII.

0 Underd. Bel. Ny Fiskeristadga, 1883, p. 155. See also Lilljeboug, 1. c., p. G74.

Scandinavian Fishes.

112

888

SCANDINAVIAN FISHES.

in the central and southern parts of this gulf it is

o

rarer, occurring down to Aland and the islands oft'

o

Abo. In Denmark, where the Grayling is called Stal-
ling, it occurs in a number of rivers from the south
to the middle of Jutland. According to Feddersen,
however, its numbers are there rapidly decreasing, an
observation which has also been made in other coun-
tries, for example in North Germany.

In European Russia, according to Grimm®, the
Grayling is met with in the small rivers, and the upper
parts of the large rivers, that flow into the Arctic Ocean,
the White Sea, the Baltic, the Black Sea, and the Cas-
pian. In Central Europe, where the Grayling has the
centre of its geographical extension in the Alps, it oc-
curs, according to E. Sciiultze6, in the basins of the
Danube, Rhine, Weser, Elbe, Oder, Vistula, and Pregel.
Westward its true range extends over Switzerland to
the mountainous tracts of Eastern and South-eastern
France. Ausonius (about 400 A. D.) knew the Gray-
ling from the Moselle0; and according to Selys-Long-
CH amps d it is common, at least in some of the small
streams, in the mountainous interior of Belgium. On
one or two occasions it is said to have been found in
Holland0. Southward the Grayling goes to Lombardy,
Venetia, Piedmont, and Istria (Canestrini, 1. c.). In
England the Grayling probably has its original home
to the west/ and north and in Wales; but at the pre-
sent time, according to Day, it is commonest in the
more southern rivers, the Avon, lichen, and Test. It
has been introduced in recent times into several Scotch
waters. In Ireland it is wanting.

Even old writers, at least from Gesner’s time, were
fairly well acquainted with the habits ot' the Grayling,
and had learnt to compare them to those of the Trout.
At most times it lies stationary and alone, in the shelter
of some jutting stone or in the shade of an overhanging
tree; but it darts oft' like an arrow, t\4i enlightened,
and is as speedy to seize any victim that comes its
way, often leaping out of the water to caj^Gjre an in-
sect. But sometimes it shows sociable tendencies, and
in Juckasjarvi, according to Fjellner, it assembles in

shoals during June and July. Small cascades it easily
surmounts by leaping, but it is deficient in the Salmons’
power of passing high falls. Large Grayling keep to the
deepest part of the water, the smaller ones stay nearer
shore.

The spawning-season usually occurs soon after the
breaking up of the ice, and may thus vary in Sweden
between April and June. In the lakes of Lapland, how-
ever, it sometimes happens, according to Trybom8’, that
the breeding Graylings repair to the mouths of brooks,
where there is open water some time before the ice
breaks up. In southern countries the spawning begins
earlier: in Italy, according to Canestrini, from January
to April, v. Si e bold describes in the breeding fish a
firm, but smooth, dermal eruption on the scales, whose
limits, however, remain distinct. This eruption is want-
ing in the sterile individuals, which are further di-
stinguished by a paler coloration. The Grayling too
discards its ordinary caution during the spawning-season.
According to Heckel and Inner the fish swim in pairs
to the spawning-place, where they scoop a hollow for
the eggs with their caudal fin, the eggs being covered
after impregnation with gravel or pebbles; and where
the spawning has taken place in March, the ova are
hatched in June7'. According to Fjellner live roe is
found between the stones in the rapids of the Torne
Elf in July, and in August the fry swim about in the
river. The diameter of ripe eggs, lying loose in the
abdominal cavity, ready to be deposited, we have found
to be 2 V4 — 2 Va mm. But Benf.cke assigns to them a
size of about 4 mm., presumably when they have swell-
ed after impregnation. According to Norback the living
female contains 5,000—13,000 ova to every pound of
her own weight. The fry grow rapidly; in England,
according to Day, they are 4 or 5 inches long at the
end of July or the beginning of August. But the
Grayling, it is stated, does not attain maturity until
three or four years old.

In spite of its feeble dentition the Grayling is di-
vstinctly a fish-of-prey; and it is no dainty feeder.
Worms and the larva; of insects, mollusks, crustaceans,

“ Fishing and Hunting on Russian Waters, p. 12.
h Geogr. Verbr. Biisswasserfisch. Mitteleur., Stuttgart 1890,
c Mosella, 1. 90.

'' Faune Beige, p. 22.

e In the little river Geul, according to v. d. Ende, Versl. Werkz. Vereenig. Bevord. Fill. Iehthyol., Deel. I, p. 39.

■T “The very best stream in the world for Grayling is the Teme” (a tributary of the Severn), according to Bullock, see Buckland,
Nat. Hist. Brit. Fish., p. 331.

,J Nordisk Aarsskrift for Fiskeri, l:ste Aarg. (1883), p. 303.
h According to Day the fry are excluded in 12 — 25 days.

WIIITE-SALMON.

889

and small fishes, in short nearly all the animal life offered
it by the water where it lives, enter into its diet; and
in its greedy haste it swallows leaves and fir-cones that
fall into the water, mistaking them for insects. It seems
to be especially partial to caddis-worms (the larvae of
Stone-flies), the coverings of which are generally to be
found in its stomach. But mollusks are also one of its
favourite foods; in a male 55 cm. long the stomach was
full of entire shells, an inch long, of pond-snails ( Limncea ).
Fishes of the Minnow’s size fall a prey to the full-grown
Grayling. Grayling large and small are besides roe-
eaters. When the Salmons spawn, they keep watch,
and feed on the ova; but in requital they are harassed
by the Salmons, in whose maw many a Grayling has
perished. This probably explains how, where food is
plentiful, Grayling and Trout can thrive in company,
but where food is scarce, the former must give way.

As the habits of the Grayling in essential respects
resemble those of the Trout, the same fishing methods
are used for them both. The Grayling, like the Trout,
readily takes a fly, and in many places gives the fly-fisher
good sport; but owing to its weak jaws a certain amount
of care is necessary not to tear the hook out and lose
the fish. Less fastidious anglers use a bait of worms
and small fishes, for example Minnows. But the Gray-

ling is taken most commonly and in the greatest quan-
tity with net and seine.

The flesh of the Grayling has always been held in
esteem. It is white, of good flavour, and easy of di-
gestion, suitable for even weak stomachs. It is there-
fore set higher even than that of the Salmon. “E'm
Ac sell ist ein Kheingraf, ein Salm isf ein Herr," is an
old saying to be found in Gesner. The Grayling is
best in autumn and winter, worst, of course, just after
the spawning. Not only its flesh has enjoyed a good
reputation; in ancient medicine its fat. ( oleum Aeschice)
was widely employed. When the fish is in good con-
dition, the whole intestinal canal is embedded in rich
fat, and the oil extracted from this has been used espe-
cially in eye and ear diseases and to cure cutaneous
diseases and burns. Linnaeus states that, the Lapps
employ the gastric juice of the Grayling in the prepara-
tion of cheese, to curdle the reindeer milk, simply laying
the whole intestinal canal in the milk.

Besides Ha spa, a name which may perhaps be
explained by the use of Asp among the Swedish fisher-
men both for large Cyprinoids and large Gwyniad-
fishes, Nilsson and Lilljeborg mention from Lake
Wetter Val, and from Norrkoping Oreval, as names
applied to the common Grayling.

Before proceeding to the Gwyniads, we may briefly | near to the limits of our fauna, though it has not yet
mention a. genus whose geographical range approaches : been found in Scandinavia.

Genus STENODUS.

Teeth small and disappearing during growth, hut at first set in two or three rows on the interm axillaries and in
the anterior part of the prominent lower jaw, in a card on the tongue, and in a dense, continuous, two-armed card
on the anterior part of the palate ( the head of the vomer and the front of the palatine hones). Length of the
maxillaries about 50 — 43 %, and, of the lower jaw about 78 — 70 %, of that of the head reduced. Breadth of the
snout across the articular knobs of the maxillaries about, equal to that of the interorbital space, which is about 1 .
(less than 23 %) of the length of the head. Base of the dorsal fin as a rule somewhat shorter than that of the
anal and less than 13 % of the length of the body, but, more than 1/2 of the length of the head reduced. Pyloric
appendages well -dev eloped. Scales middle-sized, about 90 — 110 in the lateral line, winch is complete.

This genus occupies an especially remarkable in-
termediate position between the Smelt and Vendace
groups. In external appearance it so closely resembles
the latter that the confusion between them which has
made its way into literature is easily explainable. But
Stenodus, which becomes as large as the largest Sal-

mons, does not correspond in its characters to the Scan-
dinavian Yendaces until it has attained a size far
greater than that reached by them. In the structure
of the ventral fins Stenodus is approximated to Argen-
tina, the outermost ray, which in the rest of the fa-
mily appears merely as a more or less rudimentary

890

SCANDINAVIAN FISHES.

supporting ray, being here so developed that two
simple rays may with ease be counted in these tins.

The genus is strictly Arctic, belonging both to
Asia and America; but just as the Huch, a species in-
habiting the Danube, has its nearest relation, not to
say identical species, in the Siberian Salmo fluviatilis,

so in the basins of the Volga and Ural there lives a
Stenodus hardly to be distinguished in species from
the Siberian form, which in its turn seems to be
identical with the American. However this question
of species be determined, we are here interested
only in

THE NELMA OR SIBERIAN WHITE SALMON.

STENODUS NELMA.

Fig. 221-

Fig. 221.

Stenodus nelma , from the lower

part

of the Dwina. Taken by Lieutenant H. Sandeberg on the 29th Sept.,
1 4 of the natural size.

1877.

R. hr. (9)10; D. JLJL[i4_l6]; A. —^^[(15)16-18(19)];

L. Icit. 88—109.

Syn. (?) Salmo leucichthys , Gyldenstadt, Nov. Comm. Acad. Sc. Petrop.,
tom. XVI (1771 — 72), p. 533; cett. vide Smitt, Riksm.
Salmonid., p. 207 ; adde Grimm, Fish., Hunt. Russ. Wat.,
pp. 12 et 20.

Salmo Nelma , Pall., Russ. Reis., pt. II, p. 716; cett. vide
Smitt, 1. c.; adde Grimm, 1. c. ; Lillj. (Stenodus), Sv., Norg.
Fisk., vol. II, p. 698.

(?) Salmo Mackenzii, Rich , Narr. Voy., Frankl. 1819 — 22, j
App., p. 707; cett. vide Smitt, 1. c.

(Joregonus dupeoides, Lillj., Vet.-Akad. Handl. 1850, p. 304;
cett. vide Smitt, 1. c.

The Nelma is a fish highly esteemed and much
employed in Russia. As we have already mentioned,
it probably belongs to the same species in South-eastern
Russia as in the Arctic parts of that empire. The
southern form — which is perhaps distinguished by a
somewhat smaller head, less than V5 of the length of
the body — was named by Gyldenstadt Leucichthys,
a translation of the Russian Bjelaja By biz a or Belori-
bitza, the white fish, referring to its coloration. In the

Arctic Ocean it lives like the Salmons, and at the end
of winter, according to Pallas, ascends the rivers in
multitudes. Pallas supposed it to be wanting in the
Yenisei; but Nordenskiold’s Expedition of 1876 brought
home tine specimens from that river. But most of the
Royal Museum specimens are from the Dwina, where
they were taken at Archangel by Lieutenant H. San-
deberg; and to judge by these specimens it appears as
if the spawning took place in spring or summer, for
in those caught at the end of September" the genera-
tive organs were extremely little developed. That it
is as marked a predatory fish as the Grayling, was
shown by one of these specimens, the stomach of which
contained five but partially digested Roach.

As Nilsson has remarked, it is by no means im-
possible that the Nelma may sometimes rove from the
Arctic Ocean to the fjords or rivers of Norwegian Fin-
mark; but it has never yet been found there. Accord-
ing to Grimm the annual take of nelma and leucichthys
in European Russia amounts to about 100,000 pud
(1,638,050 kilo.) and has a value of more than 1,200,000
roubles (£*190,000).

See Smitt, Rilcsm. Salmon ., lab. inetr. VIII.

SA LM0NID2E.

891

Genus COREGONUS.

All the teeth of the mouth , except those on the tongue , as a rule soon disappearing during growth , or soon con-
cealed in the gums , or persistent as scattered , fine , villiform, mobile teeth, or entirely wanting. Length of the
maxillaries 50 ( exceptionally 52) — 32 %, and of the lower jaw 70 ( exceptionally 74) — 40 %, of that of the head
reduced. Breadth of the snout across the articular knobs of the maxillaries perceptibly less than that of the in-
terorbital space , which i is at least 1/ '4 (exceptionally 24 %) of the length of the head. Base of the dorsal fin less
than 15 % of the length of the body, but more than half the length of the head reduced. Pyloric appendages
well- dev el oped. Scales middle-sized, about 80 — 90 (exceptionally about 70- — 110) in the lateral line,

which is complete.

The dental reduction which we have seen in the
preceding genera — in the strength of the teeth in
Mallotus, and both in their strength and number in
Thymallus — has advanced still further in the Gwyniad
genus ( Cor eg onus ) towards the Cyprinoid type. The
direction of the reduction is indeed not the same as
in Thymallus, the teeth on the tongue being persistent,
and sometimes even more developed than in Mallotus.
But else the reduction is a continuation of that which
we have seen in Thymallus, and in their form, the
jaw-teeth, where they are present, come nearest to
those of the Graylings. The maxillaries and the vomer
are always toothless in Coregonus ; but on the inner
(hind) surface of the intermaxillaries and on the an-
terior part of the palatines the Gwyniads proper are
usually furnished with teeth, on the former bones set in
a thin transverse row and mobile as in Mugil, on the
latter firm and pointed, but small. Among the true
Gwyniads wre find only dermal papilla instead of teeth
in the lower jaw; but among the Vendaces small teeth
sometimes occur in the anterior part of the lower jaw,
as well as on the intermaxillaries®. In the pharynx,
however, we find teeth, numerous though small, not
only on the so-called pharyngeals, but also on the
upper parts of the posterior branchial arches6.

The reduction of the teeth is partly compensated,
as usual, by the apparatus composed of the gill-rakers
(fig. 222), which are subject in Coregonus to great
variations, probably connected with variations in the
diet of these fishes. The rule is apparently that, where
the diet consists of large objects, principally fish, the

Fig.

A,

B ,

C,

D,

E,

F,

gill-rakers (45) and branchial lamellae on the first left
branchial arch in a Vendace ( Core-
gonus albula).

,, (60) on the first left branchial arch in a

Felet ( Coregonus cyprinoides).

,, (45) on the first left branchial arch in an

Asp-Gwyniad ( Coregonus aspius).

,, (27)1 on the first left branchial arch in a

,, (25)1 Gwyniad ( Coregonus lavaretus).

,, (20) on the first left branchial arch in a

Polcur ( Coregonus polcur).

Figs. B — F after Widegren.

Smitt, 1. c., tab. metr. VIII, Uoregoni, note to specimen No. 5.
See above, p. 523, note a.

892

SCANDINAVIAN FISHES.

rakers are scattered and short, but that, where the diet
is composed of small objects, principally Entomostraca,
they are finer, denser, and longer, thus forming a more
perfect filtering-apparatus. Now as the diet is com-
monly altered with the age of the Gwyniads, we may
as a rule expect to find a corresponding change in the
gill-rakers". But Gwyniad forms occur, for example
the Siberian pelet (among the Vendaces) and mulcsun
(among the true Gwyniads) or the so-called asp of Lap-
land, which attain a considerable size, but have per-
sistently dense and numerous gill-rakers. Some forms,
again, show an increase with age in the number of the
gill-rakers6. The variations do not therefore follow in
absolute succession the changes of growth. They rather
shows a certain independence in their appearance, an
independence 'which in some forms has rendered them
available as characters, at least in their extremes, espe-
cially as they are accompanied in most cases by an
external character, a difference in the shape of the snout.

By the last-mentioned character two groups have
long been distinguished within the genus Coregonus,
one of them having its best-known representative in
our Vendace, the other in our Gwyniad. To the for-
mer group Agassiz0 gave the name of Argyrosomus,
characterized essentially by the projection of the point
of the lower jaw beyond the tip of the snout, whereas
the form-series of the true Gwyniads advances to a
greater development and protrusion of the tip of the
snout, culminating in the form hence known as the
ndbbsik ( Beaked Gwyniad , Coregonus oxyrhynclnis). The
limit between the two groups it is indeed impossible
sharply to define, on account of the transition forms;
but in its two extremes the altered shape of the snout
is due to a very considerable difference in the form
and position of the intermaxi llaries. We have already
remarked in the case of Thymallus a considerable re-
duction of the intermaxillaries, to small, flat and thin,
triangular disks, set transversely in front of the tip of
the snout and the incurved articular processes of the
maxillaries, slightly thickened at the lower margin
alone, where lie the alveoli of the single row of teeth,
which are directed inwards (backwards), more or less
vertically to the plane of the disk. In the Vendaces
this structure recurs, only that the teeth are as a
rule wanting, the disk is still thinner and rests more

entirely on the front of the inward, terete tip of the
maxillary bone. This part of the maxillary bone some-
times, as in the Scandinavian Vendace, bears outside
the protuberance which articulates with the ethmoidal
cartilage, a separate process, directed downwards (for-
wards), with which the inner (hind) surface of the inter-
maxillary bone articulates, and by means of which the
said bone is raised when the mouth is opened. The
breadth (height) of the intermaxillary bone does not
exceed 2/s °f the breadth of the snout across the ar-
ticular knobs of the maxillaries, and its sharp inferior
margin then forms the osseous framework in the sharp,
transverse margin of the tip of the snout. In the true
Gwyniads the intermaxillary bone on each side of the
snout is higher, the depth of the snout being always
more than 2/5 of its breadth across the articular knobs
of the maxillaries, and more robust, with teeth or ru-
diments thereof in the above-mentioned position. It is
also more firmly articulated with the under surface of
the tip (articular process) of the maxillary bone. This
articulation is formed in the following manner: the
upper margin of the intermaxillary bone is hollowed
into a groove which surrounds the said lower margin
of the maxillary not only in front, but also more or
less far back (outwards) on that part of the latter bone
which forms the upper lateral margin of the mouth.
The intermaxillary bone thus assumes a more or less
vertical position when the mouth is closed; and when
the tip of the snout is prolongated — which often ap-
pears as a change of growth — the under margin of this
bone is turned more and more in a backward direction.

The scales of Coregonus are thin and cycloid, as
a. rule thinner than those of the Grayling, with not very
distinct radiating grooves, though both the anterior and
posterior margins of the scales are generally corrugated
and notched thereby. The scales of the lateral line,
here as in the Grayling, commonly have the anterior
margin elongated to a triangle at the middle.

The internal organs resemble those of the Salmons;
but the pyloric appendages are shorter and still more
numerous (Kroner counted about 200). The perito-
neum is silvery white.

The genus has the same geographical extension as
the Graylings. It is most numerous, and appears in
its most developed forms, in the Siberian rivers, which

a See for example the four Gwyniads, 206 — 350 mm. long, which. are included in Smitt, Riksm. Salmon., tab. melr. X, Nos. 170 — 173.
b See for example Smitt, 1. c., p. 278, the averages for Coregonus Nilssonii and Cor. Wartmanni.
c Lake Superior, p. 339.

VENDACE.

893

most of the forms ascend from the Arctic Ocean, and
in the great lakes of North America. But even in Scan-
dinavia it ranks among the fishes most important from
an economical point of view. The number of the spe-
cies it is as yet impossible to determine, for the vari-

ability of the forms is as great here as among the
Salmons, and has given rise to the establishment of
numerous, merely nominal species.

The name of Coregonus was coined by Artedi"
and has reference to the anterior angle of the pupil.

THE VENDACE ( SAV. SIKLOJAN OR RABBOXEN).

COREGONUS ALRULA.

Plate XLII, fig. 2.

Lower jaw projecting beyond the shadow (sharp) tip of the snout, at which the breadth (height) of the intermaxil-
laries is less them 2/5 of the breadth of the snout across the articular knobs of the maxillaries. Length of the base
of the anal fin less than 14 % of the length of the body. Least depth of the tail less than 8 % of the length of
the body or 45 % of the length of the head, but more than 47' % of the length of the base of the anal fin. Ver-
tical diameter of the eyes more than 71 % of the breadth of the interorbital space, ivhich is less than 30 % of
the length of the head and usually less than the length of the upper jaw, which is more than 1/3 of the length

of the head or V 5 of that of the head reduced.

a: True Sikloja (var. cdbula): distance between the dorsal fin and the tip of the snout usually more than 42 %

of the length of the body, and the postabdominal length less than 02 % of that distance. Gill-rakers on the
first branchial arch more than 40. Least depth of the tail at most 71 % of the length of the base of the
anal fin. Length of the head, even in full-grown specimens, as a rule more than 19 %, and of the head
reduced more than 15 %, of that of the body.

b: Sikvimma (var. vimba): as above, but: length of the head as a rule less than 19 %, and of the head reduced

less than 15 %, of that of the body.

Fig. 22.3. Coregonus vimba, 10 n of the natural size, o71, from Stora Nyckelviken (Lilia Vartan) in the island-belt of Stockholm.

Caught on the 12th Aug., 187 9.

Philos. Iehthyol., p. 72. xo qyj, pupil, and yiovla, angle.

894

SCANDINAVIAN FISHES.

R.br. 7—8; D. - -—(=11— 13); A. — [=13 — 16(17)];

8- -9( 10; ’ 10 — 12(13)L v 'j

P.

1

(12)13 — 15(17) ’

V.

(1)1

9 — 11 ’

C. x + 1 + 17 + 1 + x;

8(9),

Lin. lat. (70)75-90(94); L. tr. ; Vert. 58 «.

Syn. Marena, Schonev., Ichtli. Slesv. Hols., p, 46 (vide Kr., Damn.

Fisk., vol. Ill, p. 93). Vandesius, Sibb., Scot. III., pt. 2,
lib. Ill, p. 26. Goregonus edentulus, maxilla inferiore lon-
g-iore, Art., Ichthyol., Gen., p. 9, Syn., p. 18 (excl. synon.),
Spec., p. 40; Lin., Fn. Suec., ed. I, p. 119. Anims-vimma,
Lin., It. W goth., p. 231. Ooreg. spec. II, Gisler, Vet.-
Akad. Handl. 1753, p. 196.

Salmo Alhula, Lin., Syst. Nat., ed. X, torn. I, p. 310; Retz.,
Fn. Suec. Lin., p. 349; Bl., Schn., Syst. Ichthyol., p. 411;
Pall., Zoogr. Ross. Asiat., tom. Ill, p. 413; Nilss. ( Core -
gonus ), Prodr. Ichthyol. Scand., p. 17; Ekstr., Vet.-Akad.
Handl. 1834, p. 16; Cuv., Val., Hist. Nat. Poiss., vol. XXI,
p. 520, tab. 633; Lloyd, Scand. Advent., vol. I, p. 135;
Nilss., Slcand. Fn., Fisk., p. 465; Sundev., Stockli. L. Hush.-
Sallsk. Handl., II. 6 (1855), p. 81; Wgrn, Landtbr.-Akad.
Hand!., vol. 18 (1858), pp. 180 et 206; Id., Ofvers. Vet.-
Akad. Forh. 1862, p. 591, tab. IX, fig. 1; Id., Landtbr.-Akad.
Tidskr. 1863, pp. 201, 203, 211; Sieb., Susswasserf. Mittel-
eur ., p. 265; Mgrn, Finl. Fiskfn., p. 54; Gthr, Cat. Brit.
Mas., Fish., vol. VI, p. 192; Coll., Vid. Selsk. Forli. Chrnia
1874, Tilltegsli., p. 170; ibid., 1879, No. 1, p. 91; Olss.,
Ofvers. Vet.-Akad. Forh. 1876, No. 3, p. 138; Malm ( Argy -
rosomus ), Gbgs, Boh. Fn., p. 547 ; Fedders. ( Coregonus ),
Naturh. Tidskr. Kbhvn. ser. 3, vol. XII, p. 80; Bncke, Fisch.,
Fischer., Fiscliz. O., W. Preuss ., p. 152; Mela, Vert. Fenn .,
p. 352, tab. X; Mob., Hcke, Fisch. Osts., p. 133; Tryb.,
Nord. Aarsskr. Fisk., 1 Aarg. (1883), p. 295; Norb., Handl.
Fiskev., Fiskafv., p. 407 ; Reut., Sundm., Finl. Fisk., tab.
VI; Smitt, Vet.-Akad. Hand!., Bd. 21 (1886), No. 8, p. 212,
cett. ; Lillj., Sv., Norg. Fisk., vol. II, p. 706.

Salmo Vimba, Lin., Syst. 1. c., p. 311; Retz., 1. c., p. 350;
Ascan., Icon. Rer. Nat., cah. IV, tab. XXIX; Nilss. ( Core-
gonus), Prodr., 1. c.; Cuv., Val., 1. c., p. 514, tab. 632; Nilss.,
Stand. Fn., Fisk., p. 462; Gthr, 1. c., p. 193; Smitt, 1. c.
Salmo Marwnula, Bl., Fische Deutschl., pt. I, p. 176, tab.
XXVIII, fig. 3; Jard., Edinb. Journ. Nat., Geog. Sc., vol. Ill,
p. 4, tab. 1 ; Jen., Man. Brit. Vert. Anim., p. 432.
Coregonus clupeoides , Nilss., Prodr., p. 18; Stand. Fn., Fisk.,
p. 467.

Coregonus vandesius, Rich., Fn. Bor. Amer., pt. Ill, p. 213;
Gthr, Cat., 1. c., p. 194.

Coregonus Willughbii, Jard., Illustr. Scot. Salm., tab. 6; Yarr.,
Brit. Fish., ed. 2, vol. II, p. 146.

Coregonus brevis, Maklin, Ofvers. Finsk. Vet.-Soc. Forh., vol. XI
(1868—69), p. 19.

The Scandinavian Vendace is usually a small fish,
in most cases of the Bleak’s or the Baltic Herring’s size,
seldom as large as the oceanic Herring, and attaining a
maximum length of about 3 dm. But in other places, as
in some of the North German lakes, it grows to a length
of at least. 372 dm. In form and in coloration it re-
minds us both of the Bleak and the Herring. The body

is fusiform but compressed, in the true Vendace — and
among our forms most in the Wetter form — elongated
as in a Bleak, with the greatest depth, at the beginning
of the dorsal fin, about 19 % of the length, but in the
sikvimmab as a rule deeper, with the greatest depth
sometimes 24 % of the length. In the former the great-
est. thickness of the body is about s/5, in the latter
about 3V> °f the greatest depth; but gravid females of
the former approach the proportions of the latter. The
dorsal and ventral margins are broadly convex or even
flat, the former being the broader, but for some dis-
tance immediately in front of the dorsal fin somewhat
sharpened to a faintly marked carina. The upper and
lower contours are similarly arched, except in gravid
females, where the ventral profile, as usual, is more
sharply curved in the abdominal region. The average
least depth of the tail in the true Vendace is about
6 b'a % of the length of the body, 31 % of that of the
head, 42 % of that of the head reduced, or 60 % of that
of the lower jaw. In the sikvimma these averages are
respectively 7V3 %, 37 %, 49 %, and 75 %.

The head is pointed, somewhat more compressed
than the abdominal region, especially below, across the
lower jaw, the cheeks thus converging distinctly in a
downward direction. The frontal profile at first forms
an unbroken continuation of the dorsal arch, but be-
comes straighter, owing to a slight depression above
the eyes. The inferior profile of the head is somewhat
more sharply curved. The length of the head is about
Vs of that of the body, but varies partly with age,
the oldest specimens having, as usual, comparatively
the smallest head, partly in inverse proportion to the
depth of the body, the sikvimma, which has the deepest
body, having the smallest head. The variations run
between 23 and 18 % of the length of the body. The
same rule applies to the variations in the length of
the head reduced, which run between 1 7 1/2 and 13 %
of the length of the body. The decrease in the aver-
ages during growth, a decrease which is generally
uniform, is also expressed in the difference of form
given above between albuia and vimba; and though
this difference is not quite constant, it may still be
employed as a character in the great majority of cases,
if attention be paid to the age of the fish. The eyes
are furnished, as in most of the Thrissomorphs, with a
well-developed, adipose membrane, surrounding the

“ 56 — 58, according to Gunther.

b Sikvimma = Gwyniad-Zarthe, Silcloja = Gwyniad-Bleak.

VENDACE-

895

whole iris, but broadest at the anterior corner of the
eye. The pupil is fairly circular, with only a faint
angle in front. In adult Vendace the eyes occupy with
fair exactness the second quarter of the length of the
head, though the anterior corner of the eye encroaches
upon the first quarter thereof, the average longitudinal
diameter of the eye being thus about 25 %a of the
length of the head, but the length of the snout only
about 21 V2 0/0 °f the same. The vertical diameter is
somewhat less, in albula on an average about 23 %b, in
vimba 22x/2 %, of the length of the head. The breadth
of the interorbital space, which rises only a little above
the upper margin of the eyes, is in albula about equal
to the longitudinal diameter of the eyesc, in vimba
somewhat greater''. The average breadth of the snout
across the articular knobs of the maxillaries is 19 % of
the length of the head. The two nostrils on each side
of the snout are situated as in the preceding species,
somewhat nearer to the orbit than to the tip of the
snout. The hind margin of the anterior nostril is raised
in the form of an obliquely cut tube; the posterior
nostril, which is larger, may be closed from in front
by means of a semicircular dermal flap. The maxillaries
— as they appear externally — are of uniform breadth,
curved like a sabre, and bluntly rounded at the hind
extremity. Their average length from the top of the
articular processes is in albula about 35 %, in vimba
about 34 %, of that of the head, or in the former about
48 %, in the latter about 45 %, of that of the head
reduced. Their breadth is on an average 1/3 of their
length. Their supplementary (jugal) bone is similar in
shape to their lateral part and nearly 3/ 4 of its length,
being elongated to a point in front. The average length
of the lower jaw is in albula about 51 %, in vimba
about 49 %, of that of the head, or in the former about
70 %, in the latter about 66 %, of that of the head re-
duced. The dentition of the mouth and the gill-rakers
(41 — 48, exceptionally 39 or even 38) we have already
noticed. The transverse palatal folds (vela) are well-
developed, especially in the lower jaw. The gill-open-
ings extend along about the last third of the upper
opercular margin and down to about a line with the

anterior margin of the eye, to which point the bran-
chiostegal membranes are free both from each other and
the isthmus, though they cross at the extreme front,
in the same manner as in the preceding genera. The
branchiostegal rays are thin and flat, more or less cur-
ved in a sabre-like form. The operculum is obliquely
quadrangular, with the lower anterior corner most
pointed and most elongated. Back from the articular
process runs a ridge, faintly marked in the bone itself,
but more distinctly indicated by a darker colour. Within
the right angle of the preoperculum lies a pore, marked
by a darker colour, and belonging to the inner osseous
ridge of the preoperculum, which ridge it pierces, but
covered by the adipose membrane that here, as usual,
covers the ramifications of the lateral line. This pore
is always distinct in Scandinavian Vendace and gene-
rally in young Gwyniads (as in young Nelma’s); but in
adult specimens of the larger Coregoni (as in the Nelma)
it becomes more and more indistinct, usually invisible.

The dorsal fin is of the shape commonest among
the Salmonoids, trapezoidal, the first branched ray being
the longest in the fin. The distance between it and the
tip of the snoutd is on an average about 43 1/2 % of the
length of the body, usually somewhat greater than in
the closely allied Siberian form Coregonus Merlcii. The
base of the fin measures on an average in albula about
9 %, in vimba about 10 %, of the length of the body,
and its height (the length of the longest ray) in the
former about 14%, in the latter about 15 % of the same.
The anal fin differs but little from the dorsal, but is
longer and lower, most so in vimba , its base measuring
on an average about 11 or 12 %, its height about 8x/2
— 9V2 0//°i °f the length of the body. Its distance from
the tip of the snout is on an average 69 by % of the
length of the body. It terminates, like the adipose fin,
at a distance from the caudal fin that is on an average
710 of the length of the body. The caudal fin is deeply
forked, most so in vimba , the average length of the
middle rays being about 5 1/2 — 5 %, of the longest rays
about 17 — 19 %, of the length of the body.

The pectoral fins are obliquely pointed, and their
length is on an average 16 — 15 % of that of the body.

“ The changes of growth run between about 32 and 23 /.

h The changes of growth run between about 27 and 20 %.

e On an average 25‘7 % of the length of the head.

d On an average 27'7 % of the length of the head.

e We may remind the reader that here, as in the following Gwyniads, the snout is measured from the articular knob of the maxil-
lary bone.

113

Scandinavian Fishes.

896

SCANDINAVIAN FISHES.

The ventral tins are broader and truncate, and their
average length is about 141/ 2 — 13 % of that of the body.
The distance between the latter tins and the tip of the
snout is on an average about 47 %, the preabdominal
length about 28 %, and the postabdominal length about
24 %, of the length of the body.

The scales of the body we have already noticed.
Only the head and the fins are naked. The lateral line
runs from the temples along the posttemporal bone on
each side, descending at the top of the gill-opening,
but its backward course on the sides of the body is
almost perfectly straight, with only a slight downward
curve at the extreme front.

The coloration is green, shading into steel-blue, on
the back, silvery white on the sides, of a more or less
pure milk-white on the belly. The back, as well as
the occiput and snout, is slightly transparent, but far
less than in the Smelt. The tip of the snout and the
point of the lower jaw are usually coloured with a black
pigment, which extends in a thinner coat to the maxil-
laries and to the margin of the follicle of each scale
on the body. The gill-covers commonly gleam with a
brassy lustre, especially at the top. The adipose tin is
of the same colour as the back. The dorsal and caudal
tins are of a more or less dark gray, the other tins
light and transparent. The iris is silvery white, the
eyeball coal-black at the top.

The Vendace is really a Baltic fish, with the centre
of its range in the Baltic Sea and the* lakes of the
countries — principally the eastern ones — bordering on
the Baltic. But, if modern opinions as to the specific
determination be correct, it also occurs in Scotland, to
which country, says tradition, it has been introduced
from abroad. The Irish lakes (Loughs Neagh, Erne,
Derg, and Corrib) contain a form, Coregonus pollan, so
closely resembling the Vendace that the specific distinc-
tion between them can hardly lie maintained. The Pol-
lan as a rule has fewer gill-rakers (34 — 38 on the front
of the first branchial arch) and fewer anal rays (12 —
13). At an earlier age than the other Vendaces the
Pollan approaches in the depth of the snout the pro-
portions which we have already remarked as most cha-
racteristic of the true Gwyniads". The most distinctive
character of the Pollan, however, is the short base of
the anal tin, which measures at most about 3/n of the

length of the body, and is so short that the least depth
of the tail is at least 3/4 thereof. In this respect, as in
many others, the Pollan composes one extreme in the
form-series of the Vendaces (the albula group), the op-
posite extreme consisting of the Siberian Seldetkan or
Coregonus Merkii. Strangely enough, this difference in
form within the Vendace group answers to the diffe-
rence we have above observed between the Salmons of
the Atlantic and Pacific basins. The Oncorhynchus
group of the Pacific differs from the Salmo group of
the Atlantic mainly in the greater length of its anal
fin; and the nearer the range of the albula group ap-
proaches to the Pacific, the longer is the base of its anal
fin. We find the following average relations between
the length of the base of the anal fin and that of the
head reduced:

Average

Pollan.

Albula.

Vimha.

Seldetkan.

Length of the base of the anal fin express-
[ ed in % of that of the head reduced

62.5

73.9

82.5

90.3

These averages can hardly be the expression of a mere
accident, for they are attended with similar results in
several other relations; and there is thus a distinct
connexion here between the difference in form and the
geographical separation. Even in Scandinavia the true
albula and the virnba show some geographical separation,
the former being commonest to the extreme north and
in Lake Wetter, the latter in the Malar vallev and the
basin of Lake Wener; but our most typical specimens
of virnba are from Finland. These two forms are be-
sides different, as we have seen above, in most of the
average proportions ; but the changes of growth and the
sexual distinctions reduce the differences to such an
extent that constant characters can hardly be adduced.

The Vendace occurs in all the provinces of Sweden,
except Gothland, Blekinge, Halland, and BohuslarP.
From Norway it was described and figured even by
Ascanius, but it is found only in Lake Mjosen and
some of the small lakes to the extreme south-east. In
Finland it is as common as in Sweden, up to about the
69th degree of latitude. In the Arctic Ocean it is
wanting; but from Western Russia and the Baltic Pro-
vinces its range extends over North Germany to Hol-

“ Smitt, Riksmuseets Salmonider , p. 233.

6 Cf. Under cl. Bet. Forsl. Ny Fislceristadga 1883, p. 159.

VEND ACE.

897

stein. When Krcyer wrote his Danmarks Fislce, he
knew nothing of its presence in Denmark, but in 1868
he received it from Foussingo So near Renders, and
in the following year Feddeesen received it from Jul
So, another of the Jutland lakes. At about the same
date it is said to have been met with in Ulse So and
Sothorup So (Zealand). In South Germany, France,
Belgium, and Holland it is unknown.

The Vendace is no true marine fish , but it occurs
in fairly great number in the island-belts of the Gulfs
of Bothnia and Finland, at least to the islands of So-
dermanland, where it cannot be reckoned, however,
according' to Ekstrom, among the common fishes.

The Vendace is thus found in very different pla-
ces, both in brackish water and in fresh. It lives in
lakes large and small, deep and comparatively shallow.
At certain times it repairs to running water. It is a
gregarious fish, and is consequently taken at times in
great quantities, while it also supplies food to several
large fishes-of-prey. Out of the water it is not tena-
cious of life, but, if treated with care and furnished
with cool water, it may be kept alive long enough in
aquaria". To wind and weather it is very sensitive.
Excessive heat or cold drives it to deep wafer; and in
stormy weather it takes refuge under the lee of the
land. In spring it roves about in more scattered com-
panies. In summer it shoals in the lakes during July
and August, making its way in large armies towards
the surface and into shallow water, to seek its food,
which consists principally of Entomostraca , insects, and
small mollusks. During autumn and still later in the
year it spawns, its active movements at the surface
rendering it easy of observation. In the Norrstrom at
Stockholm numbers of small Vendace are taken in
spring with hoop-nets below the bridge of Norrbro; but
in autumn the larger Vendace are caught in traps just
above the bridge. The end of September and the be-
ginning of October is the best season for the last-
mentioned fishery. Two traps ( ryssjor ) have been
set, one in 2 feet of water, the other in 3, with the
tail end fastened to a stake, so that the current keeps
the trap straight, with the mouth down the stream.
The fisherman who has employed these traps for many
years declares that the V endace always follow the same
course, going hardly a foot or two to the right or left.
The Vendace swim up the stream only when the water

runs from Lake Malar to the sea, especially tvhen there
is rather a strong current; yet it sometimes happens,
though usually only one or two days a year, that a
single day’s take in these traps may amount to 190
kilo, of Vendace, averaging about 5 or 6 to the kilo-
gramme, which fetches at first hand 50 — 70 ore (63/4- —
9V2 d.). The Vendace are then on the way to their
spawning- place. “In the inner island-belt,’’ writes Sux-
devall, “oft' Vaxholm for example, the Vendace is
taken both in summer and autumn, but does not seem
to go far out. On one occasion 90 lispund (765 kilo.)
had been secured at a single haul of the seine off Fors-
vik (Ingaro) in the month of November; otherwise it is
only seldom that a few specimens are caught there.”
From Sodermanland Ekstrom states that, in the month
of December or about the time when the lakes freeze,
the Vendace resorts to shallower water, where it spawns
on a stony or sandy bottom, the spawning seeming,
however, to be of long duration, for Vendace not quite
spent are caught with ice-seines in the middle of Ja-
nuary. In the Government of Norrbotten, according
to Widegren, the spawning-season of the Vendace be-
gins at the middle of October. “It then makes its way
6 — 12 miles up the mountain streams, and deposits its
roe on a. stony or sandy bottom. Off the town of
Lulea the Vendace enters the bays and creeks at the
middle of October, and spawns on a muddy bottom in
2 — 6 fathoms of water. Off Muonioniska Village the
spawning takes place at the end of November.” In
Lake Wener Widegren found that the Vendace spawns
from the end of October in 10 fathoms of water, re-
sorting chiefly to the shallows among the islands round
Luro and Kallandso. In Lake Wetter, on the same
authority, it breeds about the 24th of October near the
bottom and at a depth of 60 fathoms. “The greatest
number spawn,” he says, “in deep water along the east
coast from Grenna to the island of Jungfru. The shoals
first met with at the spawning-places consist exclusively
of males. This would seem to be the case with the
true Gwyniads as well, for those first taken are always
males. After the spawning, which ends at the middle
of December, no more Vendace are taken in deep water,
though they probably pass the whole winter in the
depths. Early in spring they seem to have migrated
to the extreme south of the lake, and are never found
in very deep water, but remain throughout the sum-

We have kept large Vendace alive for a fortnight in a glass jar filled with the ordinary water supplied by the Stockholm Waterworks.

898

SCANDINAVIAN FISHES.

mer, from May to October, in about 30 fathoms of
water off the west coast. The fry, 4 — (3 inches long,
also stay during the greater part of the year in deep
water, but in late summer often ascend the inlets on
the north-east coast, between Hals and Hofvanas Point.”
Where several lakes communicate with each other, the
Vendace commonly roves from the lower to the upper
in order to spawn.

The ova are tine and rather numerous, somewhat,
though not firmly, adhesive. The ripe eggs which have
already dropped into the abdominal cavity are about
1V2 mm. in diameter. In females 19 — 20 cm. long
their average number was estimated by Collett at 3,600
to each female". When only 7 cm. long'', probably in
its second year, the Vendace has reached maturity.

The Vendace is caught only in nets and the like —
as we have mentioned above, in traps and hoop-nets.
It takes a hook extremely seldom, and then the bait
must be a tempting fly. Its flesh is good. “In places

where Vendace are taken in any considerable quantity,”
writes Eksteom from Morko, “the flesh is salted and
in this state supplies the poorer classes with a great
part of their winter food. Here, where only few are
taken, the fish is generally eaten fried, and in this
form its fine and rich flesh is a real dainty.” In Swe-
dish restaurants and even in the homes of the upper
classes the Vendace is better known for its roe, which is
made in Norrland into a kind of caviare, under the
name of Bleak roe ( lojrorn ).

The Vendace has many Swedish names. The vimma
cannot be regarded as a separate species, and the V en-
dace is often confounded with yonng Herring and goes
by the same names. In Wermland it is also called sil
and stint a, in Westerbotten sillack. Its other names
in Northern Sweden are Smdling ( Smarting ) and Robb-
oxen , a word which reminds us of Rceputsclika, the
current name for the species in Western Russia, the
Livonian Repsen , the Esthonian Rcebus.

THE GWYNIAD (sw. siren).

COREGONUS LAVARETUS c

Plate XLII, figs. 3 and 4.

Tip of the snout more or less deep , truncate or conical, the depth ( the breadth of the inter maxillaries in a vertical
direction) being more than 2/5 of the breadth across the articular knobs of the maxillaries. Point of the lower jaw
not projecting beyond the tip of the snout. Length of the maxillaries more than tivice their breadth.

R. hr. (7)8—9(10); D. — 1 — [(12)13 — 16(17)];

(9)10— 12 L v yj,

A. (2)3~4 [13 — 16(17)] : P. 1 ; V. — ;

10— 13(14)l j (12 — 13)14 — 16(17) (9)10—11(12)

9 — 1 1

C. x + 1 4- 17 + 1 + x\ Lin. lat. (72)75—100(106); L. tr. 1 ;

v 8—9(10)

Vert. 624

Syn. Lavaret , Belon, Nat., Div. Poiss ., p. 278. Lavaretus + Bezola
+ Farra (1. Ferra 1. Pala), Rondel., Univ. Aquat. Hist.,
part. II, pp. 162 — 164 + Oxyrynchus (p. p.), p. 195. Albula
nobilis + parva, Gesn., Hist. Anim., lib. IV, pp. 33 et 34.
Albula nobilis, Schonev., Ichthyol. Slesv. Hols., p. 12. Gui-
niad , Willughb., Hist. Pise., p. 183. Coregonus maxilla
superiore longiore plana, pinna dorsi ossiculorum quatuordecim,
Art., Ichthyol., Gen. Pise., p. 10; Syn. Pise., p. 19; Descr.
Spec. Pise., p. 37; Lin., Fn. Suec., ed. I, p. 118; Gisleh,
Vet.-Akad. Handl. 1753, p. 195. Coregonus maxilla superiore
longiore conica, Art., Gen., p. 10; Syn., p. 21.

Salmo Lavaretus, Lin., Syst. Nat., ed. X, tom. I, p. 310;
Ascan., Icon. Per. Natur., call. Ill, p. 6, tab. XXX; Bl.,
Fisch. Deutschl., part. I, p. 163, tab. XXV (4 -Salmo Thy-
mallus latus, p. 170, tab. XXVI + Salmo Marcena, p. 172,
tab. XXVII 4- Salmo Wartmanni, part. Ill, p. 161, tab. CV);
Retz., Fn. Suec. Lin., p. 348; Pall., Zoogr. Ross. Asiat.,
tom. Ill, p. 395 (+ Muksun, p. 398 + Polcur , p. 400);
Nilss. ( Coregonus ), Prodr. Ichthyol. Scand., p. 15; Yarr.,
Brit. Fish., ed. 2, vol. II, p. 142; Kr., Damn. Fisk., vol. Ill,
p. 55; Cuv., Val., Hist. Nat. Poiss., vol. XXI, p. 466 ( +
Cor. Palea, p. 477 4- Cor. Reisingeri , p. 496 + Cor. Nilsoni,
p. 497 4 Cor. sikus, p. 500); Nilss.,' Skand. Fn., Fisk., p.
458; Wdgrn, Ofvers. Vet.-Akad. Forb. 1862, p. 583 (+ Cor.
megalops, p. 589); Mgrn, Finl. Fiskfn., p. 49 ( + Cor. Wide-
greni , p. 52); Gthr, Cat. Brit. Mtts.. Fish., vol. VI, p. 178
(+ Cor. lapponicus, p. 181 4- Cor. gracilis, p. 182 + Cor.
richardsonii, p. 185 + GW. maxilla ns, p. 189 4- Cor. humilis,
p. 190); Coll., Forh. Vid Selsk. Chrnia 1874, Tilhngsh.,

° Benecke reckoned 10,000 to each female and estimated their diameter at about 2 mm., assumably in large specimens.

6 Cf. Smitt, Rilcsmuseets Salmonider, tab. metr. VIII, Coreg. No. 46.

i: The name is Old French from the neighbourhood of Geneva, and is derived by Rondelet from laser, to wash: “quod nurnquam
sordidus sit, sed bene ablutus sit, ob munditiem et candorem nomen habere videtur.”

e 58 — 62, according to Gunther; 60 — 62, according to Kroyek; 57 — 64, according to Fatio.

GWYNIAD.

899

p. 166; ibid. 1879, No. 1, p. 89; Id., N. Mag. Naturv., Bd.
29, p. 108; Malm, Gbgs, Boh. Fa ., p. 546; Feeders., Naturh.
Tidskr. Kblivti, ser. 3, vol. XII, p. 79; Smitt, Intern. Fisch.
Ausst. Berlin 1880, Scliwed. Kat., p. 17; Bncke, Fisch.,
Fischer., Fischz. 0 ., IV. Preass., p. 150; Mela, Vert. Fenn.,
p. 347, tab. X ( + Cor. Nordmcmni , p. 350); Smitt, Ofvers.
Vet.-Akad. Forh. 1882, No. 8, p. 37 : Gt. Intern. Fish. Exhib.
Loudon 1883, Swed. Cat., p. 188; Norb., Handl. Fislcev.,
Fiskafv., p. 404; M5b., Hoke, Fisch. Osts ., p. 131; Smitt,

Vet.-Akad. Handl., Bd. 21, No. 8 (1885), p. 248; Bncke

in M. v. d. Borne, Hcmdh. Fischz., Fischer.., p. 144; Lill.t.,
Sv., Norg. Fisk., vol. II, p. 733.

Salmo Oxyrinchus, Lin., Syst., 1. c., p. 311; Pall., 1. c., p.
403 (? + Salmo microstomas, p. 405); Nilss. ( Covegonus ),
Prodr., p. 14; Ekstr., Vet.-Akad. Handl. 1834, p. 12; Ivr.,
1. c., p 76; Cuv., Val. ( oxyrliynclius ), 1. c., p. 488 ( + Cor.

conorhynchus, p. 485); Nilss., Fa., 1. c., p. 453; Sund.,

Stockli. L. Hush.-Sallsk. Handl. 1855, pp. 81 et 90; Lloyd,
Scaucl. Adceut., vol. I, p. 129; Wdgrn, 1. o., p. 577; Sieb.,
Siisswasserf. Mitteleur ., p. 259 ( + Cor. Marcena , e Bl., p.
263); Gthr, 1. c., p. 173 ( + Cor. Lloydii, p. 174); v. Bemm.
in Hericlots, Bouicst. Fa. Nederl. , part. Ill, p. 386; Coll.,
1. c., 1874, p. 165; Malm, 1. c., p. 544; Winter, Naturh.
Tidskr. Kbhvn, ser. 3, vol. XII, p. 44; Fedders., ibid., p.
80; Day, Fish. Gt. Brit., Irel., vol. II, p. 126, tab. CXXI,
fig. 2; Norb., 1. c., p. 406; M6b., Hcke, 1. c., p. 130;
Bncke (M. v. d. Borne), 1. c., p. 143.

Coregonus clupeoicles, Lacep., Hist. Nat. Poiss., vol. V, p.
697; Gthr, 1. e., p. 188; Day, 1. c., p. 127, tab. CXXII
= Cor. La Cepedei, Paiin., Ann. Nat. Hist., vol. I (1838),
p. 162 cum fig. (+ Cor. microcephalus , p. 163 cum fig.);
Yarr., 1. e., p. 151.

Coregonus Fera, Jur. Mem. Soc. Phys., D’Hist. Nat., Geneve,
tom. Ill, part. I, p. 190, tab. 7 (+ Cor. liiemalis , p. 200,
tab. 8); Rapp, Jabresb. Ver. Vater. Naturk. Wiirtemb., Jahrg.
X, p. 154, tab. VI (+ Cor. Wartmanni, e Bl., p. 148,
tab. V + Cor. acronius, p. 158); Sieb., 1. c., p. 251 ( + Cor.
Wartmanni, e Bl., p. 243 + Cor. hiematis , e Jur., p. 254);
Lunel, Hist. Nat. Poiss. Bass. Lem., p. 106, tab. XI ( +
Cor. liiemalis , e Jur., p. 114, tab. XII).

Coregonus generosus, Peters, Monatsber. Akad. Wiss. Berlin
1874, p. 790.

Coregonus macrophthalmus , Nusslin, Zool. Anz., Jahrg. V (1882),
p. 164 = Cor. exiguus, Klunz., Jahresb. Ver. vat. Naturk.
Wilrttemb., Jahrg. 40 (1884), pp. 110 et 117.

Coregonus Sulzeri, NOssl., 1. c., p. 253.

Coregonus dispersus (■= Cor. Wartmanni, e Bl., + Cor. an-
nectus + Cor. exiguus, e Klunz.) + Cor. balleus (= Cor.
Asperi + Cor. Scliinzii + Cor. acronius, e Rapp + Cor. hie-
malis, e Jur.) 4- Cor. Suidteri , Fatio, Fn. Vert. Suisse,
vol. V, p. 67, cett.

Ada. De synonymis atnericanis, quum speciinina piscium pau-
ciora solum vidi, hierco quid dicam. Vix tamen Coregonum
album (Le Sueur, Richardson, vide Fn. Bor. Airier., part. Ill,
p. 195, tab. 89, fig. 2 = (dor. clupeiformem , Mitch., vide
Jordan et Gilbert, Bull. U. S. Nat. Mus., No. 16, p. 299)
a lavareto pycnocentro nostro diversum credam; neque Core-

a See Underd. Bet. Forsl. Ny Fiskeristadga 1883, p. 159.
6 See Smitt, Riksm. Salmon., 1. c., tab. IV, fig. 65.
c Smitt, 1. c., tab. VI, figg. 98 et 99.
d Prosopittm, Milner.

gonus quadrilateralis (Rich., 1. e., p. 204, tab. 89, fig. 1)
a polcure 1. brachymystace nostro differre videtur.

The Gwyniad with its numerous variations has
proved no less troublesome to the systematist than the
Salmon. A great proportion of the names enumerated
above have been applied, we may almost saw, in-
discriminately, for though different forms have, no
doubt, been distinguished — and these in many localities
very well marked — their natural relations to each
other have not been understood.

The Gwyniad is so like the Vendace in most essen-
tial respects that a detailed description is unnecessary.
The most striking difference lies in the form of the snout,
and the cause of this difference, the shape and position
of the intermaxillaries, ive have already investigated.
But the Gwyniad attains a much greater size than the
Vendace. In Lake Superior Gwyniad weighing 23 lbs.
(nearly 107a kilo.) are taken, and in Sweden" the Gwy-
niad attains a length of about 9 dcm. and a weight of
about 6V3 kilo. Specimens of this size are, however,
restricted to the northern provinces, the Baltic, and
the largest lakes, as for instance Wener and Wetter;
in the smaller lakes, where it is more or less entirely
landlocked, it is always of less considerable dimen-
sions, hardly larger than the largest. Vendace, and
retains more or less of the characters of youth. Merely
from this we can gather that the Gwyniad’s true home
lies in the north and in salt water, as well as in the
rivers flowing into the sea..

To gain a correct understanding of the relations
between the numerous forms in which the Gwyniad
appears under different conditions in Scandinavian wa-
ters, it is necessary to include in the comparison at
least two of the Siberian forms that, belong to the same
series of development. Both these forms were suspected
by Pallas, who first, described them, to be identical in
species with our common Gwyniad; but one of them,
the muksun b, shows the development of the type to the
greatest size of body conjoined with a persistency of the
resemblance to the Vendace, while the other, the polcurc,
represents the opposite extreme in the evolution of the
type, being approximated to the so-called tscliir ( Salmo
- Coregonus d — nasus, Pall.), which is principally di-
stinguished by its short and deep (high) maxillaries.

900

SCANDINAVIAN FISHES.

I

Like the Vendace forms the muksun has long and dense
gill-rakers; but the polcur, like the tschir, short and
scattered ones. Furthermore the muksun has a compa-
ratively shallow snout, the polcur a comparatively deep
snout. The two forms as well as the typical Gwyniad
live together in the Siberian rivers and the Arctic Ocean
off their mouths; and the difference between them coin-
cides in an eminent degree with the distinction drawn
bv the fishermen in several Swedish lakes between the
different kinds of Gwyniad in their catch. The muksun
type they call blasik or gronsik (in Norrland asp, when
large, or sikloja, when small); while the polcur type
bears the names of sik, fetsik, storsik, hvitsik or botten-
sik. The typical ndbbsik (Houting), with its conically
prolongated snout, also belongs in fact to the polcur
group. The shape of the snout has probably been em-
ployed time out of mind by the Swedish fishing popu-
lation as a character for separate varieties of Gwyniad ;
but in 1831 Laestadius directed attention to the gill-
rakers of the asp as a distinction from the sika. That
Widegren had also observed the last-mentioned cha-
racter, appears from the drawings which he left at his
death in 1878, and which we have been enabled here
to reproduce (fig. 222, B — F, above, p. 891).

These two groups of the true Scandinavian Gwyn-
iads we distinguished in 1879 by the names of Core-
gonus Wartmanni and Cor. lavaretusb ; and they are
sometimes well marked even where they occur in the
same lake (figs. 224 and 225). As an example we may
adduce the Royal Museum collections from Lake Stor
(Storsjon) in Jemtland. Seven bldsikar and six fetsikar
show the following constant differences:

Blasilcar.

Fetsikar.

Number of gill-rakers

on the first branchial arch of

each side

34—45

20—24

Depth of the snout in % of the length of the head

8.6 — 9.s

11.1—12.3

7? 77 77 7? 7? 77

, „ „ „ „ „ reduced.

11.4—13.2

14.8 — 16.7

77 77 7? 77 77 77

, ,, breadth of the snout

50.o — 62.5

66.6—87.5

7? 77 77 77 77 7?

, „ length of the maxillaries ...

27.3 — 35.7

40. o — 46.2

Length of the lower jaw in

% of the length of the head

4-

T

o

40.9 e — 39.3

„ „ „ „ „ „

,, „ „ „ „ „ „ reduced

62.9 — 59. bd

59.3 — 51.5

„ „ „ maxillaries,,

,, „ „ „ „ ,, lower jaw..

60.o — 66.7

68.4/ — 73.o

But on examining the Fetsik fry of this lake, we
find that, even at a length of 70 — 80 mm., only the
first three characters hold good. Thus at the outset
of their development the two forms stand nearer to
each other, even after the external shape of the body
has assumed the character of the group. As is more
distinctly shown by our figures, the bldsikar of this
lake belong to the group characterized in Gunther* 5 6'
by “snout vertically truncated,” the fetsikar, on the
other hand, belonging to the group7* in which “the
snout is obliquely truncated, with the nose protruding”.
The same difference appears, however, between true
bldsikar as well, for example in Lake Ring, where, to
the best of our knowledge, only bldsikar are found1.
It is still more difficult to maintain the distinction
between the two groups where they occur together in
the great Swedish lakes or in the sea. The Royal
Museum collection of Gwyniads from Lake Wetter
shows the most varying form of snout7, exemplifying
almost all conceivable transition forms between shallow-
snouted Gwyniads (which we have called tapinorlvynchik )
— not represented, however, by fully typical speci-

a “The Asp occurs in all the large lakes, up to Kilpisjarvi in Tornea Lappmark. Spawns in autumn, in the rivers in October. Is

exactly like the Sik, but differs in the gills, whose spines in the Asp are long and fine.” L. L. L^stapius, MS in the Royal Museum.

6 See the Swedish Special Catalogue at the Fisheries Exhibition in Berlin 1880. Besides the Gwyniads included in this catalogue, my
tables of measurements (published in 1886 in Vet.-Akad. Handl.) were accessible in manuscript, and in the course of a public discussion I
cited the results of these tables, results which render the specific determination within the lavaretus group so untrustworthy that we have
rather to deal with local varieties (See Giglioli, Annali dell’industria e del commercio 1880 (Roma 1881), num. 29, p. 38).

Subsequently NOsslin (1882), Klunzinger (1884), and Fatio (1885 — 90) have arrived at the same conclusion, with respect to the
connexion between the characters derived from the appearance of the gill-rakers and those drawn from the form of the snout, as I maintained
on the said occasion and published in 1882 in Ofvers. Vet.-Akad. Fork. But they adhere to the older opinion according to which several
species may be determined with certainty, by means of these characters, within the lavaretus group.
c 43‘5 in a single specimen.
d 58'8 in a single specimen.
e 44'4 in a single specimen.
f 62'5 in the youngest specimen.
g Cat. Brit. Mus., Fish., vol. VI, p. 187.
h 1. c., p. 178.

1 Cf. for example Smitt, Riksrn. Salmon., tail. Y, figg. 78- — 81, representing four bldsikar ( Coregonus Nilssonii ) caught at the same time.
■> See Smitt, 1. c., tafl. IV, figg. 69 — 71 and tail. V, figg. 73 — 75.
k From ran eivog, loiv, and Qiyyog, snout.

GWYNIAD.

901

Fig. 224. Blasik ( Cor eg onus Wartmanni, forma Nilssonii ), 1/2 of the natural size, taken in Lake Stor (Jemtland) on the 13th Sept.,
1877 by Mr. R. Lundberg, Inspector of Fisheries, a, head of the same specimen, natural size; b, snout of the same,

seen from in front, natural size.

b

Fig. 225. Fetsih ( Coregonus lavaretus), x/2 of the natural size, o71? taken in Lake Stor (Jemtland) on the 13th Sept., 1877 by Mr. R. Lund-
berg, Inspector of Fisheries, a, head of the same specimen, natural size; b , snout of the same, seen from in front, natural size.

902

SCANDINAVIAN FISHES.

mens — and deep-snouted ( hypselorhynchi ■“), or between
narrow-snouted (, stenorliynclii1 ) and broad-snouted ( eury -
rhynclii0), without these differences being always at-
tended by analogous differences between dense gill-
rakers ( pycnocentrid ) and scattered (: manocentri e). To
this we must add that among the comparatively pyc-
nocentrous Gwyniads of Lake Wetter specimens occur
(tig. 226) which differ in hardly any respect from the
form of the manocentrous Gwyniads, e. g. in Lake Stor
(tig. 225). An examination of the Lake Wener Gwyn-
iads has yielded the same result; but in both these
lakes, as well as in the sea and in the large rivers
flowing into the sea, there appears a still more pro-
minent alteration in the form of the snout, an alteration
which has given rise to the name of ndbbsik ( Beaked
Gwyniad, Houting; tig. 227), the oxyrhynchus of old
writers. This alteration consists partly in the prolonga-
tion and thickening of the ethmoidal cartilage, but
mainly in an agglomeration of connective tissue and
fat. It is analogous to the elongation of the point of
the lower jaw in the Salmons, and cannot in itself
justify any specific, distinction, whether it appears in
the narrow-snouted Gwyniads7 or in the broad-snouted
Its reaches its highest development in old Gwyniads
from the depths of Lake Wener, which do not differ,
however, in other respects from their companions, the
broad-snouted Gwyniads of the same lake. But the
Houting form was first observed in the North Sea on
the Belgian, Dutch, and German coasts, whence it ascends
the Rhine, Weser, and Elbe in order to spawn, and in
the south of the Baltic, where it enters the Haft’s and
the Oder.

However variable the shape of the snout may ap-
pear, it is still the most trustworthy guide to the de-
termination of the forms — as the fisherman’s experience
has long since taught him — although the characters
cannot be well defined. This indefiniteness of character
we can easily understand, for we can range the diffe-
rent forms in an almost unbroken series according to

their varying degrees of resemblance to one or other
of the above-mentioned extremes, the muksitn and the
polcur.

The shape of the snout also affects the length of
the jaws, from which we may derive important cha-
racters. The appearance of the form-series at once
suggests that the pycnocentrous forms should have longer
jaws, and the changes of growth71 show that during
youth the relative length of the jaws undergoes a re-
duction. But in old Gwyniads from Lake Wener — the
form which in Gunther bears the name of Ooregomis
maxillaris, the German Madui-Mar cme — we observe
a retrogression towards the characters of youth, a re-
trogression which ranges this form, as regards the
length of the maxillaries, even when the tip of the
snout is prolongated to the Houting type {Cor. oxy-
rhynchus), beside the pycnocentrous asp from Lapland.
The other extreme of the series — the polcur form,
which, as we have mentioned, really belongs to Siberia
and Northern Russia, but also occurs in the northern-
most rivers of Sweden — sometimes has the maxillaries
so reduced that we have conferred on this modification
the special name of brachymystax'.

The shape of the bod)’ is the same as in the Ven-
dace, but in general somewhat deeper, deepest as a
rule in the polcur and the forms that come nearest it,
the manocentrous forms as a rule having a deeper body
than the pycnocentrous. The greatest depth of the
body varies in adult Gwyniads between about 22 and
25 % (in gravid females as much as 27 %) of its length;
in the polcur the percentage is usually 26 or 27. In
young Gwyniads measuring 37 — 143 mm. we find the
greatest depth to increase with growth from 1 5 x/2 to
19 Vs of the length. But on account of the temporary
variations due to the periodical tumidity of the gene-
rative organs, the greatest depth of the body, here as
in most cases, is ill adapted to express its characteristic
form. Far more important in this respect is the least
depth of the body. This is seldom so small as in the

“ bynqloSi high.
b ovevog, narrow.
c Slqvq, broad.

d From rcvvjvog , dense , and v.tvroov, spine.
e yavog, scattered.

f Smitt, 1. c.. lafl. IV, figg. 69 and 70.
g 1. c., fig. 67.

h 1. c., tab. metr. XIII, and */s >n Nos. 362 — 364 and 397 — 403.

’• Answering in essential respects to a form which inhabits the depths of the Lake of Constance, and which has been named Core-
gonus acronius. But this latter form', judging by the two specimens we have examined (see Smitt, 1. c., tab. metr. X, Nos. 162 and 163),
has much longer pectoral and ventral fins and a shorter snout.

GWYNIAD

903

Fig. 226. Coregonus lavaretus , pycnocentrus , l/2 of the natural size, cF, taken in Lake Wetter, Nov., 1863, by Widegren. a, head of
the same specimen, natural size; i, snout of the same, seen from in front, natural size.

Fijr. 227.

Coregonus lavaretus , o.ryrJtyncJnts, spawning o71? taken in Lake Wener at Stora Eken.

a. head of the same specimen, natural size.

a

of the natural size.

Scandinavian Fishes.

114

904

SCANDINAVIAN FISHES.

Vendace, varying on an average about 7 % (in polcur
about 8 %, in tschir nearly 9 %) of the length of the
body, between 33 and 40 % (in polcur 41 — 45 %, in
tschir 52 %) of the length of the head, or between 44
and 52 % (in polcur 56 — 61 %, in tschir 73 %) of the
length of the head reduced. Some Gwyniad forms
may indeed be said to lie characterized by a shallow
tail (an elongated body); but it should be observed
that this character is also a character of age. Conse-
quently it happens that, in certain lakes, especially in
Lake Bolmen in Smaland, where it has longest been
known, but also in many other lakes from Lapland to
Smaland, we meet with a Gwyniad form (Nilsson’s
Coregonus lavaretus, Malm’s forma holmensis) which
differs, so far as we know, constantly from the allied
forms in the shallow peduncle of its tail. But in other
waters, for example in the system of small lakes in
Smaland which falls into the Baltic through the River
Morrum, this very form is developed into the form
with deeper tail, which else has its most typical repre-
sentative in the Lake Ring Gwyniad (V alenciennes’
Coregonus Nilssonii). The hlasikar in the uppermost
of the said lakes — Helga and Bergunda" — in most
respects afford an instructive example of the changes
of growth through which the Gwyniad passes in the
less advanced stages of its form-series. In particular
they show how the least depth of the body, the most
distinctive character of holmensis, increases with age so
as to approach the typical Coregonus Nilssonii. In

o

Asnen, a lake of greater size and situated nearer the
outlet of the system, the Nilssonii type6 is developed
at an earlier age (a smaller length) than in the two
said lakes. In this locality it therefore seems that
Cor. holmensis should really be regarded as a young
stage of Cor. Nilssonii , which in other places, for ex-
ample in Lake Bolin, may persist as a separate form.

The deeper form of the body is most often accom-
panied by a special arrangement of the scales that
gives the fish a singular appearance. When the skin
is distended, the rows of scales part from each other0,
and the exposed portion of each scale becomes hexagonal
instead of semicircular. The Gwyniad then displays a

striking resemblance to the Grayling. In the hlasikar
— but especially in the typical Lake Ring Gwyniad —
we also find a special form of scale, at least in the
rows nearest to the lateral line, the free (hind) margin
of the scale becoming more or less elongated at the
middle (forming a rounded obtuse angle instead of a
circular arc). But in all the Gwyniads examined by
us this difference also appears between the scales on
the sides of the tail and those on the forepart of the
body. The character assigned by Lilljeborg to Core-
gonus Nilssonii, that the exposed part of the scale in
the rows nearest to the lateral line is more than half
as long as high, may be found in most Gwyniads, at
least in solitary scales.

While it often happens, especially among the hld-
sikar , that the comparatively great size of the eyes is
persistent longer than usual — a relation which Wide-
gren expressed by establishing a separate species, Core-
gonus megalops — the fetsikar run to the opposite ex-
treme, the reduction in the relative size of the eyes
during growth proceeding more rapidly and advancing
further than usual. The form produced in this man-
ner we have called microps, less in order to indicate
the importance of the difference as such, than because
it is usually accompanied by an increased fatness which
has rendered the form economically important. It
occurs, according to the collections of the Royal Mu-
seum, in the Gulf of Bothnia, the rivers of Norrland,
Lake Wener, and Bohuslan. Malmgren sent home
specimens of a similar form from Lake Ladoga. The
same importance is shared by another form, from the

o

Angerman and Torne Elfs, which is characterized by
a comparatively small head, and which we have there-
fore named microcephalus.

None of these forms seems quite to deserve the
rank of an independent variety. They sometimes appear
to occur alone, but as a rule at least two are found
in the same lake or river — Lake Wener possesses at
least 7 of these forms — and that they intermingle
during the spawning, is more than probable. But as
a rule a distinction may be drawn between them in
the following manner:

a Smitt, ]. c., tab. metr. XI, Nos. 210 — -215.
b 1. c., Nos. 216 and 217.

c A striking example of this is Siebold’s figure of Coregonus acronius {Susswasserf. Mitteleur., taf. II), monstrously swollen owiDg
to the expansion of the air-bladder when the fish was drawn up from deep water; but the same appearance is almost as marked in gravid
females. We have above seen a similar difference in the appearance of the scales among the Perches for example: — cf. the Carass Perch
(fig. 3, p. 29) with the common Perch.

GWYNIAD.

905

A: Shallow-snouted , as a rule with dense
gill-rakers: depth of the tip of the
snout at the rostral protuberances (a3a4,
see fig. 228) less than 15 % of the
length of the head reduced (ad, see fig.);
gill-rakers on the first branchial arch
as a rule more than 30.
a: Length of the maxillaries more
than 30 % of that of the head. Gill-
rakers on the first branchial arch
more than 50. (A Siberian form) Coregonns muksun.
b: Length of the maxillaries less than
30 % of that of the head. Gill-
rakers on tire first branchial arch

less than 50.- Cor eg onus Wartmanni.

a: Depth of the tip of the snout
less than 61 % of its breadth
(cija2, see fig.).

aa: Least depth of the peduncle
of the tail less than 36 %
of the length of the head
— Cor. bolmensis.
bb : Least depth of the peduncle
of the tail more than 36 %
of the length of the head.
aa: Gill-rakers on the first
branchial arch less than
40 — Cor. Nilssonii.

(3(3 : Gill-rakers on the first
branchial arch more
than 40 — Cor. as pi us.

(3: Depth of the tip of the snout
more than 61 % of its breadth
— Cor. Wartmanni.

B : Deep-snouted , as a rule with scattered
gill-rakers : depth of the snout at the
rostral protuberances more than 15 %
of the length of the head reduced;
gill-rakers on the first branchial arch
as a rule less than 30.
a: Length of the maxillaries more

than 30 % of that of the head Coregonns maraena.

a: Length of the maxillaries less
than 76 % of that of the lower
jaw — Cor. maxi liar is.

(3: Length of the maxillaries more
than 80 % of that of the lower
jaw — Cor. oxyrhynchus.
l>: Length of the maxillaries less than
30 % of that of the head.
a: Length of the lower jaw greater

than the least depth of the tail Coregonus lavaretus.
aa : Length of the head more
than 18 % of that- of the
body.

aa: Vertical diameter of
the eyes more than 24
% of the length of the
head reduced — Cor.
lavaretus.

(3(3: Vertical diameter of
the eyes less than 24
% of the length of the
head reduced — Cor.
m i crops.

bb: Length of the head less
than 18 % of that of the body
— Cor. microeephalus.

Fig. 228. Outline drawing of a Gwyniad ( Coregonus Nilssonii ), showing how the measurements in the table of averages are taken.

Line ab = length of the body.
ac = ,, ,, ,, head.

ad = ,, ,, ,, ,, reduced.

a,a2 = breadth of the snout.

Ha\ = depth „ ,, „

ag = length „ „

gh = longitudinal diameter of the eyes.
ef = vertical ,, ,, ,, ,,

ai = length of the maxillaries.
mn ----- breadth ,, ,, ,,

Tel — length of the lower jaw.

op = length of the suboperculum.
ag = distance between the dorsal fin and the
tip of the snout.
qr = base of the dorsal fin.
st = height ,, „ „ „

uv = length of the pectoral fins.
uiv = preabdoininal length.
aiv = distance between the ventral fins and the
tip of the snout.
wee = length of the ventral fins.
wy = postabdominal length.

ay = distance between the anal fin and the tip
of the snout.

yz = base of the anal fin.
a(3 = height ,, ,, ,, „

yd = dorsal margin of the peduncle of the tail.
kxk = length ,, „ ,, ,, ,, ,.

zE = ventral margin ,, ,, ,, ,, ,, ,,

Crj = least depth „ ,, „ ,, ,,

d~b = length of the middle caudal rays.

I'A = ,, ,, ,, longest rays in the upper

caudal lobe.

906

SCANDINAVIAN FISHES.

ft: Length of the lower jaw less
than the ■ least depth of the
tail Coregonus polcur.

aa: Length of the maxillaries
more than V \ of that of the
head and than 36 % of that
of the head reduced — Cor.
polcur.

bb: Leno'th of the maxillaries

O

less than i/i of that of the
head and than 36 % of that

of the head reduced — Cor.
braehymystax.

Instead of describing these forms at length we
shall refer the reader to the two following tables of
averages, the first intended to show both the difference
of form and sex and the changes of growth in the
three forms, of which we have examined a sufficient
number of male and female specimens of various ages
to enable us to trace the last-mentioned changes.

Aver

age

N i 1 s s

o n i i

1

V a r t m a n n

i.

La

v a r e

t u s.

9

&

9

c?

?

cf

Length of the body expressed in millimetres (ab in tig. 228)

184

257

161

253

184

314

167

253

147

320

82

163

305

„ „ head (ac)

in % of the length

of the body

20.7

19.9

20.8

19.7

20.i

19.6

21.5

18.9

20.6

19.6

21.i

19.9

19.5

,, „ „ „ reduced (ad)

7? ff ff ff ff

99 99

15.7

14.6

15.8

14.7

15.3

14.8

16.o

14.3

15.5

14.7

15.8

14.8

14.6

Distance between the dorsal fin and the

tip of

the snout (aq)

ff y> ff

„ „ „

43.5

44.o

42.7

43.2

42.8

43.6

44.o

42.6

43.5

44.3

43.6

43.0

43. s

Base of the dorsal fin (qr)

ff ff ff ff ff

99 99 99

ll.i

10.7

11.0

11.5

ll.i

12.3

10.8

10.8

10.5

11.0

11.2

11.9

11.5

Height ., ., ,, „ (st)

ff f> ff ff ff

9 9 9 > 99

15., 5

15.4

15.7

15.8

15.i

15.8

15.o

14.7

14.7

15.4

11.7

14.5

14.8

Length of the pectoral fins ( uv )

ff ff ff ff ff

„ „ „

13.8

14.2

15.i

15.2

14.4

15.2

15.9

15.5

13,i

15.1

12.3

13.i

15.3

Preabdominal length (uw)

99 99 99

27.5

28.6

26.5

27.7

29.0

29.2

27.o

28.9

27.9

29.6

28.2

(27.9)

28.5

Distance between the ventral fins and the

tip of

the snout (aw)

ff ff ff ff ff

99 99 99

47.9

47.4

47.i

46.9

47.8

48.7

47.i

46.9

48.2

49.i

48.4

47.7

44.4

Length of the ventral fins (wx)

ff ff ff ff ff

99 99 99

13.8

14.3

14.6

14.6

14.i

14.3

14.1

14.4

13.3

14.3

11.4

13.i

14.2

j Postabdominal length (wy)

fj 19 9)

» „ „

25.6

26.o

24.6

25.8

24.9

26.9

24.6

25.7

24.4

26.o

23.5

24.7

25.9

Distance between the anal fin and the

tip of

the snout (ay).

ff ff ff ff

ff ff ff

71.2

71.4

70.3

71.i

71.6

73.7

71.o

70.6

71.3

73.4

70.7

71.3

72.s

Base of the anal fin (yz)

99 99 99 99 99

ff ff ff

11.3

11.3

12.i

12.i

10.7

10.7

10.7

11.7

10.8

11.0

10.1

10.7

11.0

Height „ „ „ («/?).....

99 9 9 99 99 99

ff ff ff

9.8

10.5

10.1

10.7

9.9

10.4

lO.o

10.1

8.7

10.1

7.1

8.8

lO.i

] Dorsal margin of the peduncle of the tail (yd).. „ „ „ „ ,,

ff ff ff

9.9

9.1

10.2

9.1

9.3

8.9

10. o

9.3

10.i

9.0

9.8

9.7

8.7

Length „ „ ,, „ „ „

,, ,, ,,

ff ff ff

14 i

13.5

10.6

13.7

13.2

13.3

13.3

13.3

13.3

13.2

13.o

13.4

13.5

Ventral margin,, „ „ „ „ „

{ZS) .. ,, ,, ,, ,, 99

ff ff ff

9.2

8.9

9.5

8.9

9.3

8.6

9.7

9.4

9.6

8.5

9.3

9.6

8.7

Least depth ,, ,, ,, „ „ ,,

(b7?)-- 99 99 99 9 9 99

„ „ „

6.8

7.4

7.o

7.5

7.o

7.4

6.8

7.2

6.4

7.2

6.i

6.7

7.i

| Length of the middle caudal rays (, 9h)

99 99 99 99 99

99 99 99

5.2

6.2

5.5

5.9

5.9

5.4

5.9

5.8

5.6

5.4 ’

6.3

5.5

5.4

„ ,, „ longest rays in the upper

caudal

lobe (ix)

99 99 99 99 99

„ „ „

—

20.3

17.5

19.8

18.9

19.2

19.3

18.5

16.i

18.4

-

17.5

18.3

| Length of the snout (ay)

99 99 99 99 99

„ „ head

24.3

25.i

24.8

24.9

24.8

26.o

25.i

25.o

25.5

27.3

23.8

25.7

22.s

Longitudinal diameter of the eyes (gli)

99 99 99 99 99

99 99 99

25.3

22.6

26.9

23.1

24.7

21.2

25.2

22.2

25.2

20.8

27.7

24.6

21.i

Vertical ., (ef)

99 99 99 99 99

99 99 99

22.3

19.4

22.6

19.9

22.7

18.2

21.3

18.6

21.3

18.i

23.6

21.2

18.5

Breadth ofthe snoutatthe rostral protuberances(rt,n2) ,, ,, „ ,, „

99 99 99

16.5

16.i

17.0

16.5

14.6

16.o

15.3

15.9

16.o

15.9

14.6

16.3

16.2

Depth „ „ „ „ „ „ „

(a3a*») ff ff ff ff ff

99 99 99

9.4

9.i

9.4

9.4

9.4

10.6

9.o

10.3

10.4

12.2

9.2

10.5

11.9

Breadth of the forehead at the middle of the eyes „ „ „ „ ,,

99 9> 99

28.6

30.i

27.6

31.2

28.4

30. o

28.3

30.9

27.9

29.7

27.6

29.3

30. o

Length of the maxillaries ( ai )

99 99 99 99 99

99 99 99

25.2

27.5

28.9

27.7

28.2

27.i

27.6

29.o

29.o

27.8

28.9

27.9

27.8

„ „ „ lower jaw (kl)

- ff ff ff ff ff

99 99 99

45.2

44.o

45.8

44.6

44.5

42.o

45.8

44.5

43.5

41.3

44.4

42.7

41. s

,, ., „ suboperculum (op)

99 99 99 99 99

99 99 9’

30.3

29.8

29.7

29.7

29.0

30.2

30.8

29.6

29.6

29.o

30.8

30.i

28.9

Vertical diameter of the eves

.. in % of the length of the head reduced

30. o

26.4

29.9

26.2

29.7

24.i

28.i

24.5

28.3

24.2

31.4

28.4

24.0

Depth of the snout at the rostral protuberances „ „ „ „ „ „ „

99 99

12. G

12.4

12.4

12.7

12.3

14.0

12.i

13.6

13.8

16.3

12.3

14.1

15.8

Length of the maxillaries.

-* 99 99 99 99 99 99 99

99 99

33.8

37.4

38.3

37.3

36.8

35.9

37.o

38.3

38.6

37.3

38.5

36.8

36.9

Breadth „ „ „

in % of their length

37.9

37,o

36.4

36.5

37.0

37.3

36.9

36.2

37.7

38.0

30.9

38.3

37.7

Length ,, ., .,

. in % of the length of the lower jaw

55.7

63.o

63.9

62.2

63.7

64.7

60.3

65.2

66.9

67.4

65.i

64.5

66,7

Least depth of the peduncle of the tail ...

- 99 99 99 99 99 99 99

99 9 9

72.9

84.5

72.3

85.i

78.s

89.6

69.3

85.2

71.3

89.3

64.4

78.9

87.7

Depth of the snout at the rostral protuberances in % of its breadth at the

same point.

56.8

56.3

55.3

56.3

64.i

66.4

58.9

65.i

64.8

76,c

64.o

65.3

74.o

Number of gill-rakers on the first left branchial arch

35

36

35

36

29

32 j

31

33

25

25

23

24

25

Not even in the averages do we here find any
constant difference, except in the number of the gill-
rakers; and even the form difference which in the
case of adult specimens is the most trustworthy di-
stinction between bldsikar and fetsikar, the different

depth of the snout, proves to be a character of age.
But in the last-mentioned respect, where the percent-
ages rise with age, as well as in other characteristic
features, e. g. the relative length of the lower jaw,
where the percentages sink with increasing age, we

GWYNIAD.

907

line! that, where a change of growth is clearly defined
in tendency, the fetsik ( Lavnretus ), at the same size as
the bldsik , has attained a more advanced development.

The sexual differences show that in the males the
length of the abdominal part is generally less than in
the females, the anal tin thus beginning comparatively
farther forward, and the snout broader but longer, the
males being thus ranged as representatives of the
earliest developmental stages of the type. In the dif-
ferences of sex, which may easily be traced in the

above table, the bldsikur , however, often follow a dif-
ferent rule from the fetsikcir, and consequently, when
the former become manocentrous, or the latter pycno-
centrous, the general rule for the sexual difference is
commonly reversed, a circumstance which most readily
suggests the presence of hybridism".

Guided by the alterations of development and the
differences of sex included in the preceding table, ive
may easily explain the form-series, as it appears in
the following table of averages:

.

to

O

Nilssonii

Wortmanni

Lavaretus

O

Microps

S to

Average

gT*

1 ^

s*

Co

I

Co‘

pycno-

centrus.

e> _

c* 2

s 0

pycno-

centrus.

mano-

centrus.

3

pycno-

centrus.

O -o

S. ®

g O

Cl"
§* O
^ 1

Cl K>,

g 0

maiio-

ccntrus.

C

Cl

1 !

1

Number of specimens measured.

7

6

17

33

4

17

15

5

24

101

14

5

18

14

6

Length of the body expressed in millimetres

420

358

192

234

182

235

285

371

297

244

349

383

371

325

321

Number of gill-rakers on the first left branchial arch...

53

45

32

35

27

32

24

28

34

25

29

34

25

21

19

Length of the head in % of the length of the

body

21.4

19.5

19.9

20. 0

20.4

19.9

19.3

22.o

19.6

19.9

17.9

18.8

19.7

19.4

17.4

,, ,, reduced. ,, „ ,, ,, ,, ,, ,,

15.8

14.4

15.o

14.8

15.2

15.0

14.3

I6.1

14. c

14.9

13.i

13.8

14.5

ll.i

12.6

Distance between the dorsal fin and the tip of

the snout „ ,, „ „ „

41.8

44.4

43.6

43.4

43.4

43.2

44.1

46.4

43.9

43.7

43.2

43.9

44.9

43.7

41.9

Base of the dorsal fin ,, „ ,, ,, „ „ „

5,

11.7

11.7

11.0

ll.i

10.8

11.2

11.3

10.9

11.4

ll.i

10.9

11.6

10.8

11.7

12.7

Height ,, ,, ,, ,, ,, ,, ,, ,, ?? ??

„

15.2

16.2

15.2

15.7

16.9

13.3

15.o

I6.0

15.3

14.8

13.i

ll.i

15.2

17.9

16.9

Length of the pectoral fins. „ „ ,, „ ,, ,, „

15.o

15.4

14.2

14.7

15.o

15.3

14.3

15.7

14.6

14.3

13.3

13 9

15.2

14.6

15.i

Preabdominal length ,, „ „ „ „

„

2G.7

29.3

28.o

27.9

26.o

28.6

28.6

31.5

28.9

28.5

31.i

29.6

24.9

28.g

28.4

Distance between the ventral fins and the tip of

the snout „ „ ,, „

47.3

48.9

47.3

47.2

46.i

47.5

47.7

52.3

48.3

48.2

48.5

47.3

49.2

47.9

46.i

Length of the ventral fins ,, ,, „ ,, „ ,, „

„

14.4

14.7

13.9

14.4

14.6

14.3

13.5

14.3

ll.i

13.7

12.7

13.3

ll.i

14.8

14.6

Postabdominal length ,, „ „

„

26.2

25.8

24.8

25.6

25.7

25.6

26.o

23.2

25.5

25.3

25.8

26.3

25.5

26.2

27.7

Distance between the anal fin and the tip of

the snout ,, „ „ „ ,, „

72.5

73.0

70.5

71.i

72.0

71.6

72.5

73.8

72.6

72.3

72.7

72.7

73.3

72.s

72.4

Base of the anal fin. „ „ „ ,, „ „ ,,

' „

10.5

ll.i

11.8

11.8

11.6

ll.i

11.0

10. G

11.0

10.9

10.5

ll.i

10.7

11.2

11.5

Height ,5 ,, ,, ,, ,, 5, ,, 5, ,, ,5 55

„

10.9

10.2

9.8

10.4

10.5

10.1

9.7

11.1

9.9

9.5

8.5

9.5

10. 0

11.6

11.2

Dorsal margin of the peduncle of the tail „ „ „ „ ,, „ „

,,

9.5

9.5

9.2

9.4

9.7

9.4

8.9

7.9

8.7

9.3

9.7

8.5

8.8

9.1

8.4

Length ,, ,, ,, ,, ,, ,, — ,, ,, ,, ,, ?? 55 ,5

„

13.4

13.2

13.8

13.2

14.o

13.3

13.2

12.5

13.5

13.3

13.8

13.3

12.9

13.i

12.8

Ventral margin „ „ „ „ „ „ - „ „ „ „ „ „ „

8.9

8.6

9.3

9.0

9.4

9.3

8.8

7.9

8.6

9.0

9.6

8.7

8.3

8.0

8.1

Least depth ,5 .5 55 ,, ,, 55 55 55 55 ?? ?? ??

9?

7.3

7.8

6.7

7.3

7.2

7.1

7.0

7.2

7.1

6.9

6.5

7.2

7.2

8.0

7.8

Length of the middle caudal rays „ „ ,, „ „ ,, „

n

5.2

5.2

5.3

5.8

5.6

5.8

5.2

6.1

5.4

5.6

4.8

4.9

5.4

5.4

5.6

„ „ „ longest rays in the upper caudal lobe „ „ „ ,, ,, „ „

„

17.0

18.5

18.6

19.3

19.3

19.o

18.3

18.8

17.9

I8.1

15.6

17.3

17.3

19.i

I8.1

?? ?? ?? snout 55 55 55 55 55 55 55

head

27.0

25.7

24.5

24.9

24.1

25.2

25.7

28.i

26.4

26.4

26.9

27.o

28.i

27.1

25.7

Longitudinal diameter of the eves „ „ „ ,, ,, ,, ,,

n

16.2

18.4

25.5

23.6

26.3

23.1

21.2

19.4

20.n

22 2

19.o

18-3

18.2

19.6

20.5

\ ertical ,, ,, ,, ,, 55 55 55 .5 55 55 55

„

14.0

I6.0

21.8

20.4

23.0

19.9

I8.0

17.o

18.3

19.7

16.7

16.4

15.7

17.8

I8.1

Breadth of the snout at the rostral protuberances .. „ „ ,, „ „ „ ,,

18.0

16.4

16.5

16.6

16.5

15.5

15.1

20.3

15.6

16.2

16.9

16.7

15.9

I6.0

14.6

Depth ,, ,, ,, ,, ,, ,, ,, .. ,, 55 ,5 ,5 55 55 55

10.5

9.3

9.2

9.3

9.4

9.9

10.1

13.9

11.6

11.2

12.2

11.3

12.7

12.3

12.6

Breadth of the forehead at the middle of the eyes.. „ „ „ „ „ „ „

9J

27.8

30.6

30.i

30.2

31.2

29.6

29.3

30.1

29.0

29.4

31.5

31.4

29.0

32.1

31.3

Length of the maxillaries - „ „ „ „ „ „ „

9»

31.7

27.8

27.9

27.6

28.i

28.i

27.3

32.3

27.8

28.i

27.6

27.1

28.6

26.7

24.2

„ lower jaw ., „ „ „

9 9

44.8

44.4

44.7

44.7

43.o

44.2

42.o

41.7

43.4

42.4

41.7

40.9

40.9

38.7

36. g

„ suboperculum „ „ „ „ „ ,, „

„

29.3

29.5

30.1

30. 0

31.8

29.9

29.9

28.0

29.i

29.5

29.9

29.4

28.6

30.3

30.5

Vertical diameter of the eyes in % of the length of the head reduced

19.i

21.6

28. s

27.1

30.9

26.2

24.1

23.2

24.5

26.5

22.6

22.3

21.9

24.3

24.8

Depth of the snout at the rostral protuberances „ „ „ „ „ „ „ „

99

14.i

12.6

12.1

12.6

12.6

13.i

13.9

19.0

15.6

15.o

16.4

15.4

16.9

16.9

17.4

Length of the maxillaries , „ „ „ „ „ „

43.2

3 / . G

36.7

37.2

37.7

37.2

36.5

44.1

37.4

37.3

37.5

36.9

37.8

36.5

33.4

„ „ lower jaw , „ „ ,, „ ,, „ „

99

60.9

60.1

59.1

60. 0

57.7

58.5

56.i

57.1

56.5

56.6

56.5

55.6

55.2

52.9

50.4

Least depth of the peduncle of the tail „ ,, „ „ „ „ „ „

„

46. 1

54.2

44.1

49.2

47.8

47.4

48.2

45.2

48.8

46.4

49.0

52.4

49.3

56.2

61.5

Breadth of the maxillaries in % of their length

35.4

37.4

35.5

36.7

36.o

36.8

36.7

37.5

37.2

37.4

38.8

38.o

39.1

40.5

40.7

Depth of the snout at the rostral protuberances in % of its breadth at the same point.

6O.1

56.2

55.7

56.0

57.i

63.8

69.i

68.7

75.2

70. 0

72.3

67.9

79.1

76.7

87.i

„ „ „ „ „ ,, „ „ „ ., „ the length of the maxillaries.

33.8

33.6

33.o

33.7

33.4

35.3

38.i

43.6

42.2

40.4

44.0

41.8

45.2

45.9

52.0

Length of the head reduced in % of the distance between the dorsal tin ami
tip of the snout.

the

37.7

33.1

34.2

34.3

35.0

34.8

32.7

34.5

33.3

34.0

30.5

31.4

32.5

32.4

30.2

Smitt, 1. c., pp. 281 seqq.

908

SCANDINAVIAN FISHES.

The clearest expression of the evolutional course
of the form-series we here find in the relation between
the depth of the snout at the rostral protuberances
and the length of the maxillaries: the former increases
and the latter decreases, so that the averages rise with
great regularity from left to right in the last line but
one of the table. Yet these averages, as we have men-
tioned, must be considered with reference to the size
of the specimen: among small specimens both of Nils-
sonii and lavaretus for example the manocentrous show
a smaller average than the pycnocentrous; but the law
of evolution tells us, according to the preceding table,
that if the former had been permitted to attain a
sufficient size, their percentages for this relation would
undoubtedly have risen enough to fill their place in
the series.

Bearing this result in mind, we can easily deter-
mine the systematic value of the peculiarities which
seem to characterize the Gwyniad where it lives under
exceptional conditions. In Enare Traesk for example
(Finnish Lapland) it sometimes develops the beaked
form" at a length of 14 or 15 cm. and with a depth
of snout persistently answering to the shallow snout
of the typical blasikar, though it is quite manocentrous,
with no more gill-rakers than a polcur. But to coin
a special name for such a form, is of questionable
utility, for a similar prolongation of the snout, though
not quite so great, may be observed in young blasikar ,
for example from Qvickjock, where they attain matu-
rity at about the same size'.

The relation between the two groups, the onec
collected round the bldsik type, the other** round the
fetsik type, is evidently the same as that between
trutta and salar among the Salmons. They come so
near each other that constant characters to distinguish
them cannot be adduced; the one is a more advanced
development of the other, and they intermingle in their
spawning operations *. However unlike the differentia-
tion of form may be in localities at a distance from
each other, variations as great may be observed in the
same water, the extremes of the form-series appearing
side by side. No wonder then that ichthyologists have

failed sharply to define local varieties, though a more
than adequate number of names have been proposed
to this end.

The coloration of the Gwyniad is essentially the
same as that of the Vendace, and is subject to the
same variations of light or dark tone. The back is of
a, lustrous steel-blue, whence the name of bldsik , or
black, in which case the latter colour also extends to
the upper part of the head and forward over the snout,
or, like the top of the head, of a more greenish gray
tint and more or less transparent, the transparency being
most noticeable in the occiput and the snout, though
the tip of the latter is commonly brownish black. The
sides of the body have a silvery lustre, but often pass
into a dirty gray, which colour has given rise to the
name of grdsik , and is sometimes uniformly distributed
over the middle of the sides, but generally more pro-
minent at the limits between the longitudinal rows of
scales below the lateral line. The ventral side is of a
purer white, at the middle milky white. The sides of
the head partake in the silvery lustre of the sides of
the body; but on the gill-cover this hue frequently
passes into a brassy lustre; and the articular process
of the operculum is of a brownish black colour, which
is continued back in a longitudinal stripe. The yel-
loAvish colour often extends to the cheeks and jaws.
The lower part of the head is of the same colour as
the belly. No less variable is the coloration of the
fins. Sometimes all of them are entirely black; but in
the lighter varieties they may all be light, greenish
gray or transparent. As a rule, however, they are
light (gray or dashed with red) at the base, but the
outer part of the fin-membrane is dark, grayish or
brownish black, this being due to a rarer or denser
agglomeration of brownish black pigment, which is
sometimes merely besprinkled on the tips of the pec-
toral and ventral fins. The same pigment is frequently
scattered in fine dots over the skin of the whole body7,
and usually collects on the middle or bottom of the
dorsal fin, on the upper part of the head, on the inner
(hind) surface of the pectoral and ventral fins, and
sometimes here and there on the sides of the body,

“ Smitt, 1. c., tafl. V, fig. 89.

6 Smitt, 1. c., tab. metr. X, No. 169.
c Fatio’s Coregonus dispersus.
d Fatio’s Coregonus balleus.

e Fatio has recognised a biending of characters in his “species composita”, Coregonus Suidteri ( Fne Vert. Suisse, vol. V, p. 270).
f They are sometimes entirely wanting in the light varieties.

GWYNIAD.

909

in irregular dark spots. Sometimes the pectoral fins
are of a plain grayish red, while all the others are
black. During the spawning-season the scales of the
sides, most commonly below the lateral line, are coated
with dermal tubercles, such as we have noticed in the
Cyprinoids. These tubercles, of a, white or reddish
gray colour, form longitudinal rows (see Plate XLII,
fig. 4), one to each row of scales, but they are not
present on all the rows of scales, sometimes only on
5, sometimes on as many as 13, most of them below
the lateral line. They are sometimes present both in
males and females, sometimes only in the males; and
many breeding Gwyniads, even males, in the same
shoal are without them".

The Gwyniad, as we have already mentioned, is
spread throughout Sweden; and the rule is that the
blasikar belong to fresh water, the fetsikar to the sea,
the rivers flowing into the sea, and the great lakes.
So in this respect too the relation between the two
main groups of Gwyniad forms is fully analogous to
that between trutta and salar among the Salmons. The
range of the Gwyniad probably extends over all
northern countries both of the Old and New Worlds6.
Throughout Finland and Norway the species occurs;
but in the latter country, according to Collett, its
range is interrupted between Trondhjem and Tromso,
which is also the case with several other fresh-water
fishes. In Denmark, where it is called Hcelt and Snce-
bel (the German Schndpel = Sw. ndbbsik), it is common
on the mainland, less so on the islands and round
their coasts. It is also common throughout Germany
north of the Alps — where it bears many names, Ma-
rdne, Blaufelchen, Henke, Kilch, etc. — and in the
Baltic Provinces of Russia. But in South-eastern
Europe it is wanting. In Switzerland it occurs in the
numerous forms (24 sub-species) described by Ratio.
In Savoy it inhabits the Lac du Bourget; and it has
been introduced in recent times into Lakes Maggiore
and Como on the southern slope of the Alps. In
France proper, with the exception of the most eastern
districts, it is wanting; but to the fish-market of Paris,
where it is known by the name of Outil, it is supplied
from Belgium and Holland, where in its ndbbsik form,

called Holding and Adelvisch , it inhabits the North
Sea and the Zuider Zee and ascends the Rhine. In
England it is common in Ulleswater and the other
Cumbrian lakes, up to a height of 2,600 feet above
the sea-level, being here known as the Schelly, in
Scotland it inhabits Lake Lomond, where it is called
the Powan, and in the Welsh lakes it bears the name
of the Gwyniad. The flouting form has been taken
on the east and south coasts of England. The North
American White-fish has its true habitat in the region
of the Great Lakes, but its range extends to the Arctic
Ocean. But in Greenland and Iceland the species is
unknown.

In this wide geographical extension there are three
centres where the Gwyniad attains its most robust
development: Siberia, the Baltic countries, and the
basin of the Great Lakes in North America. The first
home of the species lay in the north, and to the south-
ern tracts in the interior of Europe where it notv oc-
curs, it had free access during the Glacial Period.
But when the waterways became landlocked, and the
extent of the seas was reduced, it was compelled to
adapt itself to the various environments of its abode,
and it has thus been differentiated into a number of
forms which cannot, however, be distinguished by con-
stant characters.

“The Gwyniad’s habits,” writes Ekstrom of the
Baltic form, “differ little from the Salmon’s. Like
the latter it ascends from deep water in the spawning-
season, and is said to observe a certain order in its
evolutions.” “The roading Gwyniads,” wrote Gisler
from Norrland, “gather in shoals with such consent
that hardly a single fish can be found on the coast
between the shoals. They mostly repair to the coast
in violent gales from the south and south-east, which
drive them into fjords and estuaries. As soon as the
wind veers to the west, they ascend the rivers with
eager haste, and are then observed to shape their
course in two wings or lines converging to a point,
as is related of the Salmon c.” “In the island-belt of
Morko,” continues Ekstrom, “the Gwyniad ascends from
deep water, where it has passed the winter, in spring-
time, when the Baltic Herring spawns. It follows the

“ According to Gisler they arc wanting in the males which first reach the spawning-place.

b Bean (Oat. Coll. Fish. Exh. bij U. S. Nat. Mas., Gt. Intern. Fish. Exhib. London 1883, p. 8) and Turner ( Contrib . Nat. Hist.
Alaska, Arct. Ser. Publ. Sign. Serv. U. S. Army, .No. II, p. 104) mention Coregonus clupeiformis ( albus ) from Alaska: and as we have
stated above, we know of no specific difference between this form and our European Gwyniad.
c See above.

910

SCANDINAVIAN FISHES.

Herring-shoals, and devours their roe. Afterwards it
returns to deeper water, and does not re-appear until
the close of summer, at the end of September. Except
during the spawning-season the Gwyniad is so cunning
and cautious a fish that its sagacity has given rise to
a proverbial expression, a sly person being called an
arg sik (a thorough Gwyniad). It is also timid, and
so voracious as to devour not only the roe of other
fishes but also its own. It dies soon after it is taken
out of the water.” This greatly depends, however, on
the depth at which it is caught. When drawn up from
very deep water, it dies at once, the air-bladder swells,
and the belly is distended, sometimes to monstrous
dimensions.

The lake Gwyniads also assemble in shoals twice
a year. The first period is during summer, when in-
sect life is most thriving, gnats and dragonflies sport
at the surface or drop into the water, and when the
small crustaceans ( Entomostracn and fresh-water Garn-
maroids) are most plentiful. Shoals of Gwyniads then
seek their food at the surface and in the shallows.
Fhe other period is in autumn and winter, when the
Gwyniads gather to spawn. Similar observations have
been made in North America". In Lake Ontario, for
instance, the Gwyniad is taken early in spring far out
in the lake, in about 200 feet of water. At the be-
ginning of June it repairs in shoals to the coast in
quest of food. At the beginning of August, when the
heat of the upper layers of water is too oppressive, it
speedily retires to the deeper and cooler parts of the
lake. About the middle of October it again approaches
the coast, but on this occasion in order to spawn, an
operation which lasts from the middle of November
to the beginning of December, and is performed in
water of a temperature of about 40° Fahr. (+ 41/2 C.).
But in certain lakes the summer shoals do not appear,
for example in Lake Erie, according to Milner, where
the surface temperature sometimes rises to 75° Fahr.
(-*- 24° C.), and where the Gwyniad resorts in summer
to the deepest parts of the lake. Here, however, it is
by no means condemned to starvation.

a See Mr. P. Kiel in Be. Goode, Fisher ., Fisher. Industr. U.
h According to Beehm’s description of the fishery in the R. 0
spawn at the end of the summer, and return in August, usually at t
Obi ( Thierleben , Bd. VIII, p. 235). The Gwyniads brought home b;
Riksm. Salmon ., tab. metr. XIII, No. 424, cett.) that the eggs were
been performed in the course of the summer. In some of the very (
and August.

The largest and most developed Gwyniad forms,
with their more or less ventral mouth (situated on the
under surface of the snout), are evidently in the first
place bottom-feeders. They seize the larger Gamma-
roicks and the mollusks ( Limncea , Planorbis, Pisidium ,
etc.) which live among plants and in the mud, not to
mention, as we have already stated, that they are vora-
cious roe-eaters. Small fry also enter into their diet.
Young Gwyniads and the smaller forms, with their
more terminal mouth (situated at the tip of the snout),
are in general shore and surface fishes, and have to
content themselves with smaller prey. But in many
Scandinavian lakes, especially in Norrland and Lapland,
small crustaceans ( Entomostraca ) and gnat-worms occur
in such multitudes that even the large Gwyniads grow
fat on them.

The spawning-season occurs, as we have mentioned,
in autumn and winter6, as a rule from the middle of
October until after the beginning of January. The
young spawn in comparatively shallow water, only 1 — 3
fathoms in depth, and earliest in the year; older spe-
cimens generally breed later and in the deeper parts
of the lakes. The males usually come first to the
spawning-place, the females following them. The breed-
ing fishes display eager excitement, especially at night.
The males even cling fast with their teeth to the fe-
males, seizing them under the gills or by the pectoral

fins. Or the pair sometimes two or more males and

one female — pressed close together, run in sinuous
curves towards the surface or even above it, “belly to
belly, with their sides gleaming bright in the water,
where they strive and wriggle until they are rid of the
roe and milt, which adheres to the bottom, and on
stones or nets” (Gisler). In stormy, snowy, or very
cold weather the spawning is performed in the same
wav, but several metres below the surface. The water
round about is dyed with loose scales, and after the
spawning the exhausted fish seek out resting-places.

The eggs are light yellow, and their dimensions
vary considerably, according to the size of the mother
fish. In Udsikar from Lake King the ripe eggs measure

S., sect. I, p. 510.

i, the Gwyniads ascend the river from the sea when the ice breaks up,
ie end of the month, to the Arctic Ocean, or at least to the Gulf of
Theel and Tbybom from the Yenisei in 1876 also show (see Smitt,
so large at the beginning of July that the spawning might well have
eep Swiss lakes too, according to Fatio, the Gwyniad spawns in July

GWYNIAI).

911

l3/4 — 2 mm. in diameter, in the larger fetsikar as
much as 3 mm. At first they are highly sensitive,
especially to warmth ; when the embryo is half-developed,
they are easier of transportation. In water of a tem-
perature varying from December to April between
+ 1° and + 5° C. (34° and 41° Faiir.), the fry are not
hatched and capable of swimming until 5 months after
the deposition of the ova. Just after their hatching
Gwyniad fry are 9 mm. long, when a fortnight old,
11 or 12 mm., at the age of one month, 15 mm., and
when three months old, about 30 mm. (Norback). In
the following year the young measure about 12 — 17 cm.
From his examination of American Gwyniads Milner"
deduced the following results:

Weight of the
mother fish.

Weight of the
ovaries.

Number of eggs.

Number of eggs per lb.
in the living fish.

2 lbs.

53/4 oz.

21,229

10,614

93/

/ 4 5?

7'Vj »

28,500

10,361

4 „

16

48,100

12,000

77,

25

66,606

8,881

In the largest Gwyniad this result pretty nearly cor-
respond to Norback’s calculation for Scandinavian Gwy-
niads, that on an average each female Gwyniad has
7,000 — 8,000 eggs to each Swedish pound* of her
own weight.

The Gwyniad does not yield to the Salmon as a
food-fish. In many parts of Sweden, especially in Norr-
land, it is more important to the fisherman. It is
taken principally with net and seine, but also on long-
lines, and at the spawning-places the leister is em-
ployed. The small Gwyniads of Lake Wetter that
frequent the shallower parts of the lake are netted
with the so-called stronot (= strewing seine , but really
a net, being without pocket), which is plied in the
same way as the trammel-net c, with a beater, but
without any outer net. One end is made fast by
the shore or on a shoal, and the fisherman rows the
other end of the net in a spiral with many curves,

one within another. In the Gulf of Bothnia Gwy-
niads are trapped in large ryssjor, with mouths and
hoops a fathom wide and with a land arm some-
times a thousand feet long, extending out from the
shore to deep water even where the beach is shelving.
On the Swedish coast these traps are called Finnish
storryssjor , from the country where they were first
used, and they are considered to be the most effective
engines of all, for not only Gwyniads, but also every
other kind of fish, especially Salmon — and sometimes
a seal or two — enter into the catch. But their use
readily becomes an abuse, if they close channels, or
are set at the mouths of rivers to bar the passage of
the ascending fish. In Lake Wener the long-line is
also employed; it may be baited with worms, shellfish,
or small Gammaroids. But in the Lake of Constance
Gwyniads take a hook baited with nothing but a black
horse-hair, which is bent so as to have some slight
resemblance to a fly. When fishing for Gwyniad, how-
ever, the angler should be cautious, for the fish struggles
violently to escape, and often tears its mouth loose
from the hook.

In flavour the large fetsik may be mistaken for
Salmon, if boiled fresh and served with suitable sauce.
Smoked fetsik too is a dainty food. The large bldsik
is also excellent eating — the muksun in particular —
and the Lake Ring Gwyniad, small as it is, is com-
mended for its fine flavour; but it will not bear keeping.
Salted Gwyniad is common in the markets of Norrland,
and in this form or dried the Gwyniad is an important
winter food among the Lapps.

The Gwyniad fisherman suffers greatly from the
depredations of the seal, which appears, says Ekstrom,
to be highly partial to the flesh of this fish. “The is-
lander often finds his nets stripped by the seal, and
considers that he has got off cheaply, if the nets are left
whole. When once the seal has found a net, and been
allowed to make a good meal there, it often repeats the
visit, and unless the net be removed, it comes every
night without fail for its share of the fisherman’s take.”

a See Brown-Goode, 1. c., p. 519.
l> A Swedish pound is about 15 oz.
c See above, p. 741, note a.

Scandinavian Fishes.

115

912

SCANDINAVIAN FISHES.

Genus ARGENTINA".

Jaws toothless , but the palate armed in front ivitli a semicircular transverse row of small but pointed , recurved
teeth , and the tongue furnished within its fleshy rim with a similar , but sparser row of somewhat larger teeth.
Length of the maxillaries at most about 30 % (26 — 291/2 %) of that of the head reduced , and the length of the
lower jaw at most about 50 % (43—4 91 j 2 %) of the same. Breadth of the snout at the articular knobs of the
maxillaries perceptibly less than that of the interorbital space, which is more than 1/i (26 — 29 %) of the length
of the head. Base of the dorsal fin less than 8 % of the length of the bodyb, or than half the length of the
head reduced. Pyloric appendages little or moderately developed. Scales large, for the most part curved , and
armed with small spines. About 60 (58) — 70 scales in the lateral line, which is complete, and the scales of
which are rendered more or less heart-shaped by an indentation at the hind margin.

In the genus Argentina we find reminiscences both
of the Smelts and the follotving family: a certain de-
gree of transparency and cucumber-like smell (not
quite absent, however, in the Gwyniads) and a com-
paratively small0 number of pyloric appendages point
to the former; the stiff, but fragile tin-rays and the
singular shape of the scales remind us of the latter.
The last-mentioned peculiarity, which is also connected
with the formation of the longitudinal ridges that
appear on the sides of the body in Argentina, and are
each covered by a row of curved scales indented at the
hind margin (similar in this respect to those of the
lateral line), ranges Argentina beside the extinct genus
Osmeroides of the Cretaceous Period**, though in other
respects the latter genus, with its numerous branchio-
stegal rays and its strong, toothed jaw-bones, was
more like the Salmons. In Argentina the jaws are
comparatively weak, and there are no jaw-teeth, while
the maxillaries are without supplementary (jugal) bones.
But the most striking among the other characteristics
of Argentina is the reduction of the rostral region
proper, which both in the Salmons and the Gwyniads
is sometimes so considerably elongated. Here, on the
other hand, the ethmoidal region, with its lower cover-
ing-bone, the vomer, is well-developed and compara-
tively long; but the intermaxillaries are so greatly
reduced that the head of the vomer, in almost the

same manner as in the Eels, seems to form the firm
anterior margin of the mouth. Lastly we will mention
a characteristic which is indeed not quite absent in the
Gwyniads, but is there much less developed, almost
rudimentary. Outside (under) the pelvic bones and in
front of the insertions of the pectoral fins, the large
ventral scales form a flap, free behind, which covers a
great part of the ventral fins when they are folded.

Argentina belongs to the deep-sea fishes — as the
large eyes indicate — but not to their most character-
istic types. The young are frequently met with in the
upper marine zones, and even ascend, according to
Nilsson0, to the mouths of rivers. In the depths of
the ocean the genus probably has an extensive range.
It has longest been known, for a long time exclusively,
from the North Atlantic and the Mediterranean ; but
in 1878 it was found off the coast of New Zealand, and
this in a form that can hardly be distinguished from
one of the Atlantic species. Such a find as this most
naturally suggests the possibility of discovering the spe-
cies in intermediate localities as well; but it is remark-
able that New Zealand is also the only region in the
Southern Hemisphere which possesses among its auto-
chthonic fresh-water fishes a member of the Salmon fa-
mily (Retropinna Richardsonii ).

Only two species of the genus Argentina are
known with certainty* 6 7.

“ Artedi, Ichthyol. , Gen. Pise., p. 8.

6 In a damaged specimen Brown-Goode and Bean found the length of the base of the dorsal fin to be 8'4 % of that of the body.
c According to Nilsson, however, sometimes 20.

d Agassiz, Reck. Poiss. Foss., tome V, lime Partie, p, 105, PI. 60 b.
e Observationes lchthyologicce , p. 7; Skand. Fna , FisJc., p. 476.

f Valenciennes’ Argentina leioglossa (without teeth on the tongue) from the Mediterranean off Algiers has already been identified by
Giglioli with A. sphyrama, and Hutton’s Argentina elongata from New Zealand is based on a single young specimen in bad condition (see
GtiNTHER, Deep Sea Fishes, Chall. Exped., p. 218).

SIIL-SMELTS,

913

THE GREATER SIIL-SMELT.

ARGENTINA SILUS.

Fig. 229.

Base of the anal fin , as well as the longitudinal diameter of the eyes , less than half the length of the head re-
duced. Height of the dorsal fin less than 2/s of the preabdominal length. Lower posterior margin of the oper-
culum straight. Gill-rakers on the first branchial arch about 20. Branched rays in the pectorals 16 or 17, in

the ventrals 11 or 12. Scales of a horny yellow.

WM

Mil

’

Fig. 229. Argentina situs , 9) 1 Y24 the natural size. Taken on a Fladdock-line at a depth of 50 fthms., between the Roster and Tisler
Is., on the 11th Nov., 1880; C. A. Hansson. a , scale from the base of the dorsal fin; b, scale from the lateral line; c, scale from

the first row below the lateral line.

All the scales from the right side of the body and twice the natural size.

2 3—4

R. br. 6 ; D. — ; A. — ;

9“ 10—12’

P.

16 — 17

V.

11 — 12

6. x +■ 1 + 17 + 1 + x\ L. lat. 66 — 70; L. tv. —1; Vert. 65® — 68c.

3

Syn. Salmo Situs, Ascan., Icon. Rer. Nat., fasc. Ill, p. 3, tab.

XXIV; Cuv. ( Uoregonus ), Regn. Anirn., ed. 2, tom. II, p.
308; Nilss., Prodr. Iclittiyol. Scand., p. 19; Id. {Argentina),
Observ. Iehthyol., p. 3; Kroy. ( Acantholepis ) in Gaim.,
Voy. Scand., Lap., cett., Zool., tab. 17; Id., Damn. Fisk.,

vol. Ill, p. 98; Cuv., Val., Hist. Nat. Poiss., tom. XXI,
p. 421; Nilss., Skand. Fna, Fisk., p. 469; Gtbr., Cat.
Brit. Mus., Fish., vol. 6, p. 202; Olsson, Lunds Univ.
Arsskr., tom. VIII (1871), p. 6 (sep.); Coll., Forh. Vid.
Selsk. Chrnia 1874, Tillsegsh., p. 173; 1879, No. 1, p. 93;
N. Mag. Naturv., Bd. 29, p. 109; Winth., Naturh. Tidskr.
Kbhvn, ser. 3, vol. XII, p. 45; Storm, Vid. Selsk. Skr.
Trondhj. 1883, p. 29; Sm., Vet.-Akad. Handl., Bd. 21, No.
8, p. 196; Peters., Vid. Meddel. Naturh. For. Kbhvn. 1884
(1886), p. 159; Lillj., Sv., Norg. Fisk., vol. 2, p. 679.

0 More commonly 10, according to Lilljeborg.

6 According to Nilsson; 66 according to Khoyer.
c According to Lilljeborg.

914

SCANDINAVIAN FISHES.

Salmo immaculatus ( Blankesten ), Mull., Zool. Dan. Prodr.,
p. 279, vide StrAm, Skr. Naturh. Selsk., Bd. 2, H. 2 (1793),
p. 12, tab. I, fig. 1.

Situs Ascanii, Reinh., Maanedskr. Literat. Kbhvn 1833, p. 239.

Argentina syrtensium , Br. Goode, Bean, Proc. U. S. Nat. Mus.,
vol. I (1878), p. 261; vide Gthr, Deep Sea Fish., Chall.
Exped., p. 217.

Obs. In the preceding- pages (see above, p. 59) we have em-
ployed the name of Blanlcesten, which StrAm first applied (in Sond-
mores Beskrivelse ) to a fish of the genus Sparus, as a synonym of
Fldckpagell (the Common Sea-Bream), following 0. F. Muller and
Brunnjch. But as StrAm has subsequently come to the conclusion
that the former name properly belongs to the species now in point,
it is best omitted in the said passage, especially as its application in
Norway seems to be vague.

The Greater Siil-Smelt attains a length of nearly
half a metre". The body is elongated and fairly thick,
the greatest depth, which occurs at the beginning of
the dorsal lin, being in adult specimens about 18 % of
the length, and the greatest thickness about 2 / 3b of the
greatest depth, or slightly greater than the postorbital
length of the head. The dorsal and ventral contours
form about equal curves. They meet in a point at
the tip of the snout, and converge behind till, in front
of the caudal tin, the least depth of the body measures
about 6 1 3 % of its length or '/3 of the length of the
head reduced. In transverse section the body forms,
according to its degree of distension, a more or less
regular rectangle, the back and the belly being broad,
the former more so, and flatly convex or even flat,
and the sides more or less compressed. A further
characteristic, which we have already mentioned, are
several longitudinal, low ridges. Two of these run,
parallel in front, converging on the tail, on each side
of the dorsal margin and on the uppermost part of the
sides, the lower starting from the upper angle of the
operculum; two corresponding ridges follow each side
of the ventral margin and the loAvest part of the sides,
the upper starting from the pectoral fins, the lower
from the ventrals; one ridge is formed by the lateral
line; and the median line of the dorsal margin between
the dorsal and adipose flns, as well as of the ventral
margin between the ventral tins and the anal aperture,
rises in a similar ridge.

The head forms a four-sided wedge, but is so
contracted below as strongly to remind us of that of
the Capelin. Its upper and lower surfaces converge for-

wards to the shallow, but rather broad tip of the snout,
which is rounded in an horizontal direction. Its sides
are flat, even the upper, the interorbital part of which
is depressed between the somewhat tumid supraorbital
parts; and at the articulations of the lower jaw a break
is formed when the mouth is closed, the upward slope
of the lower jaw being somewhat sharper than the
downward slope of the snout. A thick adipose mem-
brane with numerous ducts and pores is spread from
the occiput and forehead down over the operculum,
the preoperculum, and the suborbital bones, and for-
ward on the snout, surrounding the large orbits both
in front and behind. The eyes occupy about 1/3 (33
— 30 %), the postorbital part rather more than 2/5
(42 — 46 %), and the snout about 29 — 30 %, of the
length of the head. The nostrils lie about half-way
between the tip of the snout and the eyes. The an-
terior is round; the posterior, which is larger and more
oblong, transversely set, is entirely covered by a semi-
circular dermal flap from its anterior margin. The
breadth of the snout at the nostrils is only slightly
less than the i nter orbital width; but the breadth across
the rostral protuberances (articular knobs of the ma-
xillaries) is only about 1/2 — 2/3 thereof. The rostral
protuberances are indistinct. The preoperculum is di-
stinguished by its rectangular shape and its long arm,
which runs in a forward direction, coasted by the nar-
row interoperculum, which is of the same length. The
operculum is trapezoidal, but its hind margin is sinuate.
Its lower margin, which forms the fairly straight and
obliquely ascending suture with the suboperculum, is
almost equal in length to the diameter of the eye. The
suboperculum is of uniform breadth and rounded at
the lower posterior angle. The mouth is most like that
of the Grayling, but the point of the lower jaw projects
a little beyond the tip of the snout, and is furnished
with a distinct, though small, symphyseal protuberance
above and a small chin-protuberance below. The lips
are not very fleshy. The curved anterior margin of
the upper jaw is entirely formed, right across the mouth,
by the narrow intermaxillaries, which are without nasal
process, the upper jaw being thus incapable of protru-
sion. The maxillaries are narrow in front, so far as
they are covered by the intermaxillaries, but behind

“ Sometimes 2 Sw. ft. (59 cm.), according to Nilsson. StrAm’s specimen was 43 cm. long; our largest specimen, which has lain
for several years in spirits, has now shrunk to a length of about 47 cm,, but measured when caught 48 1/2 cm. according to Mr. C. A.
Hansson; and the largest specimen Collett had seen was 47 cm. long, in each case to the extreme tip of the caudal fin.
b According to Kroyer, however, sometimes hut slightly more than I/2-

GREATER SIIL-SMELT.

915

they expand in a downward direction so greatly that
their breadth is about 28 % of their length, which
measures about 20 — 23 % of that of the head or 28 —
29 V2 % °f that of the head reduced. The length of
the lower jaw is about 8fe- — 83/4 % of that of the body,
32- — 34 % of that of the head, or 44 — 48 % of that of
the head reduced. The dentition of the mouth we
have already described. The most striking characteristic
is that, at the extreme front of the palate, the head of
the vomer and the anterior extremities of the palatine
bones, which touch the head of the vomer on eaeh
side, form an arch furnished with a row of small, bent,
subulate teeth of uniform size. This arch has the ap-
pearance of an inner upper jaw, compensating the lack
of teeth on the upper jaw proper. In old specimens
these teeth are more scattered, but longer and more
villiform. The teeth on the tongue also seem to become
more scattered with age: in the semi-elliptical row we
have counted 10, 8, and 6. There is no palatal fold
in the usual place, just behind the jaws ; but behind
the head of the vomer and the incurved ends of the
palatine bones the skin of the palate lies in a pad-like
transverse fold. The gill-rakers are set in one row,
and resemble pointed, rather long, triangular disks,
which are destitute of the small spines that commonly
fringe them in the Gwyniads. The pharyngeals are
armed with several rows of subulate teeth. The pseudo-
branchim are large. The gill-openings are of consider-
able size, each extending from the occiput on a level
with the upper orbital margin and almost in a line
with the hind margin of the preoperculum down to a
line with the centre of the eye. The two branchiostegal
membranes are free both from each other and the
isthmus. The branchiostegal rays are six in number,
the last five being sabre-shaped and gradually increas-
ing backwards both in length and breadth, the first
narrow and styliform, but curved like the others.

The dorsal fin is trapezoidal, but its upper posterior
margin is more or less convex. The first two rays are
simple; the first branched ray is the longest, being
twice as long as the first simple ray, but only slightly
longer than the second. The fin begins at a distance
from the tip of the snout measuring about 41 or 42 %
of the length of the body, and the length of the head
is about 60 or 61 % of the said distance. Its base is
less in our specimens than 8 % (about 73/4 %) of the
length of the body, or about 30 — 32 % of the length
of the head- Among the preceding members of the

Salmonoid family the Smelts alone have so short a
dorsal fin. The greatest height of the fin is about
twice the length of its base. The anal fin is also tra-
pezoidal, but lower and at least somewhat longer than
the dorsal. At its anterior margin there lie three or
four simple (undivided) rays, the first quite short, about
half as long as the second, the others gradually in-
creasing in length to the hindmost, which is about equal
in length to the first branched (longest) ray, or in other
words to the base of the dorsal fin. The distance be-
tween the beginning of the anal tin and the tip of the
snout is about 78 % of the length of the body, and
the length of its base is about V3 (33 — 31 %) of that
of the head. The adipose fin, which resembles that of
the Salmons, lies above the 4th — 6th or 5th — 7th
branched ray in the anal. The caudal is deeply forked,
the length of its middle rays being only about 1/3 of
that of the longest ones, or about 4 % of that of the
body.

The pectoral fins resemble in shape and position
those of the Gwyniads. Their length is about 14 — 15
% of that of the body. The first (simple) ray does not
reach quite to their tip; the second or third branched
ray is the longest. The blunt ventrals also remind us
of the Gwyniads, but are distinguished by their nu-
merous rays — a characteristic generally belonging to
the lower (earlier) grades of differentiation among the
Teleosts — and by the covering lobe, formed by large
scales on the ventral side in front of their base, under
which they may be concealed to a great extent. Their
length is about 1 10 — 1/9 of that of the body, or 3/4
(78 — 74 %) of that of the pectorals. They are set half-
way along the body (at a distance from the tip of the
snout measuring about 49 — 51 % of the length of the
body), exactly below the termination of the dorsal fin
or a little farther back. The preabdominal length is
less than in most other Salmonoids (24 — 26 % of the
length of the body), but the postabdominal length is
greater (29 — 27 % of the same).

The shape and texture of the scales, which are
rather thick but flexible, vary according to their situa-
tion. They are densely imbricated, and inserted so far
into the follicles that most often only 1 4 of their sur-
face is uncovered by the next scale in front. Their
exposed part is closely set with small spines, pointing
in a backward direction, which render the body as
rough to the touch as in many Sharks. On the other
hand, this part of the scale is without concentric stria;,

916

SCANDINAVIAN FISHES.

or, if these are present, they are faint or at least scat-
tered, while on the anterior (inserted) part of the scale
they are all the more numerous. Radiating grooves
there are none; but the anterior margin of the scale
in particular is often sinuate. The nucleus of some
scales, for example on the forepart of the back and of
the belly, is so large as to occupy the greater part of
the scale, with only a comparatively small number (20
— 30) of concentric striae in front of, above, and below
it. But in other cases, for example in the large scales
on the sides just below the lateral line (fig. 229, c),
the nucleus is all the more reduced, so that about 80
or 90 concentric striae may be counted, about each 8th
— 10th one coarser than the rest, on the inserted part
of the scale. Most of the scales are irregularly round,
of a rounded quadrangular or broadly elliptical shape,
with the longitudinal axis set vertically; and in these
cases the anterior margin as well is arcuate, or elong-
ated in an angular form at the middle. Where they
cover any of the longitudinal ridges on the body, they
are indented, as we have mentioned above, at the
middle of the hind margin. This also applies to the
scales of the lateral line (fig. 229, b). The large scales
set in a row just below the lateral line, largest on the
forepart of the body (fig. 229, c), are distinguished by
their more rectangular shape, with straight anterior
margin and with the anterior angles right angles. Along
each side of the base of the dorsal fin lies a row
of obliquely linguiform scales, with the exposed part
obliquely triangular (fig. 229, a). We shall meet with
similar scales in the following family, as well as a certain
resemblance in the cucullate shape characteristic of the
duct in the scales of the lateral line (fig. 229, b).

The coloration of the body is determined by the
horn-yellow tint of the scales; but the inner surface of
their exposed (posterior) part is lined with a thin coat
of silvery lustre, which gives to the sides of the body
an iridescent gleam of silvery, golden, or brassy hue.
The back is darker, brownish; the bellv “silvery white,
with a reddish lustre” (Hansson). The adipose mem-
brane on the head is yellowish, but under it the cheeks
and opercula gleam with a silvery lustre. The fins share
in the general coloration, being of a grayish yellow or
paler (whitish, according to Kroyer). The adipose fin is
yellow, but at the base of the same colour as the back.

The abdominal cavity, and the pharynx forward to
the root of the tongue, are black; but the black lining
of the peritoneum has a silvery layer underneath. In all

our specimens, however, the viscera are so greatly da-
maged that we cannot describe them with any minuteness.

The Greater Siil-Smelt belongs to the deep — though
probably not to the deepest — parts of the North At-
lantic. In company with the Norway Haddock ( Sebastes )
it lives off the Norwegian coast (Ascanius, Strom, and
Nilsson) and off the east coast of North America (Sable-
Island Bank off Nova Scotia, Brown-Goode and Bean)
at a depth of about 80 — 200 or 300 fathoms. The deep
fjords of Norway afford it a constant abode. In the
Skager Rack it is occasionally found. Juke other deep-
sea fishes it is sometimes suddenly carried in some way
or other to depths where the pressure is inadequate to
its requirements, and it is then borne helplessly to the
surface, where it drifts about, unable to regain its home.
Under these circumstances it has been cast ashore on
the west coast of Jutland. It has most frequently been
met with in the neighbourhood of Bergen; but its range
extends, according to Collett, from the entrance of
Christiania Fjord along the whole coast to Tromso.
According to Storm it is not rare in Trondhjem Fjord,
and has been found, especially in late years, at several
places in the fjord, to the very head thereof. In the
Skager Rack, off Dynekil, between the Foster and Tisler
Islands, a specimen Avas caught on a long-line in No-
vember, 1890, Avhieh was forwarded to the Royal Museum
by Mr. C. A. Hansson, and which is represented in our
figure. From the SkaAv to the extreme south of the
west coast of Jutland three specimens have been found,
according to Wintiiee and Petersen, since 1871. The
species consequently seems to have the southern limit
of its range here in the deeper, northern part of the
North Sea, for farther south it has never been met Avith.

Of the Greater Siil-Smelt’s habits we know no more
than of the life led by most deep-sea fishes. As a rule
the stomach is forced into the pharynx Avhen the fish is
secured, so that nothing can be ascertained as to its food
from the contents of the stomach. But to judge by its
mouth, Avhieh calls to mind the Grayling, its dentition,
Avhieh is somewhat more poAverful than the Graylings’s,
and its fairly coarse gill-rakers, its predatory habits are as
marked as those of the said species, and we know that it
takes a hook baited with mussels ( Mytilus , according to
Nilsson) or a bit of fish (Herring, according to Olsson).
It is fat, and its flesh is said to lie of good flavour. Some-
times a score or so are exposed for sale in the fish-
market at Bergen; but else it is taken far too seldom
to be of any economical importance to the fisherman.

SALMONOIDS.

917

THE HEBRIDAL SIIL-SMELT (sw. stromsillen).

A RGENTIN A SPHYRCENA.

Fig. 230.

Base of the anal fin perceptibly greater than the longitudinal diameter of the eyes , and more than ‘/g the length,
of the head reduced. Height of the dorsal fi,n more than 2/ 3 of the preabdominal length. Lower posterior mar-
gin of the operculum indented, in an elongated 8-form. Gill-rakers on the first, branchial arch about 13 or 14.
Branched, rays in the pectorals 12 or 13, in the ventrals 9. Scales transparent, with a rich coat of silvery

pigment on the inner surface.

b

Fig.

230. Argentina sphyrcena, 9> natural size. Taken on a long-line off Helso, near Stromstad, on the 12th Dec., 1879. C. A. Hansson.
a, one of the scales in the lateral line, magn. 2 diam.; b, section of the body just in front of the ventral fins, natural size.

Pl . br. 6 :

„ 3“ , 36

1

(1)1

I) . — ; A. — ;
8 10’

l *r*1
l ^

V. — — — ; C. +14-17 + 1+ #;

Fin. lot. 58 c

; Lin. transo

3

. — 1 ; Vert.
3

51 J — 52.

Syn. Sphyrsena parva sive

Sphyrseme

secunda species, Rondel., De

Pise., lib. VIII, cap. II, p. 227. Pisciculus Rom* Argen-
tina dictus, Willughb., Hist. Pise., p. 229; Ray, Syn.
Meth. Pise., p. 108. Argentina , Art., Ichthyol. , Gen. Pise.,
p. 8; Syn. Pise., p. 17.

Argentina Sphyrcena, Lin., Syst. Nat., edit. X, tom. I, p. 315;
BrOnn., Iclithyol. Massil., p. 79; Cuv., Mem. Mus., vol. I,
p. 228, tab. 11, fig. sup.; Risso, Hist. Nat. Ear. Mer.,
tom. Ill, p. 462; Esh. in Nilss., Stcand. Ena , Fisk., p.
47 6; Gthr., Cat. Brit. Mas., Fish., vol. VI, p. 203;
Canestk., Fna D’ltal., part. Ill, p. 129; Coll., Forh. Vid.

Selsk. Chrnia 1874, Tillsegsh., p. 171; 1879, No. 1, p.
92; N. Mag. Naturv. Chrnia, Bd. 29, p. 109; Malm, Gbgs ,
Boh. Fn., p. 550; Hansson, Ofvers. Vet.-Akad. Forh. 1880,
No. 4, p. 22; Mon., Ilist. Nat. Poiss. Fr ., tom. Ill, p.
554; Day, Fish. Gt. Brit., Irel ., vol. II, p. 136, tab. CXXV;
Storm, Vid. Selsk. Skr. Trondhj. 1883, p. 29; Sm., Vet.-
Akad. Handl., Bd. 21, No. 8, p. 196; Lillj., Sv., Norg.
Fislc., vol. II, p. 692.

Argentina Silus, Nilss. (p. p.), Obs. Ichthyol., p. 7 ; Edw.,
Journ. Lin. Soc., Zooh, vol. XV (1881), p. 334.

Osmerus Hebridicus, Yarr., Hist. Brit. Fish., ed. 1, Suppl. II,
p. 16; Nilss. ( Argentina ), Skancl. Fna, Fisk., p. 474;
Gthr., Cut., 1. c.

Argentina Cuvieri, Cuv., Val., Hist. Nat. Poiss., vol. XXI, p.
413 + Arg. leioglossa, p. 417, tab. 624 (vide Giglioli,

1—2

a - according; to Day.

8— 9 &

9 .9.

b _£ A

9— 10 ” ” ”

c 52 — 53, according to Nilsson and Gunther.
d Sometimes 50, according to Nilsson.

918

SCANDINAVIAN FISHES.

Espos. intern. Pesca, Berlin 1880, Sez. Ital., p. 102) +
Arg. yarrelli, p. 418.

Goniosoma argentinum, Costa, Fna Regn. Nap., Malacotterigii
Abdominali, Genere Goniosoma, tab. XXXVI.

Argentina decagon , Clarke, Trans., Proc. N. Zeal. Inst., vol. XI
(1878), p. 295, tab. XIV, fig-, infer, (vide Day, 1. c.).

(?) Argentina elongata, Hutton, Ann., Mag. Nat. Hist., ser. 5,
vol. Ill, p. 53; G^chr, Deep Sea Fish., Chall. Exped., p.
218, tab. LV, fig. B.

The Hebridal Siil-Smelt attains at most about half
the length of the preceding species". It is besides, in
most cases, of a more elongated form, the greatest
depth in adult specimens — with the exception of gra-
vid females — being about 13 % of the length of the
body; but in this respect there is no constant distinc-
tion In other respects too both species are so like
each other that the essential differences are but few.
The Hebridal Siil-Smelt has a comparatively longer anal
fin, a higher dorsal fin, smaller eyes and less developed
adipose membrane on the head — it is no true deep-
sea fish — fewer rays in the pectoral and ventral fins,
fewer scales, vertebras, and gill-rakers, but a somewhat
longer peduncle of the tail (behind the adipose and the
anal fins). To these we should add a characteristic
first remarked by Lilljeborg, the faint S-shaped cur-
vature of the lower posterior margin of the operculum
in the Hebridal Siil-Smelt. But in coloration this spe-
cies differs widely from the Greater Siil-Smelt, owing
to its thin and more silvery scales, most of them de-
ciduous.

The length of the head in adult specimens is some-
what less than * 1/i of that of the body, the longitudinal
diameter of the eyes rather more than ||4, and the
postorbital length about 2/5, of the length of the head.
The forehead is flat, the supraorbital parts being hardly
at all tumid, but their outer margins indented in a
rounded manner above the middle of the eyes. The
nostrils lie rather near to the orbits, and they are of
fairly equal size, or the anterior is even a, little larger
than the posterior. They are circular, or oblong in the
longitudinal direction of the body. The breadth of the
snout at the nostrils is perceptibly less than the width of
the interorbital space, and its length is about equal to the
longitudinal diameter of the eyes. The jaws are relati-
vely somewhat shorter than in the Greater Siil-Smelt,

the length of the maxillaries being about 19%, that of
the lower jaw about 3072 %, of the length of the head.
The transverse row of palatine teeth is sharply defined
at both ends, the margin of the palatine bones being
indented just behind them. The gill-rakers are not
only more scattered but also comparatively shorter
than in the preceding species. The pseudobranchise
are rather large here as there.

The height of the dorsal fin is about equal to the
length of the head reduced, or nearly 18 % of the
length of the body, and perceptibly more than twice
the length of the base of the fin. Here too the anal
fin begins at the termination of the third quarter of
the length af the body, but the length of its base is
nearly 2/5 of that of the head. The adipose fin begins
above the 2nd or 3rd branched ray in the anal.

The pectoral fins, the length of which is about
13 7 2 % of that of the body, are rather shorter, the
ventrals, on the other hand, somewhat longer, than in
the preceding species, the length of the latter being
only about Vio Gss lhan that of the former.

The form of the scales varies as in the preceding
species, but their spines are less developed.

In fresh specimens, according to Edward, the
body is so transparent that on holding it up to the
light the vertebra? can be distinguished. The divisions
of the brain are even more distinctly visible through
the parietal and frontal bones than in the preceding
species. The back is of an olive gray, the sides and
operc.ula are silvery, the former, however, according to
Edward, with different shades of colour between the
several longitudinal ridges. The space between the two
ridges nearest to the back is of a deep amber, the
next strip greenish blue, the third space, just below
the lateral line, silvery white with a bright metallic
lustre; the lower spaces are similar to the upper, but
their colours are fainter. The belly is then grayish white,
with a dash of greenish blue and purple. The fins are
for the most part transparent, the dorsal pale gray, the
anal white, the caudal of a deeper gray, with “a dark
longitudinal mark along either lobe near to its outer
edge” (Day). The paired fins are pale, “of a faint rose
colour” (Moreau). “A black spot on the upper edge
of the orbit, and a smaller one on the snout” (Day).

“ According to Moreau the ordinary length of this species in the Mediterranean Sea is 14 — 20 cm. Collett’s largest specimen
from Norway was 265 mm. long.

1 Collett states as a change of growth that, in young specimens (with the length of the body to the end of the caudal lobes less

than 15 cm.), the greatest depth is about '/g, in the oldest specimens (more than 21 cm. long) about 1,/6, of the said length of the bodj7.

HEBRIDAL SIIL-SMELT.

919

The abdominal cavity of this species too is black,
but the pharyngeal cavity white with a coat of silver.
The black pigment of the abdominal cavity, which here
too rests on a layer of silvery colour, extends to the
stomach, which bends abruptly downwards and forwards
in about a line with the tips of the pectoral tins. Close
up to the diaphragm the intestine bends no less abruptly
downwards and backwards, to the right of the liver,
which is comparatively small, and it is there furnished
with rather long and thick appendages, varying, as it
appears, in number. Nilsson counted 14 — 20 of these
pyloric appendages, we have found 12, and Moreau
found 10 — 12; Wjlllughby (Ray) counted 6 or 7, and
Day only 5. The intestine then runs in a straight
line to the vent, just in front of the anal tin. The
gall-bladder is thin- walled and oblong. The spleen is
triangular and lies behind the stomach. The air-bladder
is long and fusiform, pointed at both ends, and seems
here, as in the preceding species, to be without con-
nective duct with the oesophagus. It has long been
remarked for the rich, highly lustrous coat of silvery
pigment both without and within its thick wall.

In the North the Hebridal Siil-Smelt is best known
as an inhabitant of Christiania Fjord, where it was also
first observed, by Esmark. From this fjord, says Col-
lett, where it is caught in Sprat-seines and nets, some-
times in hundreds at a single haul, it is brought al-
most daily during autumn to the Christiania market.
In winter solitary specimens are hooked on the clayey
bottom. It was in this manner that the specimen re-
presented in our figure was secured, the only authen-
ticated instance of the occurrence of this species on the
Swedish coast. The said specimen was caught on a
Haddock-line off Helso, 3 miles from Stromstad, about
Christmas, 1879. Along the Norwegian coast the spe-
cies has been met with at several places, though in no
great number, up to Trondhjem Fjord, where it bears,
according to Strom, the name of Stavsild. Stromsild
( Stream Herring ), the name by which it is known in
Christiania Fjord, is derived, according to Nilsson,
from the circumstance that “the young at least, like
young Herrings, at certain seasons enter the mouths
of the rivers, and are found at the outlets of the streams. "

On both sides of Scotland and off the Yorkshire coast
solitary specimens have been taken; but the species is
most common and has longest been known in the Me-
diterranean, on the coasts of Italy and France. As we
have already mentioned, it has also been found on the
coast of Algiers; and it apparently belongs to the New
Zealand fauna,, far from its European home. Its bathy-
metric range extends to a depth of at least 200 fathoms,
for Malm received three specimens which had been
found in the stomach of a Ling caught at that depth
on the fishing-bank of Storegg off the Norwegian coast.

In the stomach of the Hebridal Siil-Smelt Edward
discovered small crustaceans and Sertularice, Collett
Anneloids. But as it is also hooked on lines set for
Haddock and the smaller Codfishes, it apparently feeds
like them on mollusks and, probably, on small fishes.
It is a sociable and lively fish, and in the seine it
wriggles till the scales fall off; but after this, says
Collett, it floats about helplessly at the surface. Of
its spawning-season widely different accounts are given.
According to Risso it, breeds in the Mediterranean in
spring, and appears on a sandy bottom in April. Yar-
rell mentions a female, 8 inches in length and full of
roe, that was taken on the west coast of Scotland in
June. On the east, coast of Scotland Edward caught
a male fairly full of milt in October. Collett received
from the neighbourhood of Stavanger a gravid female
taken in June; but in Christiania Fjord he found the
females full of roe in October. The female caught by
IIansson had extremely small eggs in December, but
did not seem to have spawned in the course of the
autumn, in which case the ovaries would probably
have been still more shrunken".

In Scandinavia the Hebridal Siil-Smelt has never
been put to any practical purpose, except perhaps as
bait on a long-line or hand-line. Its flesh is good,
according to Risso, but its cucumber-like smell per-
haps uninviting. Greater advantage has been taken of
this fish in Italy. There the silvery pigment of its
air-bladder and scales has long been collected to afford
material, under the name of essence d'orient, for the
manufacture of imitation pearls, as we have above
described when treating of the Bleaks.

“ Cf. Smitt, Riksmuseets Salmonider , tab. inetr. VII.

116

Scandinavian Fishes.

920

SCANDINAVIAN FISHES.

Fam. SCO PE LID M.

Form of the body variable, generally Herring-like but also irregular (. strongly compressed and deep). Scales, where
present, thin , middle-sized or large. Sides of the body (in all the Scandinavian forms but onea) furnished with
luminous spots. Dorsal margin of the tail furnished with an adipose fin. Margin of the upper jaw formed
either by the intermaxillaries alone or partly (behind) by the maxillaries as well. No barbels. Air-bladder, where
present, simple and without connexion with the cranial cavity. Branchial cavity (in all the Scandinavian forms )

furnished with pseud obranchiw. Ovaries with oviduct.

Most of the fishes belonging to this family have
a common characteristic which is connected with their
manner of life, and of which we have seen no instance
among the preceding fishes. Nearly all the Scandi-
navian Scopeloids and most of the others possess or-
gans by means of which they more or less voluntarily
emit a phosphorescent light, and which are therefore
known as luminous spots. As a physical reflex or a
chemical phenomenon such phosphorescence is found
in certain plants; as a real manifestation of a vital
function it is not uncommon in the animal kingdom.
Many — if not most — of the lower and the lowest
marine animals are phosphorescent; and among the
higher evertebrata the glow-worm ( Lampyris ) is familiar
to us all. In the glow-worm the light extends not
only to the fatty bodies in the abdomen, but also to
the eggs, i. e. the fat of the yolk, and consequently
this phenomenon — in the last case at least — is not
unconditionally an attendant of nervous activity. Nor
is this the case with the light that radiates from the
dermal mucus of certain Batrachians6, though the more
copious secretion thereof produced by irritation is sub-
ject to the control of the nerves. Among fishes too,
in some of the organs more highly developed for phos-
phorescent purposes, the light seems to be at least par-
tially independent of the immediate action of the will.
But in some fishes special nerves have been found
whose extension in these organs suggests that the light
radiates in the direction chosen by the fish and at the
moment when the fish desires its aid.

In their more developed forms the luminous spots
closely resemble the so-called parietal organ of lower
fishes, Batrachians, and reptiles. Together with the

said organ they have been included among so-called
ocellate organs; and where it has been declared with
certainty that a special nerve for such an organ might
be traced, their function may best be compared to that
of electric or pseudoelectric organs, i. e. that, as ner-
vous force by means of fine nerve ramifications upon
a so-called electric plate excites electricity in a finely
granular mass (a transformation of the sarcoplasma of
a muscular cell) on each side of an elastic plate (a
transformation of the rhabdia — rod -substance — of a
muscular cell), so the nervous force of the luminous
spots, by its action upon a strongly refractive and co-
agulant mucus (a transformation of cells in a slime-
gland of the skin), causes this mucus to burn and shine.

In their simplest form the luminous spots are de-
scribed by Lendenfeld0 as depressions or elevations of
the skin, scattered or distributed in rows on the sur-
face of the body, the smaller (0‘1 — 0'3 mm. in dia-
meter) without pigment, the larger (0‘3 — 0'5 mm. in
diameter) with a layer of pigment within their base.
They are then covered only by the thin epidermis, and
are furnished with nerves and fine capillary vessels
from the skin. Internally they are indeed hollow; but
the inner surface is lined on the outer part (distally)
with a layer of closely packed fusiform cells, at right
angles to the skin, and on the inner part (proximally)
with a layer of gland-tubes, arranged in a similar
concentric manner and closely packed. These tubes
are closed and rounded at the basal (proximal) end,
open towards the centre of the organ, and full of gland-
cells, which secrete the granular mass that fills the
hollow in the centre. This mass, a strongly refractive
and coagulant slime, is produced by an immediate

a The obscure and extremely rare Sadis atlanticus.
b Boie, Isis 1827, p. 726.

c Gunther Deep Sea Fish., Rep. Voy. Challeng., Zool., vol. XXII, App. B.

SOOPELOIDS.

921

transformation and dissolution of the gland-cells, which
are afterwards ejected through that end of the tube
which opens into the hollow.

From this comparatively simple structure, which
as yet reminds us of slime-glands of ihe skin, the first
step to higher differentiation is that, as the sac deepens
more and more, its inner (bottom) part is separated by
a constriction of the wall and becomes spherical in
shape, while the outer part remains more or less re-
gularly cup-shaped, a rotation-paraboloid with cir-
cular mouth, in which case the axis is vertical to the
surface of the body (fig. 231), or elliptical and more
or less elongated", in which case the axis is obliquely
set. Throughout the wall of the organ, within the
layer of pigment, lies an extremely thin, but power-
fully refractive membrane (m), which even extends over

the mouth of the organ, where we also find a double
membrane, consisting of an inner, cellular leaf ( d ) and
an outer, structureless one (c) explained as a kind of
lens, and situated within the thin, structureless epi-
dermis ( b ), which has been compared to a cornea. The
internal structure is otherwise essentially the same as
in the simpler luminous spots — though both connective
tissue and nervous elements enter more plentifully into
its composition — but the shining mass (/c) here lies
as a more or less lens-shaped body in the annular
constriction between the spherical (inner) and cup-
shaped parts of the organ, forming a disk between
these two parts but itself divided into two parts (Jc
and 1) by a thin membrane, which is distended like
a diaphragm right in the constriction. To this disk
run the extreme ramifications of the spinal nerve (n)

Fig. 231. A: Section of a compound luminous spot in Astronesthes niger, X 100. After Lendenfeld.

I, cornea-like, transparent part of skin over the organ; c, transparent membrane outside the lens-shaped body d; e, gland-tubes of the sphe-
rical part; /, pigment layers; g and g. pigment coating of the outer and inner parts of the luminous body; h, gland-tubes in the outer part
of the luminous body; Jc, granular mass and cells, inner part; l, outer part; m , shining membrane within the pigment layer; n, nervous cells.

Fig 231. B: Diagrammatic section of the luminous spot represented in fig. A, showing the reflection and refraction of the rays

of light in the luminous spot. After Lendenfeld.

ai a4 ar,d h &4, rays of light emitted from the central part (1c) of the luminous spot and reflected in the lower, spherical part of the

same; Cj c3, dy d3, — e3, and fy — f3 , rays of light from the same source, emitted in a centrifugal direction, reflected from the sides of

the outer, parabolic portion of the organ, and refracted by passing through the cornea-like layer (g); xxy, axis of the optical system;

Q , cornea; h, shining tissue of the outer, parabolic part; i, glands of the inner, spherical part.

See for example the upper spots in our figure of Argyropelecus Olfersii.

922

SCANDINAVIAN FISHES.

that innervates the organ; and the rays of light emitted
from the shining mass are reflected, as shown in fig.
231, B , collected in a cone and concentrated at their
exit through the dermal lens.

These luminous spots" regularly appear, among
the fishes now under consideration (see for example
our figure of Maurolicus Mulleri), in two rows, an
upper and a lower, along each side of the body. In
their more highly developed form, which consists mainly
in the addition of a more powerful reflecting appara-
tus* 6, it is the rule, however, that the upper row breaks
up into two or more groups of 3 — 6 spots gathered
in a row, arid more or less entirely coalescent inter-
nally (with the elsewhere spherical parts). Similarly

A b

Fig. 232. A: The confluent lower abdominal spots of Sternoptyx
diaphana, twice the natural size, and seen from below.
a, ventral fins. After Lendenfeld.

Fig. 232. B: Section of the luminous spots shown in fig. A.
a, basal canal; b, outer parabolic cups. After Lendenfeld.

the inner parts of the luminous spots in the lower
(ventral) row frequently coalesce; and this confluence
may extend to still more spots, even till the spots on
one side of the body coalesce with those on the other.
This is the case, for example, with the luminous spots
set at the very edge of the belly, in front of the vent-
ral fins, in Argyropelecus Olfersii. Here the inner
parts of 12 pairs of luminous spots coalesce, forming
a common, longitudinal, basal canal (cf. fig. 232, B, a),

while the outer (distal) parts of each spot (cf. fig. 232,
B , h) remain independent on each side of the sharp
median carina of the ventral margin (cf. fig. 232, A).
Besides the above-mentioned reflecting apparatus, which
lies within the pigment layer, and which here consists
of a comparatively thick layer of closely packed and
prismatic, calcareous spicules, Lendenfeld has described,
from the outer (cup-shaped) portions of these organs,
cylindrical and prismatic structures within the refract-
ing mass (analogous in situation to the above-men-
tioned gland-tubes). In these structures he distin-
guished two kinds of radiating cells, one of them
elongated and club-shaped, with an oval, highly re-
fracting body (probably a vesicle) in the thicker, distal
end; and these cells, the proximal peduncles of which
he supposes to be in direct connexion with the nervous
fibrils, are in his opinion “the special phosphorescent
elements”.

These more highly developed luminous spots appear
not only on the sides of the body in longitudinal rows,
but also on the halves of the lower jaw and on the
branchiostegal membrane. The largest of them are
isolated shining spots on the sides of the body, on the
dorsal or the ventral margin of the tail, or on the
head, especially below the eyes, on the snout, and on
the forehead. The large suborbital spots that occur in
certain species are innervated by a special branch of
the fifth pair of cranial nerves. That their function
is subordinated to the will of the fish, seems therefore
indubitable. Lendenfeld explains their purpose as
twofold: 1) the illumination of the water round the

fish, whether their possessor be seeking food in the
dark abysses of the ocean or at the surface in the
night-time — possibly too as a signal between the sexes
during the spawning-season — 2) the intimidation of
a pursuing enemy, when a sudden flash of light may
save the fugitive. The latter interpretation applies in
particular to the luminous spots situated on the hind
part of the body.

The parietal organ of Amphibians and reptiles,
according to our present knowledge of its structure,
seems also capable of explanation as a luminous spot.
The largest luminous spots known are found in a deep-
sea fish of the Scopeloid family, Ipnops Murrayi c, whose

° Composite, ocellar, phosphorescent organs, without reflector, Lendenfeld.

6 Composite, ocellar organs, with special reflector, Lendenfeld.

c Gthr, Deep Sea Fish., 1. c., p. 190, pi. XLIX, fig. B, and Moseley, ibid., App. A, pi. LXVII and LXVIII.

SCOPELOIDS.

923

eyes and optic nerves are entirely aborted, the fish
being consequently quite blind". But under the ex-
tremely thin and transparent roof of the skull (an-
swering to the parietal and frontal bones, which are
hardly distinguishable), and forward over the wide pa-
latine arch, thus extending above nearly the whole
surface of the flattened head to the posterior limit of
the nasal region, there lie a pair of lamelliform lumi-
nous spots, separated from one another by a longi-
tudinal median septum rising from the roof of the
mouth. Above each of these luminous spots run both
the frontorostral branch of the cephalic system of the
lateral line and the nasal branch of the fifth pair of
cranial nerves. The position of these luminous spots
is thus, like that of the parietal organ, intracranial.
With regard to their function Gunther remarks: “The
power of producing light, and thereby attracting other
creatures, must be of great use to a fish, which, de-
prived of organs of sight and touch, would be unable
to procure its food.”

As we have mentioned above (p. 826, note/), Gun-
ther'' has proposed to range the Scandinavian members
of this family in two families, the Stern op tychidce and
Scopelidce. But the difference between these two fa-
milies is hardly appreciable. They occupy an inter-
mediate position between the Salmonoid and Clnpeoid
families. In common with the former they possess an
adipose fin behind the true dorsal fin, and their ovaries?
like those of the latter, are furnished with a special
oviduct. The adipose fin is often very small, but in
most cases furnished with fibrillous (cartilaginous) rays.
The distinction adopted by Gunther consists merely
in the greater or less length of the intermaxillaries,
the posterior part of the margin of the upper jaw being
formed in his S ter nop tychidce by the maxillary bones.
But these bones often coalesce so closely with the in-
termaxillaries that the limits between them can hardly
be detected. The utility of the character is thus con-
siderably reduced. On the other hand it happens, in
Gunther’s Scopelidce , that the hind part of each inter-
maxillary bone is firmly applied to the outer surface
of the maxillary, above the lower margin of the latter.
Consequently the said margin may form part of the

margin of the upper jaw in this family as well, though
in such instances it is toothless. We therefore prefer
the older opinion, embraced in recent times by Moreau*'
and retain the Scopeloid family in its entirety. It
then contains about eighty species of marine fishes,
most of them deep-sea forms or nocturnal surface-fishes
but some belonging to the littoral regions of the tropic
seas. The Scopeloids belonging to the Scandinavian
fauna may be distinguished as follows:

1 : Snout shorter than the postorbital
part of the head. Dorsal fin situated
about half-way along the body.

A: Abdominal part of the body ex-
ceedingly deep and compressed ;
its ventral margin sharp and se-
parated by an abrupt break from
the under surface of the tail.

(Subfamily S ter nop ty chince) Argyropelecus Olfersii.

B: Abdominal part of the body pass-
ing uniformly (without sharp
break) into the caudal part, and
with more or less terete ventral
margin.

a: Maxillaries furnished with
teeth behind, and entering
into the structure of the mar-
gin of the upper jaw. (Sub-
family Cocciinui). Preabdomi-
nal length greater than the

postabdominal. _ Maurolicus Mulleri.

b: Maxillaries toothless, and as
a rule forming no part of the
margin of the upper jaw (Sub-
family Saurince). Preabdomi-
nal length less than the post-
abdominal.

a: Length of the base of the
dorsal fin more than ]/5 of
that of the body. Longi-
tudinal diameter of the eyes
less than 1/2 of the post-
orbital length of the head. Myctophurn elongatum.
(3: Length of the base of the
dorsal fin less than l/6 of
that of the body. Longi-
tudinal diameter oftheeyes
more than ‘/2 of the post-
orbital length of the head. Myctophurn glaciate.

2: Snout longer than the postorbital part
of the head. Dorsal fin situated far
back (Subfamily Paralepidince). No
luminous spots Sudis atlanticus.

“ Also without olfactory nerves, according to Moseley.
b Cat. Brit. Mus., Fish., vol. V, pp. 384 and 393.
c Hist. Nat. Poiss. Fr., tome III, p. 491.

924

SCANDINAVIAN FISHES.

Subfamily STERNOPTYCHINJE.

Body of a singular irregular form. , deep and compressed , especially in the abdominal part , ivhose sharp ventral
margin is abruptly cut off by a posteriorly ascending break ( postabdominal pari) from the under surface of the
tail. Cleft of the mouth sharply ascending. Snout shorter than the postorbital part of the head.

By its singular form of body, deep as that of the
Dory or the thinnest Carangoids, but with an abrupt-
break in the inferior protile between the abdominal
and caudal regions, this subfamily is so well distin-
guished from all other fishes that no detailed diagnosis
can be necessary. The resemblance in the form of the
body to Zeus (see above, p. 305, note a), and the si-
milar structure of the ventral margin in Trachichthys,
a Berycoid form (see Lowe, Fish. Madeir p. 64),
have suggested that these fishes should be included
among the Acanthopterygians. But merely their lu-

minous spots and the distribution thereof ought, at the
first glance, to assign them to their right place, beside
the other Scopeloids. Only four species, distributed
among three genera, are known with certainty. All
are nocturnal surface-fishes, which by day descend to
some depth below the surface, but at night ascend to
higher strata,. They belong to the basins both of the
Atlantic and the Pacific, strictly to the warmer regions
thereof; but on the Norwegian coast is occasionally
found a species of the

Genus ARGYROPELECUS h

Forepart of the back surmounted by an erect, thin and transparent, osseous structure, resembling a dorsal fin,
and composed of the neural spines of the anterior abdominal vertebrce and their chondrified interspaces.

Jaiv-teeth set in one row.

Of this genus, which is known exclusively from
the Atlantic and the Mediterranean, Valenciennes6
has indeed adopted four species; but two of them are

known only from his somewhat defective descrip-
tions, and the other two come extremely near to each
other.

a Cocco, Giorn. Scienc., Lett., Art. Sic., No. 77, Palermo 1829. The name signifies, according to Agassiz, with silver helmet (ccq-
yvqog, silver, and 7zrfrf„ helmet). To Cocco’s paper we have not been able to refer.
b Cuv., Val., Hist. Nat. Poiss., tom. XXII, pp. 392, scqq.

SC0PEL01DS.

925

ARGYROPELECUS OLFERSII.

Fig. 233.

Length of the maxillaries about equal to the horizontal length of the head Distance between the dorsal fin proper
and the tip of the snout about half the length of the body , but perceptibly less than the greatest depth of the
body , which is about s/5 °f Ls length , and only slightly less than (at least 9/10 of) the distance between the ventral
fins or the anal fin and the tip of the snout. Length of the base of the dorsal fin proper about equal to the
vertical diameter of the eyes1, and the least depth of the tail about 4/5 of the said diameter c. Silvery lustre of

the body extending over the sides of the tail.

Fig. 233. Argyropelecus Olfersii , natural size. Taken off Haagholm, near Bergen, on the 28th Jan., 1890.

Specimen in the possession of the Bergen Museum.

R. hr. 9; D. 9; A. 12; P. 10; V. 6; C. x + 1 + 1 8 d + 1 + x.

Syn. Sternoptyx Olfersii , Cuv., Reyn. Anim., ed. 2, tom. II, p. 316,
tab. XIII, fig. 2; v. Dub., Kor., Vet.-Akad. Handl. 1844,
p. 80, tab. 3, fig. 6; Lowe ( Pleurothyris ), Fish. Madeir.,
p. 64; ( Argyropelecus ), Proc. Zool. Soc. London 1850, p.
247; Cuv., Val., Hist. Nat. Poiss ., vol. XXII, p. 408;
Nilss. ( Sternoptyx ), Skand. Fna , Fisk., p. 486; Gthr
(. Argyropelecus ), Cat. Brit. Mus., Fish., vol. V, p. 386;
Coll., Forh. Vid. Selsk. Chrnia 1874, Tillasgsh., p. 149;
1879, No. 1, p. 84; N. Mag. Naturv. Chrnia, Bd. 29, p.
102; Storm, Vid. Selsk. Skr. Trondhj. 1883, p. 29; Gthr,
Deep Sea Fish., Chall. Exped., p. 167; Lillj., Sv., Norg.
Fi.'k., vol. III. p. 3.

Argyropelecus Olfersii attains, according to Collett,
a length of at least 82 mm.; but most of the speci-
mens known are of smaller size. Cuvier’s specimen
was 68 mm. long. The two specimens that we have
been enabled to examine — kindly lent us by the Mu-
seums of Christiania and Bergen — measure, from the
tip of the snout to the end of the middle caudal rays,
respectively 63 and 64 mm., being thus about equal in
size to the specimen described by v. Duben and Koren.

The species is one of the most singularly shaped
fishes, deep and compressed as a Dory, but, like its
congener, with a sharp, posteriorly ascending break in
the inferior profile between the abdominal region and
the tail. In the abdominal region the depth of the
body is about equal to the entire length of the trunk,
or about 58 — 62 % of the length of the body; whereas
at the vent the depth is only half of the greatest
depth, and about equal to the length of the tail. The
least depth, in front of the supporting (spinous) caudal
rays, is only about 7 10 of the length of the body.
The thickest part of the body is the head. According
to v. Duben and Ivoren, the thickness is greatest in
the inferior part of the body; but in our specimens
it is fairly uniform almost throughout the sides of the
head, measuring across the upper articulations of the
preopercula about V8 of the length of the body or
43 — 46 V2 % of the length of the head. Both the
dorsal and the ventral margins are compressed and,
in part, sharp, the latter, however, from different

a Somewhat greater or less; in Argyropelecus hemigymnus only about of the said length.
b In Argyropelecus hemigymnus about 2/3 thereof.

C 1 / •> '

5? y> ?? / 2 3 ”

d 17, according to v. Duben and Koren. In Argyropelecus hemigymnus we have counted 16.

926

SCANDINAVIAN FISHES.

causes. In front of the dorsal tin the dorsal margin
rises in a transparent, cartilagino-osseous carina, re-
sembling an anterior, posteriorly ascending dorsal tin,
but formed by the produced and coalescent tips of 7
neural spines. The ventral margin, on the other hand,
is sharpened at the isthmus, in the abdominal region,
in the region of the anal tin, and just in front of the
caudal tin, by the median carina between the rows of
luminous spots situated at these points, a structure
most nearly corresponding to the ventral carina of the
Herrings. But between the ventral tins and the anal
tin the ventral margin is channelled by a groove open
below. In the rest of their extent the margins of the
body are indeed thin, but blunt. The dorsal protile
follows a fairly regular curve in front of the true
dorsal tin, but from the beginning of this tin it slopes
almost in a straight line, until, at the caudal tin, it
becomes horizontal. The ventral profile is even more
sharply curved in front than the dorsal; but behind
the above-mentioned break it ascends at the same angle
as that formed by the dorsal profile in its descent,
though with a perceptible break at the end of the anal tin.

The head is deeper than long. Its singular shape
i due to the short snout, the vertically ascending cleft
of the mouth, and the deep, but short opercular appa-
ratus; and to these avc should add the comparatively
deep isthmian region, which extends under the greater
part of the length of the head. At the top the head
is convex from the occiput to the snout, but flat and
grooved in the middle, with a sharp, longitudinal ca-
rina on each side, and strongly constricted between
the eyes. Its length in an horizontal direction from
the middle of the tip of the snout measures in our
specimens about 28 — 29 % of the length of the body.
The eyes are rather large and round, the vertical dia-
meter being only a little greater than or equal to the
longitudinal, and varying in our specimens between
43 and 45 % of the length of the head, or about 1/8
of that of the body. They are set high, their upper
margin rising almost to the frontal plane, and so near
to each other that the least interorbital width is only
1 ,/4 of their vertical diameter. The postorbital length
is only about 1/3 (32 — 34 %), and the length of the
snout rather more than V4 (27 — 28 %), of the length

of the head; and the lower margin of the pupil lies
on a level with the tip of the snout or a little higher.
The suborbital ring is represented only by an extremely
thin, oblong, lamelliform, and vertically set preorbital
bone, which coasts the anterior margin of the eye.
The nasal cavities are oblong, and lie close to the an-
terior upper corners of the orbits and below the knob-
shaped tip of the ethmoid bone. The nostrils are di-
vided on each side by a thin, but flat septum of skin,
without any elevation of the margin.

The mouth ascends almost vertically, and the tip
of the upper jaw projects slightly beyond the ethmoid
(rostral) tip, but admits of hardly any protrusion. The
nasal processes of the intermaxillaries are short, and
the latter bones, each by means of its lateral branch,
are so firmly united to and coalescent with the maxil-
laries that the limit between them is distinct only in
front, for a distance measuring about 1ji of the dia-
meter of the eyes. The motion of the intermaxillaries
is thus reduced to a considerable extent: they can only
follow the forward and backward swing of the maxil-
laries. flow far the margin of the upper jaw is formed
by the intermaxillaries, it is impossible to determine
without dissection, an operation of which our materials
have not permitted. But the whole margin of the up-
per jaw is set with a row" of pointed, subulate and
curved teeth, rather scattered and small, but of various
sizes. The last four or five teeth'' are directed for-
wards, the anterior ones pointing backwards. The ma-
xillary bone, which is extremely thin, reminds us most,
both in shape and structure, of that of the Herrings.
Just behind (below) the constricted articular part it is
curved like a sabre at the very margin of the mouth,
but from this point back it is straight and of uniform
breadth, forming an oblong rectangle. The median
space, however, is not ossified, but consists of a silvery
skin, distended between the true maxillary bone and its
supplementary (jugal) part, which coalesces behind with
the maxillary, and which extends, in the form of an
osseous rod, forward to the articular part. The entire
length of the maxillary is somewhat more than ’/4
(about 26 — 29 %) of the length of the body, and its
breadth in the oblong part about ]/4 of its own length.
The lower jaw is about equal in length to the maxil-

“ Sometimes we find in front, on the intermaxillaries, a tooth or two lying inside or outside this row.

6 Situated on that part of the upper jaw which, when the mouth is wide open, lies outside the branch of the lower jaw, and thus
below the true cleft of the gape. In Argyropelecus hemigymnus the length of the intermaxillaries may be traced backward to this point,
which there answers to about the middle point in the length of the maxillaries.

ARGYROPELECUS OLFERSII.

927

laries. Each of its two branches resembles an acute-
angled, isosceles triangle, with the short (hind) base
turned downwards, where the angular part forms a
projecting, spine-like angle. Within (above) and pa-
rallel to the thin lower margin runs a ridge on the
outer side. The symphysis is straight, and projects
downwards in a chin-protuberance, channelled behind.
Most of the teeth in the lower jaw, which are also set
in a single row, are about twice as large as those in
the upper jaw; but one of the anterior teeth on each
side is considerably larger than the rest, and at the
extreme front, where the teeth are of about the same
size as in the upper jaw, they are more distinctly ar-
ranged in a double row. The tongue is rudimentary,
and the cartilaginous tip of the hyoid bone is smooth,
without teeth. Only the upper jaw is furnished with
a palatal fold, and this is not very broad. The vomer
is toothless, but on the anterior part of the palatine
bones, as well as on the upper pharyngeals, small teeth
may be felt. The pseudobranchiae on the inner surface
of the opercular and hyomandibular bones form an oblong
patch, in which about 14 filiform, transversely set la-
mellae may be counted. The gill-slits are large, ex-
tending far forward in the mouth; but the upper parts
of the hind branchial arches coalesce with the front side
of the scapular arch, the hindmost gill-slit thus being
closed above and rather short. The gill-rakers are long
but scattered, numbering about 14 on the first branchial
arch, besides a few small, hardly perceptible spines at
the lower (anterior) end. Where the horns of the hyoid
bone touch the tip of the basi-hyoid series (the lingual
bone), they are furnished with an erect protuberance.
The outer gill-openings are large, their upper angle
lying on a level with the centre of the eyes, and the
branchiostegal membranes being free from the isthmus
arid united to each other not quite to the full length of
the first branchiostegal rays, so that the anterior angle of
the openings lies in about a line with the tip of the snout.
The branchiostegal rays are slender and rod-like. The
first six lie rather near each other, and the first of all is
set close to the corresponding ray on the other side; the
last three are somewhat larger and set further apart.

The opercular apparatus, in accordance with the
shape of the head, is characterized by its height. The

vertical arm of the rectangular preoperculum is about
three times as long as the horizontal. The preoper-
cular angle is prolongated to a short, flat (thin), tri-
angular spine, pointing downwards. The other oper-
cular bones are extremely thin and flexible. The
operculum is rectangular, but forms an oblique arti-
culation above. Ifs surface is obliquely crossed by a
ridge running from the anterior upper corner down-
wards and backwards. Its breadth is equal to the length
of the snout. At its lower margin lies the triangular
suboperculum, with a more or less deep indentation
behind. Below the latter and below the preoper-
culum lies the interoperculum, which is divided into
two parts, the anterior (below the preoperculum),
elongated and united with the branchiostegal mem-
brane only behind, the posterior quadrangular with
rounded corners or circular, and entirely coalescent
with the branchiostegal membrane, on which it lies
like a scale.

The greater part of the head, as well as the rest
of the body, is covered with a thin, silvery epidermis;
but the upper temporal region, between the eyes and
the occiput, is naked, the surface of the bones being
finely grooved and granulated.

The fins resemble in quite essential respects those
of the Hemibranc.hs and Lophobranchs. As in the said
fishes, the soft rays, which are thin and transparent,
with the tips strongly compressed in the longitudinal
direction of the body, show but little developed rami-
fication and have only scattered joints". In the sup-
porting apparatus of the paired fins too, we find points
that remind us most of the Sticklebacks.

The true dorsal fin is obliquely rounded, the first
rays being the shortest, but gradually increasing in
length to the fifth or sixth, which is the longest in
the fin. It begins at a distance from the tip of the
snout measuring about half (50 — 51 %) of the length
of the body, and the length of its base is about equal
to the vertical diameter of the eye or a little greater,
about 12V2 — % of the length of the body. A little
in front of this fin ends the fin-like osseous ridge
formed by the projecting tips of 7* neural spines be-
longing to the abdominal vertebrae, together with the
thin and transparent, feebly ossified inferneural mem-

“ We are indeed ignorant of the manner of locomotion employed by Argyropelecus Olfersii ; but from this structure of the fin-rays
we may conclude that the fins perform their function by means of vibrations. This is the case, as we have seen above, not only in the
Sticklebacks and Pipefishes but also in the Dory, where the second dorsal and the anal fins are furnished with similar rays.

6 In Argyropelecus hemigymnus belonging to the 3rd— 9th vertebrae.

Scandinavian Fishes.

117

928

SCANDINAVIAN FISHES.

brane by which they are connected. The adipose tin
behind the dorsal tin is long and low.

The anal tin, which begins just behind the vent,
is longer but lower than the dorsal fin proper, and the
hind part of its margin is deeply concave. The distance
between it and the tip of the snout is about 61 — 63 %
of the length of the body, and its base measures about
17 — 18 % of the same. In the distribution of the rays
we observe that the first seven and the last four are
set more or less close together at the base, but that
the eighth ray is more widely separated both from
the seventh and the ninth, and points straight down-
wards, its direction thus converging with that of the
seventh ray.

The caudal fin is deeply forked. The length of
the middle rays is 8 % of that of the body, and ap-
pears to be perceptibly less than half (at most about
46 %) of that of the longest rays. At the upper mar-
gin of the fin lie 9, at the lower 6, short but com-
paratively thick and hard supporting rays, the anterior
directed outwards (respectively upwards or downwards).

In the suspensory apparatus of the pectoral fins, the
uppermost bone, probably answering to the supra-cla-
vicular bone — in which case the posttemporal bone is
wanting — is bent at a right angle. Its upper, shorter,
horizontal arm is straight and of uniform breadth,
attached to the median line of the occiput — not at
the side, where the posttemporal bone is attached in
other fishes — and its lower, vertical arm, which is
somewhat broader and forms a thin expansion in front,
rests upon the upper part of the clavicle. The angle
between the arms forms a short spine, directed back-
wards, and both arms are coursed on the outside by
a longitudinal groove, bounded by ridges and hollowed
into grooves similar to those in the upper part of the
temporal region (see above). The ridges and grooves
are continued on the outside of the vertical part of the
clavicle in front of the pectoral fins, and below the
latter the clavicle sends out, in the skin, a backward
process, flat and of uniform breadth, but curved up-
wards like a sabre, and with similar grooves on the
outer surface. The inferior part of the clavicle ex-
pands forwards and inwards to a thin, vertical blade,
projecting straight downwards in a pointed spine, which
marks the limit between the sloping isthmian region

and the horizontal ventral margin. The pectoral fins
are obliquely set; when at rest, they point upwards
and backwards. They are rather long, but narrow and
obliquely pointed. The third or the fourth ray is the
longest, but only slightly longer than the second. The
first ray, which is the only simple one, is a little
shorter than the second; the last (10th) occupies about
2/lt of the length of the fin, which measures somewhat
more than 1/4 (26 — 27 %) of that of the body.

The ventral fins are set undermost on the ascend-
ing break in the ventral margin, with their insertions
close together and vertically situated, the first ray,
which is simple but, like the others, compressed, being
thus the lowest, and the under side of these fins in
other fishes being here turned inwards". In shape the
ventral fins are oval, and their length is about equal
to the height of the anal fin, 8 1/2 — 9V2 % of the length
of the body. The lowest point in their insertions
lies at a distance from the tip of the snout measuring
nearly 2/3 (62 1/2 — 65 %) of the length of the body,
and the distance between this point and the foremost
point in the insertions of the pectoral fins (the pre-
abdominal length) is about 29 — 30 % of the same
length. The postabdominal length, which ascends but
breaks off at an angle, measures in a straight line
about 14V2 — 15 % of the length of the body. The
thin pelvic bones, which are vertically set, share in
the structure of the septum between the luminous spots
of the preabdominal margin. But at the lower poste-
rior corner each of the pelvic bones projects in a spine,
and sends out above this spine an ascending process*
which joins the lower part of the hindmost preabdo-
minal rib, while the lower tips of the preceding ribs
touch the squamous growths that enter into the struc-
ture of the sharp ventral margin and bear its lumi-
nous spots.

The greater part of the body — except the black
or grayish brown back — is covered with a thin, sil-
very skin; but according to v. Duben and Koren
scales are also present, though these are thin and deci-
duous. “These scales are rather large (about 2x/2 mm.
long in a specimen 65 mm. in length), smooth with
entire margin, and of a bright silvery lustre like their
underlayer. Under the microscope they gleam with all
the colours of the rainbow, but not even then does

a This is also the case, as we have seen above (p. 426), in Botkina, but is there due to an entirely different structural peculiarity.
b Cf. the analogous structure in the Sticklebacks, see above, p. 635, fig. 157, A, vs.

ARGYROPELECUS OLFERSII.

929

their surface show any regular streaks or the like.”
Of the lateral line v. Duben and Koren saw only a
trace, “in the form of an elevated line beginning be-
low the origin of the true dorsal tin, and running
along the middle of the body to the base of the
caudal tin.”

The luminous spots are distributed as follows.
On the under side of the lower jaw — on the soft
chin" — below the tip of the lingual bone, lie two ra-
ther undefined spots, which are probably luminous6,
though in this species, to the best of our knowledge,
they have not yet been examined. On the ceratohyoid
bones, in the skin between the bases of the first (low-
est) 7 branchiostegal rays, there are 6 small luminous
spots on each side of the body. Behind these each
side of the isthmus is furnished with 6 larger luminous
spots, gradually increasing in size as the isthmus grows
deeper behind. These are succeeded, on each side of
the sharp preabdominal margin of the belly, by a row
containing 12 luminous spots of fairly uniform size,
somewhat smaller than those on the isthmus and small-
est at the ' beginning and end of the row. Above the
rows on the isthmus and the preabdominal margin each
side of the body bears a row of 10 luminous spots;
but this row is interrupted and belongs partly to the
head, partly to the abdominal region. One spot lies
within the anterior (horizontal) arm of the preoper-
culum, but spreads downwards over the anterior part
of the interoperculum. Another, smaller spot occupies
the lower part of the suboperculum and the hind part
of the interoperculum. Two large spots adjoin to the
hind margin of the clavicle above the insertion of the
pectoral fin, and are indeed vertical in direction, but
set so that their tops form a continuation of the curve
followed by the upper parts of the spots on the isth-
mus. The remaining six abdominal spots are about
equal in breadth to the last-mentioned pair but shorter
than they, and lie in a continuous, horizontal row, but
lower than the last-mentioned pair. Each of them
occupies the space between two ribs, and lies vertically
or obliquely above one of the spots in the lower pre-
abdominal row, the penultimate spot, however, being
situated above the penultimate two in the lower row,
both of which occupy the same intercostal space. The
segmental arrangement of the luminous spots is thus

not observed with complete uniformity. Each side of
the postabdominal margin bears a continuous row of
4 luminous spots. Owing to the singular form of the
body these spots are indeed set at the ventral margin,
i. e. their lower parts extend thither, but also on a
level with the two large spots above the insertion of
the pectoral fin. In size they answer most nearly to
the spots in the upper preabdominal row, which they
also resemble in the circumstance that the spots on
one side of the body do not coalesce internally (supe-
riorly) with those on the other side, as is the case
with the lower preabdominal row. The postabdominal
spots on one side of the body are also separated in-
teriorly from those on the other: their defining walls
do not coalesce below, like those of the preabdominal
spots, into a median edge. On each side of the body
the walls coalesce into a more or less high, thin rim,
and together form in this manner a groove or deep
channel, open below, at the hind extremity of which the
vent is situated. The luminous spots on the tail form
two groups, the one, along the posterior two-thirds of
the base of the anal fin, being a row of 6 rather large
spots, the 2nd, 3rd, and 4th smaller than the rest, the
other a row of 4 smaller spots at the inferior margin
of the tail, just in front of the caudal fin. All these
luminous spots thus belong to the lower part of the
body. Higher up we find, however, an isolated lumi-
nous spot on each side of the head, on a level with
the tip of the snout. This spot is situated on the
uppermost part of the preoperculum, just below its
articulation.

The above description of the luminous spots in
Argyropelecus Olfersii applies in every detail to the
analogous organs in Argyropelecus hemigymnus , a Me-
diterranean and Central Atlantic form which in other
respects too so closely I’esembles A. Olfersii that it
evidently represents a lower stage in the same course
of development.

Argyropelecus Olfersii has been named after a
Prussian diplomatist who, while on a voyage to Brazil,
secured an example of this species that had become
entangled among weeds to a lead-line, “between the
Canary Islands and Brazil.” The specimen figured by
Cuvier (1. c.) was found by Dussumier some kilom.
south-east of the Cape of Good Hope, among a floating

a Mention, pars inter gnathidia mollis (Sundevall).

b Cf. Argyropelecus hemigymnus and Sternoptyx diaphana, Lendenfeld in Gunther, Deep Sea Fish., 1. c., p. 314.

930

SCANDINAVIAN FISHES.

mass of “zoophytes” (Hydrozoa on seaweed?). It lay
on one side at the surface, but was still alive. Its
colour was described as nacreous on the sides of the
body, reflecting various hues, principally azure blue.
The back was black, with the same play of colours;
the fins were transparent; the eye was of a handsome
green.

The species is thus known originally as a surface
fish. By the Challenger Expedition, however, it was
taken in a dredge that had been drawn at the bottom
in 1,125 fathoms of water, off Cape Finisterre. In
most instances, as in the case of deep-sea fishes, the
specimens secured have drifted helplessly at the sur-
face and been cast ashore by the waves. The congener
of this species — if they are both entitled to an inde-
pendent, specific rank — was taken in 1869 by the Por-
cupine expedition, in 540 fathoms of water, between
Scotland and the Faroe Islands", and in 1880 and
1882 by the American expeditions on the Blake and
Fish- Hawk, in 225 and 245 fathoms of water, off the
coast of Florida6. Argyropelecus Olfersii is also fre-
quently found off the Norwegian coast in the stomach
of Cod, a circumstance which seems to indicate that
it keeps near the bottom, as is further suggested by
its large eyes. But at night its congener is often met
with at the surface off the coast of Sicily; and it is
highly probable that Argyropelecus Olfersii shares this
habit. To the true deep-sea fishes it cannot thus be
referred; and the above-mentioned instances of its oc-
currence at great depths do not exclude the possibility

of its having been intercepted by the dredge while the
latter was being drawn up. It seems most likely that
in the daytime it avoids the light by retiring to depths
where the sun exercises no appreciable influence.

Its geographical range extends from the Cape of
Good Hope to North Cape, but appears to be restricted
between the limits of the great ocean currents. It has
thus been carried by the Gulf Stream on several re-
corded occasions to the Norwegian coast north of Ber-
gen, but has never been found further south in Scan-
dinavia. Collett mentions 8 specimens as having been
found during the last thirty years, cast ashore at dif-
ferent points on the Norwegian coast, or taken in the
stomachs of Cod or Coaltish.

Argyropelecus Olfersii is a voracious fish-of-prey,
as evidenced by its dentition. In the stomach of a spe-
cimen 82 mm. long Collett found a half-digested
Maurolicus Mulleri that had probably measured 50 mm.
Small fishes and crustaceans are, no doubt, its principal
diet. Its spawning-season is unknown; but in a fe-
male about 8 cm. long Collett counted about, 1,000
eggs x/2 mm. in diameter, a much greater number and
relatively smaller size than we found in a female
Argyropelecus hemigymnus, taken in February off the
Sicilian coast. In the latter specimen both ovaries
were rather tumid but of different lengths, the right,
which was the longer, measuring hardly 6 mm. The
eggs were V2 mm. in diameter, and lay so loose in
the ovaries that they were probably almost ready for
deposition.

Subfamily COCCIINJE.

Body of a slightly irregular Herring-form. Ventral margin more or less terete. Cleft of the mouth more or less
ascending , with the margin of the upper jaw formed in front by the intermaxillaries and behind by the maxil-
laries, which are here armed, with teeth. Teeth in the mouth of fairly uniform size. Preabdominal length greater
than the postabdominal . Pseudobranchice well developed. Snout shorter than the postorbital part of the head.

The known forms of this subfamily are not nume-
rous. Gunther c enumerates 5 species, distributed among
two genera, one of which, the Mediterranean Coccia,

containing one species with the most reduced inter-
maxillaries and jaw-teeth, has conferred its name on
the subfamily. In form and coloration, and probably

° Day, Fish. Gt. Brit., Irel., vol. II, p. 48.

b Brown-Goode and Bean, Bull. Mus. Comp. Zool., vol. X (1883), p. 250.

c Cat. Brit. Mus., Fish., vol. V, p. 387. Whether Diplophus (Gunther, Mus. Godeffr., Heft,. II, p. 101) belongs to this subfamily
is uncertain, so long as we are ignorant whether it possesses or is without pseudobranchiae. The subfamily Chauliodontince, to which, accord-
ing to Gunther, it should else be referred, is destitute of pseudobranchiae, and in general has a larger mouth with a more powerful dentition.

PEARL-SIDES.

931

in their habits, these fishes closely resemble the small
Herrings in whose company they are frequently found;
and they sometimes appear, like the latter, in shoals.
All of them are small fishes. Most of them belong to
the Mediterranean, only one species being known from
the Atlantic; and this is so like one of the Mediter-
ranean forms that its right to the rank of a separate
species seems questionable. This same species has also
been claimed, probably with justice, for the fauna of
New Zealand. The subfamily has thus an extensive
geographical range, and probably occurs in other in-
termediate regions both of the Atlantic and the Pacific.

Another species, Maurolicus tripunctulatus, described
by Esmark from the neighbourhood of Madagascar, is
stated by Lutken" to occur in “Denmark Sound” be-
tween Iceland and Greenland (66° N. lat. ; 28° W. long.).
The same peculiarity, the occurrence of a species in
localities so remote as, on the one hand, the North
Atlantic and the Mediterranean, on the other, New Zea-
land, without its being known from intermediate regions,
we have already observed in the case of Argentina spliy-
rcena, which also associates with the Herrings. And the
Herring family, as we shall soon see, contains some of
the most widely diffused piscine species.

Genus MAUROLICUS* 6.

Snout and forehead forming an unbroken profile, slightly sloping. Tip of the lower jaw prominent. Breadth of
the maxillaries less than half their length and less than the breadth of the operculum ( measured in the longitudinal
direction of the body). Jaw-teeth distinct, though small. Pseud obranchice present. Air-bladder wanting c.

This genus is distinguished, at the first glance,
from Coccia by the form of the snout, which in the
latter genus comes nearest that of tschir among the
Gwyniads, being short and blunt, tumid and truncate.
With this is connected the form of the maxillaries.
Coccia, whose gape is thus rendered much smaller, is
also without true jaw-teeth, these being merely fine
serrations of the sharp and thin edges of the jaws. In
Coccia, on the other hand, the luminous spots are more

numerous: the upper preabdominal row is continued,
unbroken and dense, on the postabdominal region, the
row thus containing 25 spots; and on the branchiostegal
membrane, which behind coalesces so completely with
the operculum as to be almost indistinguishable there-
from, Coccia ovata possesses 12 luminous spots. Coccia,
with its large eyes, which almost touch at the forehead,
where the interorbital width is extremely small, also
seems to be a still more marked nocturnal fish.

BOREAL PEARL-SIDE.

MAUROLICUS MULLER],

Plate XLIV, tig. 3.

Upper row of ventral luminous spots interrupted at the ventral fins and containing 0 or 10 spots between these
fins and the pectorals. Length of the lower jaw less than 15 % of that of the body, or than 16 % of that of the
body minus the caudal fin , and the postorbital length of the head more than 3/5 of the length of the lower jaw.

R. br. 10 U D.

2 1

A.e — ; P.

22 (15) 16-

V.

-18' 6’

C. x -I- 1 + 17 + 1 + x\ Lin. lat. 26 — 28; L. tr. 5; Vert. 32L

Syn. >Salrno inaxillis edentatis, inferiore longiore; ventre punctato,
Mull., Zool. Dan. Prodr., p. 49.

a Vid. Meddel. Naturli. For. Kblivn 1891, p. 211.

6 Cocco, N. Ann. Sc. Nat. (Bologna), Ann. I, tom. II (1838), p. 192 (p. 32, sep.) — “segnalato col nome di un celebre letterato
siciliano” (Bonap.).

c According to Gunther.

d 9, according to Nilsson, Prodr.; 8, according to Observ. Ichthyol.; 8 or 9, according to Gunther. 9 or 10(?), according to Kroyer.
e A. 26 — 33, according to Day.

J According to Day.

932

SCANDINAVIAN FISHES.

Sheppy Argentine , Penn., Brit. Zool. (ed. Warr. 1776), vol.
Ill, p. 286, tab. LXV, No. 156.

Salmo Miilleri, Gmel., Syst. Nat. Lin., ed. XIII, tom. I, p.
1378; Kr. ( Maurolicus ), Danm. Fisk., vol. Ill, p. 113.

Argentina Pennanti , Walb., Ichthyol. Art., pt. Ill {Gen. Pise.),
p. 47 ; Cuv., Val. {Scopelns), Hist. Nat. Poiss., vol. XXII,
p. 436; Yarr. (Richards.), Brit. Fish., ed. 3, vol. I, p.
330; Day {Maurolicus), Fish. Gt. Brit., Irel ., vol. II, p. 49,
tab. CIX, fig. 2; Colt,., N. Mag. Naturv. Clirnia, P>d. 29, p.
104; Petersen, Vid. Meddel. Natnrh. For. Kbhvn 1884 (1886),
p. 158; Lillj., Sv., Norg. Fna, Fisk., vol. Ill, p. 10.

Scopelns borealis, Nilss., Prodr. Ichthyol. Scand., p. 20; Id.,
Observ. Ichthyol., p. 9; Id., Skand. Fna, Fisk., p. 479;
Gthr ( Maurolicus ), Cat. Brit. Mus., Fish., vol. V, p. 389;
Coll., Forli. Vid. Selsk. Chrnia 1874, Tillsegsh., p. 150;
1879, No. 1, p. 84; Malm, Gbgs, Boh. Fna, p. 533; Winth.,
Naturh. Tidskr. Kbhvn, ser. 3, vol. XII, p. 42; Storm, Vid.
Selsk. Skr. Trondlij. 1883, p. 30; Jord., Gilb., Bull. U. S.
Nat. Mus., No. 16, p. 284.

Scopelus Humboldtii , Yarr., Brit. Fish., ed. 1, vol. II, p. 94;
ed. 2, vol. II, p. 161 (nec Risso, nec Cuvier); Storer,
Mem. Amer. Acad. Arts. Sc., n. ser., vol. II, p. 450; vol.

VI, p. 328, tab. XXV, fig. 5.

Forma mediterranea (an species distincta?), maxillis et mandibula
longioribus:

Maurolicus amethystino-punctatus , Cocco, 1. c.; Gthr, 1. c.,
p. 390.

Maurolicus australis , Hector, Trans., Proc. N. Zeal. Inst., vol.

VII, p. 250, tab. XI, fig. 90, D, vide Gunther, Ann. Mag.
Nat. Hist., ser. 4, vol. XVII (1876), p. 399.

Obs. In 1766 Strom found on the shore, at Wolden parson-
age in Sondmor, a specimen of this species (see his MS, quoted by
Collett), a drawing and description of which lie sent ‘'more than
20 years before 1791” (see Skr. Naturh. Selsk., Bd. 2, H. 2, p. 15)
to O. F. Muller, who appends to his above-mentioned diagnosis,
“Cl. StrOm misit.” Afterwards Strom confused this species with

Myctophum glaciale, which he distinctly figured and described in
Skr. Naturh. Selsk. (1. c.) — a circumstance first pointed out by
Nilsson in his Skand. Fna. But this cannot affect the determination
of Salmo Miilleri in Gmelin, who merely copied Muller.

The Pearl-side belongs to the small fishes. Our
largest specimen from Bohuslan indeed measures 72
mm. from the tip of the lower jaw to the extreme end
of the caudal fin; but the distance from the tip of the
snout to the end of the middle caudal rays, the mea-
surement termed in the present work the length of the
body, is not quite 68 mm.

The body is moderately elongated, with terete
back and belly, flattened, vertical and parallel sides.
In the forepart of the body, in front of the dorsal and
ventral fins, the ventral contour is perceptibly more
curved than the dorsal; but in the hind part of the
body these contours are straighter and converge with

fair uniformity towards the base of the caudal fin.
The greatest, depth of the body, which occurs at about
the middle of the preabdominal part, is about 1/5 (18
— 2l72 %) of the length of the same"; and the least
depth of the tail is about l/3 of the said depth or 1/.15
— Vie (about 6 V2 %) °f the length of the body*. The
greatest thickness, which is fairly uniform throughout
the forepart of the trunk (the preabdominal region),
measures about 8 — 10 % of the length of the body.

The outlines of the head run in uninterrupted
continuation of those of the body, its sides being also
flat and parallel; but the lower jaw, which sharply
ascends when the mouth is closed, renders the inferior
contour of the head more curved, while the superior
contour is straighter. The forehead is also grooved
longitudinally, its edges rising in a ridge that runs
along each side, and extends forward above the margin
of the eye, at the anterior upper angle of which it
divides to enclose the nasal cavity; but the middle of
the forehead is coursed by a lower longitudinal ridge.
The length of the head is about 1/4 (26 — 221/s %) of
that of the body. The eyes are round, though their
longitudinal diameter is somewhat greater than the
vertical, and measures about 38 — 35 %c of the length
of the head. They are set high, their upper margin
lying almost in the plane of the forehead, and are only
very slightly and obliquely upturned. The least inter-
orbital width is about 2/5 of their longitudinal diameter,
which is equal to the postorbital length of the head.
The nostrils are set close together, each pair at the
extreme front of the triangular nasal cavity, above the
dark top of the preorbital bone and behind the arti-
cular knob — also dark-coloured — of the maxillary.
The anterior nostril on each side is round, the posterior
an obliquely-set., transverse slit. The suborbital ring,
which forms the silvery cheek, is thin but complete
alon®- the lower half of the orbit, in front of uniform
breadth, only slightly broader at the penultimate sub-
orbital bone, the hindmost suborbital bone, on the
other hand, being small and triangular. The tip of
the snout is sharp (shallow) but truncate, with a slight
indentation at the middle. The length of the slender
intermaxillaries is equal to the vertical diameter of the
eyes. The maxillaries are formed as in the preceding

" 20 — 23 % of the length of the body excluding the caudal fin.

b About 7 % (according to Lilljeborg nearer 8 /) of the length of the body excluding the caudal fin.
c According to Liljeborg 32 % of the length of the head from the tip of the lower jaw.

BOREAL PEARL-SIDE.

933

species, but are curved in a sabre-like fashion through-
out their length, and their supplementary bone, which
is elliptical behind, projects in front in a tip that ex-
tends to a line with the anterior part of the preorbital
bone. Their median breadth is rather more than 1/3
of their length, which is equal to that of the lower
jaw, and less than 15 % of that of the body or than
16 % of the length to the middle of the base of the
caudal fin. In this last relation we have a character,
constant to the best of our knowledge, to distinguish
this form from the Mediterranean Maurolicus amethys-
tino-punctatus, which is otherwise deceptively like M.
Miilleri. The branches of the lower jaw are deep, but
have an arcuate upper margin; and the symphysis is
shallower than in Argyropelecus Olfersii and has a
feebly developed chin-protuberance, which however pro-
jects distinctly, when the mouth is closed, beyond the
tip of the snout". As in Argyropelecus Olfersii, a
sharp ridge runs along the outside of each half of the
lower jaw, parallel to the thin inferior margin. The
branches of the lower jaw approach rather near to
each other at the inferior margin behind; but in front
(behind the symphysis) the space between them widens
into a clavate mentum (chin-space). The feeble den-
tition of the mouth consists of small, pointed teeth,
set in a single row and of uniform size, on the inter-
maxillaries, on the hind (free) part of the lower edge
of the maxillaries, on the upper edge of the lower jaw,
as far as it is free when the mouth is open, and in
an arcuate transverse row on the head of the vomer.
There is no tongue; but the anterior (lower) part of
the basihyoid bones forms a transversely-set rim, rising
within the mouth like a tongue. The upper palatal
fold is extremely narrow, the lower better developed.
The pseudobranchiae are rather large, but consist of
only 9 or 10 transversely-set, filamentous lamellae.
The gill-rakers are set in one row, pointed, rather
long, and so dense that they number about 30 on the
front of the first branchial arch. The first gill- slit is
long, the others rapidly decrease in length. The outer
gill-openings are large in this species too, the two
branchiostegal membranes being hardly at all coalescent.
Above they extend to a level with the upper margin
of the eye, below to a line with the anterior extremity

of the preorbital bones. The branchiostegal rays are
10, the first extremely small, the others slender, ex-
cept the last (uppermost), which is broad and sabre-
shaped. Among the thin bones of the opercular appa-
ratus both the operculum and suboperculum are qua-
drilateral, rectangular, the former being rather more
than twice as large (deep) as the latter, but with the
upper posterior angle rounded. In the lower margin
of the suboperculum there is a rounded indentation.
The interoperculum is more oblong, growing narrower
in front, and lies along the lower arm of the pre-
operculum. The preoperculum resembles a somewhat
open (obtuse-angled) carpenter’s square, with both arms
of nearly equal length.

The dorsal fin is trapezoidal, with the upper pos-
terior margin rapidly sloping. It begins at a distance
from the tip of the snout measuring about 55 — 48 %
of the length of the body. The length of its base is
about 9 — 71/2 %, its height in front about 1 0 1/2 — 12 %,
of the length of the body. The adipose fin, situated
above the posterior part of the anal fin, is long and
low, as in Argyropelecus Olfersii. The anal fin is much
longer than the dorsal, but so deeply incised at the
margin that its anterior part is almost similar to the
latter fin, though much lower, while its posterior part
is very low and of uniform height, only the last rays
being slightly prolongated. Its distance from the tip
of the snout is about 63 — 60 %, its base about 1 8 1/a
— 20 %, and its greatest height about 1/9, of the length
of the body. The caudal fin is forked, though not so
deeply as in Argyropelecus Olfersii', its middle rays
occupy about 7 — 8 % of the length of the body6. At
the upper margin we have found 8 supporting rays, at
the lower 5. The pectoral fins are bluntly pointed,
semioval. Their length is about 16 — 14 % of that of
the body. The ventral fins are of almost the same
shape, but much shorter, their length being about 9 —
10 % of that of the body, and only slightly greater
than or even equal to the diameter of the eye. They
are set at a distance from the tip of the snout measur-
ing about 52 V2 — 50 V2 % of the length of the body,
sometimes a little in front of, sometimes behind the
perpendicular from the beginning of the dorsal fin.
The preabdominal length is about 30 (31a/2 — 29 V2) %

a Both the maxillaries and the lower jaw are so thin arid transparent that the whole of the former and the upper posterior part of
the latter are invisible in v. Wright’s figure.

h In v. Wright’s figure, which evidently represents a fresh and perfect specimen, the caudal fin is much less forked than in our
specimens.

934

SCANDINAVIAN FISHES.

the postabdominal about 10 (11 — 9) %, of the length
of the body.

The whole body, forward to the interorbital space,
is covered with extremely thin, transparent, rather
large, and rounded scales, somewhat higher than long,
and with scattered, concentric strife on the posterior
part. There is no special lateral line, the scales of
the median line being without pores.

The luminous spots are distributed in the same
manner as in Argyropelecus Olfersii, but more regu-
larly. A comparison with Coccia ovata shows most
clearly of all that they lie in three rows, though the
middle row should probably be regarded as a ramifi-
cation of the lowest, or vice versa. The uppermost
row begins at the extreme front of the chin, and in
Coccia runs back without a break along the branches
of the lower jaw and on the branchiostegal membrane,
one spot being situated between each pair of branchio-
stegal rays, till the membrane coalesces behind with
the interoperculum and suboperculum so completely
that the luminous spots apparently lie on the inner
surface of these bones. Above the row, on the inside
of the lower preopercular angle, we find one spot,
suggesting an upward ramification of this row. In the
Pearl-side the row is interrupted along the branches
of the lower jaw, only two spots, probably answering
to the second pair in Coccia , appearing on the soft
chin-space, and further back each of the branchiostegal
membranes is furnished with 6 spots, which shine through
the maxillaries and the horizontal (anterior) arm of the
preoperculum. On the inner surface of the opercular
apparatus the remaining spots are the same as in
Coccia ". The lowest row of luminous spots starts from
the anterior extremity of the isthmus, in Coccia quite
at the mental angle, where the two lateral halves of
the first mentioned row meet, and follows the ventral
margin, in this genus without interruption, merely
forming a curve at the beginning of the anal fin, back
to the lower corner of the base of the caudal fin. But
in front of the base of each pectoral fin and above this
row, lies one spot, suggesting an upward ramification
of the row, and on a level with the last-mentioned
spot runs the upper ventral row, pair by pair with the
spots of the lower row, and unbroken in Coccia to a
line with the beginning of the anal fin. In the Pearl-

side the lowest row begins similarly at the anterior
extremity of the isthmus; but this extremity is situated
further back, hardly in front of the perpendicular from
the anterior margin of the eye, and the row runs on
each side of the body, in an upward curve containing
6 luminous spots, towards the base of the pectoral fin.
At the ventral margin between the last pair of these
spots, begins the true (lower) ventral row, which should
thus be regarded here as a downward and backward
ramification of the isthmian row, and runs on each
side of the body to the lower corner of the base of
the caudal fin. This row contains 43 spots, the pair
at the base of each ventral fin (the 13th and 14th in
the row) somewhat obliquely arranged, and the remain-
ing 4 on the postabdominal region forming a slight
upward curve. A slight break in the row occurs at
the beginning of the anal firi (between the 18th and
19th spots), and a similar break at the end of the same
fin (between the 34th and 35th spots). The upper
ventral row of luminous spots, which in the Pearl-side
— to judge, at least, by its direction and the form of
the spots — is a more immediate continuation of the
isthmian row, begins in the upper part of the axilla,
and runs, containing 9 spots, to a line with the base
of the ventral fin, from which point it is interrupted
throughout the postabdominal region, but re-appears
in a single spot above the beginning of the anal fin.
The spots in the upper ventral row, those between the
branchiostegal rays, and most, if not all, of the pos-
terior spots in the isthmian row, are obliquely cut and
prolongated in a downward direction (see above, p.
921, with note a).

The coloration of the Pearl-side is almost the same
as that of the smaller Herrings, a lustrous silvery white,
but dark, shading into blue on a greenish ground, on
the back and the top of the head. So long as the
scales adhere, they give the sides of the body a tinge
of yellow. The black rims of the luminous spots stand
out sharply from the pale bluish (in spirit-preserved
specimens yellowish) lustre of their interior; and where
these rims touch, along the base of the anal fin and
the inferior margin of the tail, there appears a coal-
black band. The end of the tail itself is of the same
colour as the back, and outside (behind) the tail a
narrow, black band crosses the base of the caudal fin.

“ The luminous spot which Moreau {Hist. Nat. Poiss. Fr ., tome 3, p. 511) observed in M aur aliens ameihystino-punctatus on each
side of the snout, near the nostrils, we have not been able to detect in our preserved specimens of the Pearl-side; but it is distinctly shown
in v. Wrights figure, small, but prolongated along the lower margin of the anterior part of the preorbital bone.

BOREAL PEARL-SIDE.

935

All the tins are light and transparent, with only the
anterior margin or, in the caudal tin, the two outer
margins, of the same colour as the back, or dark.

Of the internal organs we shall merely remark,
after Day, that the stomach is ciecal, the duodenum
furnished with 8 rather large pyloric appendages, and
the whole intestinal canal forms only two convolutions.
The testes and ovaries are paired and long.

The Pearl-side is a gregarious fish, and though it
is most often met with in solitary specimens among
small Herrings (Sprats and Herring-fry), it also appears
in shoals of its own, instances of which have been ob-
served on the coasts both of Finmark and England.
Like Argyropelecus Olfersii, it is a pelagic and noc-
turnal fish of extensive geographical distribution. It
has been found on both sides of the North Atlantic,
and differs but slightly, if indeed it be specifically
distinct, from Maurolicus amethystino-punctatus, a Me-
diterranean species with which Gunther has combined
Maurolicus australis, a New Zealand form described
by Hector.

Within the limits of the Scandinavian fauna the
Pearl-side was first discovered by Strom off Sondmor,
and on the Swedish coast it was first taken by Nilsson,
near Gothenburg, in a seine drawn for small Herrings.
It has since been found at many points, both in Bo-
huslan and Norway, cast ashore by the waves, or in the
stomach of God and Herring". South of Bohuslan it is
unknown on the Swedish coast, and only one find has
been recorded on the west coast of Denmark, consisting
of a specimen lying on the shore at Hjorring in the

north of Jutland. On the Norwegian coast it is much
commoner, north to Hammerfest, and ascends the fjords.
In the Eyng, one of the Finmark fjords, east of Tromso,
there appeared in 1866, according to Collett, a whole
shoal of Pearl-sides. They could be scooped up in
bucketfuls from the surface, and at least 50 specimens
were preserved. Similar shoals have been seen on the
coasts of Great Britain and Ireland, in the North Sea,
the Channel, and the Irish Sea. According to Edward* * 6
the Pearl-side is a regular winter visitor off Banff,
never failing to appear in the month of January; and
from January to March Mr. Sim gathered in two years
nearly 200 specimens on the shore near Aberdeen. It
would thus seem to be most common to the north; on
the English coast south of Yorkshire only solitary spe-
cimens have been found, and on the west coast of
France, as well as in Holland, the species is unknown.
On the east coast of North America it is stated to
occur but seldom.

The small teeth and fine gill-rakers of the Pearl-
side show that it lives on minute animals. These con-
sist principally, we may assume, of the small crusta-
ceans (Entomostraca) that are so abundant in high
northern latitudes', and which compose the chief food
of the Herring too, though the latter, as we have seen,
also preys, like Argyropelecus Olfersii, on the Pearl-side.

During winter, when the Pearl-side appears in such
numbers off the Scotch coast, Day examined its gene-
rative organs, and found the comparatively large eggs
and the milt almost ripe in February. We may hence
conclude that the spawning is performed in spring.

Subfamily S A ( i! I N ffE.

Body of only slightly irregular Herring- form, or more terete and elongated. Ventral margin more or less terete.
Cleft of the mouth large and horizontal or only slightly ascending. Inter maxillaries extending to the termination
of the margin of the upper jaiv. Teeth in the mouth of fairly uniform size. Preabdominal length less than the

post abdominal. Snout shorter than the postorbital part of the head.

By. the more forward situation of the ventral fins | ceding ones, so far as regards the general rules for the
and the greater elongation of the intermaxillaries this J development of the Teleosteous type in these respects,
subfamily is raised to a higher rank than the two pre- But in the first-mentioned character these fishes come

“ Trybom found a Pearl-side about 7 cm. long in the stomach of a female Herring measuring 311 mm. that was taken in a drift-

net off Vinga on the 8th Sept., 1885.

6 See Day, 1. c., p. 51.

c Collett also found in the stomach of the Pearl-side Ualanus fnmarchicus.

Scandinavian Fishes.

118

936

SCANDINAVIAN FISHES.

nearer to the Herrings, and in luminousness they
are generally less developed. Several of them are
destitute of luminous spots, and the spots in the rows
most regularly present, the ventral, are here, as a rule,
smaller and simpler, whereas the spots of more irregular
occurrence are sometimes highly developed. Most of
these fishes have marked predatory habits, but in this
respect they are surpassed by the fourth subfamily,
the Chauliodontince, which are distinguished by the
large canine teeth in the mouth, and which in the
situation of the ventral fins more nearly approach
to the preceding subfamily. The situation and deve-
lopment of the true dorsal fin separate this subfamily,

as well as the others, from the Paralepidince, where
the said fin is small and set far back, and from the
Alepisaurince, where the dorsal fin extends almost
throughout the dorsal margin.

The name of Saurince has been coined by Gun-
ther", after Sauries , a genus of extensive range in the
Mediterranean and the tropics. The subfamily is the
richest in forms among the Scopeloids, containing
about 50 species, which are distributed among 12 ge-
nera, with or without luminous spots, most of them
belonging to the deep-sea fauna, and some being pe-
lagic fishes. Within the Scandinavian fauna there
occur only two species of the

Genus MYGTOPHUM.

Body Herring- shaped, laterally compressed , covered with

As a generic name need only be a name, and ac-
cording to the rules of the current nomenclature must
be recognised as such, even if it must be condemned
on linguistic principles, the name of Myctophum has
the right of priority, though its formation is erro-
neous or the result of a misprint. Nyctophusb, the
name which Cocco afterwards c suggested as the cor-
rect one, was probably the form which Rafinesque^

rather large scales , and furnished with luminous spots.

intended to write; but an alteration seems hardly ne-
cessary. The genus has been best known under the
name of Scopelus (Cuvier, 1 8 1 7 e) , from which the name
of the whole family has been coined, though as a ge-
neric name it must be rejected.

The genus contains about twenty species of noc-
turnal surface-fishes from the open seas of tropical and
temperate latitudes.

“ Cat. Brit. Mus., Fish., vol. V, p. 394.
b Night-light, Gr. vt'% and (ftog.

c N. Ann. Sc. Nat., An. 1, Torn. II (Bologna 1838), p. 180.
d Bid. d’lttinl. Sicil., 1810.
e Regne Animal, ed. 1, tom. II, p. 169.

SC0PEL01DS.

937

THE GREATER SCOPELUS.

MYCTOPHUM ELONGATUM.

Fig. 234.

Fig. 234. Myctophwn elongatum, together with the under side of its lower jaw. Natural size. From Trondhjem Fjord. Specimen in the
possession of the Zoological Museum of Christiania University, a, b, c, three scales from the left side of the body, X 10. a, from the
row next below the lateral line; b , from the lateral line; c, from the row along the base of the dorsal fin.

im

Ml

■--'S'A'rh''

1 -A

Base of the dorsal fin longer than that of the anal and more than /4 of the length, of the body. Length of
the pectoral fins less than that of the ventral, than either the preabdominal length or the postabdominal , and than
half the length of the maxillaries, which is about half the distance betiveen the dorsal fin and the tip of the snout
or a little greater. Longitudinal diameter of the eyes less than half (< 2/5?) of the postorbital length of the
head, which is more than 3/5 of the entire length thereof. Least depth of the tail more than 8 % of the length
of the body or than 2/5 °f the greatest depth of the latter. Scales of the lateral line slightly, if at all, larger

than those of the body in general.

R. br.S; D. — ;A.— p, P. 1 ; V. 2 ; C. x + 2 + (15— 17) + 2 + x\

’ 19a 1 6 6 10—11’ 7’ v 7

3 1/

L. lat. 43°. L. tr. — - 1.

5

Syn. Scopelus elongatus, Costa, Fna Regn. Nap., Pesc., Malacott.,
Scop., tab. XXXV; Steind., Stzgber. Akad. Wiss. Wien.,
Math. Naturw. Cl., Bd. LNXXIII, i (1881), p. 397; Storm,
Vid. Selsk. Skr. Trondh. 1883, p. 30; Coll. ( Myctoplmm ),
N. Mag. Naturv. Chrnia, Bd 29, p. 104; Lillj. ( Scopclus ),
So., Norg. Fna, Fislc., vol. Ill, p. 25.

Lampanyctus (ex Bonap.) resplendens, Richards., Voy. Ereb.,
Terr., Iclithyol., p. 42, tab. XXVII, figg. 1G — 18; Gthr

( Scopelus ), Cat. Brit. Mas., Fish., vol. V, p. 415; Coll.,
Forh. Vid. Selsk. Chrnia 1880, No. 8, p. 3 cum. tab.; Storm,
1. c. 1879, p. 130; 1880, p. 78.

(?) Scopelus caudispinosus, Johns., Proc. Zool. Soc. Lond. 1863,
p. 42; Gthr, 1. c., p. 416; — Adn. tamen (sec. Jolm-

2F4

somum ) oculos minores (loDg. diam. - long, capitis), pinn.

dors, longiorem (longit. basis = 9/10 distantiaj ex apice rostri;

77-7

long, capitis = longit. basis illius; longit. bas. pinnae

analis = 2/3 longit. bas. pinn. dors.; longit. postabdominalis
= J/2 longit. bas. pinn. dorsalis).

a 1 8, according to Collett.

b 17

x ‘ >7 75

c 38(39) — 50, according to Collett.

938

SCANDINAVIAN FISHES.

Scopelus Kroyerii, Malm, Forh. Skand. Naturf. M. Kbhvn 1860,
p. 617; Vett., Vitt. Samli. Hand!. Gbg., H. 8 (1863), p.
100; Gbgs, Boh. Fna , p. 534; Sundstrom, Fna Sveriges
Ryggradsdj., p. 262 (descr. erron.).

The Greater Scopelus, the larger Scandinavian
species, attains a length of about 15 cm.® As the
form of the Pearl-side reminds us of the Sprat, so the
Greater Scopelus resembles an Anchovy with shortened
snout. The body is rather elongated and compressed.
The greatest depth, which occurs at the ventral fins,
though the whole abdominal region is of fairly uni-
form depth, measures about 1/5 (19 % in the only per-
fect specimen we have been able to examine) of the
length of the body, and the greatest thickness is about
3/5 of the greatest depth. The dorsal contour descends
forward from the beginning of the dorsal tin in a gra-
dual curve to the tip of the snout; but the ventral
contour is straighter in front. The caudal contours
converge at equal angles and to such an extent that
the least depth of the body (in front of the caudal
tin) is about 7u (8‘9 %) of its length, or about i/9
(46'6 %) of its greatest depth. In front the dorsal
margin is convex, the ventral rather flat; behind they
are both about equally compressed, neither being sharp,
however, in front of the spine-like supporting rays of
the caudal fin.

The head is middle-sized, its length being 74 of
that of the body; but the greatest part thereof is oc-
cupied by the temples (posterior cheeks) and opercula.
The postorbital part measures about 63 — 66 % of the
length of the head, and is of fairly uniform thickness,
the sides being almost parallel, but slightly converging
in a downward direction. The forehead is slightly
convex, with a median carina between the anterior
parts of the eyes and on the snout, the sides of which
converge in front almost at a right angle. The eyes,
the length of which is somewhat greater than their
height, are of moderate size in proportion to the length
of the body'', but large in proportion to that of the
snout, which measures at most about 2/3 of their longi-
tudinal diameter. There are, we may almost say,
no true cheeks, only a thin and very Ioav suborbital
ring separating the orbit from the upper jaw; but the
lower margin of this ring is sharply marked both in

front and (very sharply) behind, and entirely conceals
the maxillary throughout the greater part of its length,
when the mouth is closed. The orbital margin is also
sharply defined and especially prominent above, where
it rises at the anterior angle to the frontal plane. The
slightly convex interorbital space is rather broad, though
it becomes narrower in front; at the middle of the eyes
it is equal to, or even greater than, their longitudinal
diameter. The nostrils on each side of the snout lie
on a level with the centre of the eye, just in front of
and beloAv the projecting margin of the anterior frontal
bone, Avhich forms the anterior orbital margin, and are
separated only by a thin dermal septum. The mouth
is large, as in the Anchovy, and the horizontal cleft
thereof extends throughout the greater part of the
length of the head, the length of the upper jaw from
the tip of the snout being 77 — 79 %, the length of the
loAver jaw 80 %, of that of the head. The margins of
both jaAvs are straight and without lips. When the
mouth is closed, only the narroAv intermaxillaries are
visible in front, forming the Avhole margin of the upper
jaw, from the very tip of the snout, where their hardly
perceptible nasal processes meet the rostral tip of the
ethmoid bone. Above them lie the maxillaries, which
are thin and sIioav comparatively little, though gradual,
expansion behind. The latter do not extend quite so
far back as the intermaxillaries, a break being thus
formed at the hind extremity of the upper jaAV. The
loAver jaAv, Avhich projects as far as the upper, resembles,
as a Avhole, a flat boat, when its tAVO branches adjoin
to each other beneath. Their loAver margins are in-
curvated, so that they cover the isthmus underneath
throughout its length, Avith only a narrow, longitudinal
opening between them. The jaw-teeth are small,
pointed, and of uniform size; they are set in a dense
card, narrow but slightly broader at the anterior ex-
tremity, throughout the margins both of the upper
jaw (the intermaxillaries) and the loAver. They also
extend over the outer surface of these margins, rendering
it rough even when the mouth is closed, the inner sur-
face, on the other hand, being smooth. Similar teeth
are set on the long palatine, the entopterygoid, and the
epibranchial bones, the upper and loAver pharyngeals,
and the Avhole series of the copular parts of the hyoid

“16 cm., according to Johnson, assuming that it is this species which he has described.

6 Their longitudinal diameter measures in adult specimens about 61/., % of the length of the body, 23 — 26 / of the entire length
of the head, and 37 — 3 9 1 /2 % of the postorbital length of the head.

GREATER SCOPELUS.

939

bone. There is no true tongue, but the cartilaginous
tip of the lingual bone is rather high and prominent
right in the angle between the branches of the lower
jaw. Here the hyoid arch of each side is attached,
and just behind it the first branchial arch, the inferior
part of which is so long that it extends behind the
branches of the lower jaw. The anterior gill-slits are
also very large; but the last (fourth) branchial arch,
with only one row of lamella?, is united throughout
the greater part of its length to the shoulder-girdle,
the hindmost gill-slit thus being quite short and open
only below. The gill-rakers are setiform. The outer
row on the first branchial arch contains about 25 gill-
rakers, 17 of them belong to the lower, front part of
the arch. The pseudobranchise consist of a longitudinal
row containing about 22 transversely set, filamentous
lamella? on the inner, upper surface of the hyomandi-
bular bone. The opercular apparatus is considerably
prolongated in a backward direction. The preoperculum,
which has no lower (forward, horizontal) arm, lies, as
in the Anchovy, in an oblique, backward and down-
ward direction. Behind, this bone expands into a thin,
dermatoid, membranous margin, which lies over the an-
terior part both of the operculum and the suboperculum,
and this membranous margin is covered with scales
pierced by a branch of the lateral line. The shape of
the quadrangular operculum is adapted to that of the
preoperculum, the former being also oblique, rhombic,
and obliquely set, in the same direction as the preoper-
culum. The hind extremity of the opercular apparatus
is formed by the upper posterior corner of the trian-
gular suboperculum, the lower posterior side of which
is indented, and at the lower angle of which lies the
small, rounded, and scale-shaped interoperculum. The
black branchiostegal membrane of each side is entirely
separate from that of the other side, but is closely ap-
plied and united by connective tissue to the inner sur-
face of the branch of the loAver jaw, thus forming, as it
were, a lining thereof, and hardly projecting beyond its
margin. The eight slender branchiostegal rays increase
in length backwards, but are comparatively short. On
the hyoid arch lie the three luminous spots that shine
through the branch of the lower jaw, the first at the
base of the foremost three rays, the second on the space
between the fifth and sixth rays, and the third on the
membrane between the bases of the seventh and eighth
rays. The gill-opening on each side extends upwards
beyond the superior margin of the operculum, and the

distance from the tip of the snout to the upper angle
of each gill-opening is equal to the postorbital length
of the head.

The true dorsal fin is rather large and of a quad-
rangular shape, with undulating upper posterior margin.
The fin begins at a distance from the tip of the snout
measuring about 38 % of the length of the body. The
length of its base is about 28 %, and that of its longest
ray (the fourth or fifth) about 18 %, of the length of
the body. The first four rays are simple — the first
ray, which is very small, also unarticulated. Among
the remaining rays the anterior (13 or 14) are branched
only once and at the tip, the posterior (6 or 5) twice
and more deeply. The adipose fin, which is short but
rather high, only slightly lower than the hindmost dorsal
rays, lies about half as far from the dorsal fin as from
the first upper supporting ray of the caudal. The anal
fin is similar in shape and structure to the true dorsal,
but much shorter, with only three simple rays at the
beginning. The first ray is so small that it easily es-
capes attention, and the third is the longest in the whole
fin, or equal in length to the first branched ray. The
distance between the beginning of the anal tin and the
tip of the snout is about 54 %, and the base of the fin
measures about 20 %, of the length of the body, the
base thus terminating a little behind the perpendicular
from the adipose fin. The height of the anal fin (the
length of the third or fourth ray) is about 16 % of the
length of the body. The caudal fin, which is deeply
forked, is remarkable in this species too for the short
but strong supporting rays, true spinous rays, that arm
the dorsal and ventral margins of the tail in front of
the true base of the fin. In the only perfect specimen
within our reach there are both above and below 10
spinous rays, gradually increasing in length behind, and
2 articulated but simple rays, the first about one-third
as long as the second, which extends nearly to the tip
of the fin-lobe. The remaining 16 rays are articulated
and branched, the middle ones about one-third as long'
as the outermost, and measuring about 7 % of the length
of the body; but the base of the fin is covered at the
sides, at least half-way along the fin, with elongated
scales, concealing the roots of the rays.

Among the paired fins the pectorals are pointed,
shorter and narrower than the ventrals, which are rounded
and of average size. The length of the former is about
9 of the latter nearly 12 of that of the body.
The first (uppermost) ray in the pectoral fins is simple

940

SCANDINAVIAN FISHES.

and only a little shorter than the second, which is the
longest in the fin and branched like the remaining 9 or
10 rays, which rapidly and regularly decrease in length
towards the bottom of the fin. In the ventral fins the
first (outermost) two rays are simple, the first ray
being, however, extremely small (rudimentary). The
second simple ray is but slightly shorter than the third
ray, which is branched and only a little • shorter than
the next, the length of which is equal to, or only a little
less than, that of the third and fourth branched rays.
'Fhe other branched rays (5th — 7th) gradually decrease
in length. Along the inner margin of the last ray runs
a thickened growth of fin-membrane, with the appear-
ance of a defectively developed ray. The under sur-
face of the ventral fins seems, like the sides of the
caudal fin, to be clothed with elongated scales.

The whole body is densely covered with scales,
which extend forward on the head over the whole inter-
orbital space, but are deciduous, the specimens taken
being generally scaleless, or with only a part of the
lateral line left. The scales (fig. 234, a) are thin, trans-
parent, and flexible, rather large and of equal size,
rounded or of a broad elliptical shape, set in the trans-
verse direction of the body (with the vertical diameter
greater than the horizontal); their nucleus is central;
they are furnished with dense concentric striae and a
few (4 — 6) grooves radiating forwards and backwards.
These grooves, between which the concentric striae (thick-
ened and elevated lines) are sharply curved, render the
anterior margin of the scale sinuate, the posterior (free)
margin irregularly erose. In the nucleus of the scale and
the immediate neighbourhood of the nucleus, as ’well as
in an angular patch coinciding with the margin of the
dermal follicle, the concentric striae are joined at right
angles by transverse striae, and in the hind (free) part of
the scale they are irregularly broken up here and there,
crossing each other in a network. The scales of the
lateral line (fig. 234, b) have a short, pear-shaped duct
at the centre of their outer surface, and are indented
in a heart-shaped form, but deeply, at the hind (free)
margin, by a narrow sinus, which widens, however, in
front (towards the centre of the scale) in a stilliform
manner. Pointed and elongated scales appear not only,
as we have mentioned above, on the sides of the caudal
fin and the under surface of the ventrals, but also in a
row along the base of the dorsal fin, where they are
more or less deformed (fig. 234, c), as if they were
only half-scales, with laterally set nucleus and an in-

creased number of grooves (about 9) obliquely radiating
towards the anterior margin.

According to Costa’s figure the coloration of the
whole dorsal side (including the upper part of the head)
and the lower posterior part of the tail is brownish red,
of the opercular apparatus shifting green and black, of
the ventral side dark brown, of the sides of the head
silvery. The few scales still adhering to the specimen
kindly lent us from Christiania Museum suggest that the
silvery lustre extended throughout the sides of the body;
but where the scales are wanting, the skin of the sides
(in spirit-preserved specimens) is also brownish red.
Back from the occiput, sometimes to a line with the
adipose fin (according to Storm), there run on each side
one or two rows of orange spots, caused, according to
Collett, by the fact that each scale in the two rows
next above the lateral line bears a similar spot at its
tip. These spots are visible, however, though faint, in
the skin, after the scales have dropped off. Costa
figures some of them, along the base of the dorsal fin,
as luminous spots.

The true luminous spots are here (in spirit-pre-
served specimens) of a greenish lustre. The largest on
each side lies on the lower part of the hind preoper-
cular margin. The three spots belonging to each hyoid
arch we have already mentioned. The remaining spots
are arranged, in the specimen now before us, as follows.
Behind the shoulder-girdle, down to the isthmus, where
the lowest spot lies just behind the posterior extremity
of the lower jaw, runs a row of spots, the third from
above being situated at the lower angle of the pectoral
fin. In front of the ventral fins (on the preabdominal
region) we find, on each side of the median line of the
belly, a row of 3 luminous spots, and up the side of
the body this row is continued obliquely backwards by
a spot just before the outer angle of the insertion of
the ventral fin and another set halfway up towards the
lateral line. Behind the ventral fins (on the postabdo-
minal region) there runs, on each side of the median
line of the belly, from the hind (inner) angle of the
insertion of the ventral fin to a line with the anal aper-
ture, a row of 5 spots, and this row is continued up
the side of the body by 3 spots, the uppermost of which
is set on the lower portion of a scale in the lateral
line. Along each side of the base of the anal fin is a
row of 9 spots, and on the lower part of each scale in
the lateral line above the end of this row there lie 2
spots. After a break measuring twice the distance be-

ARCTIC SCOPELUS.

941

tween each pair of spots in the row at the base of the
anal tin, there follows, on each side of the lower caudal
margin, a row of 5 spots, both rows approaching each
other behind; and after new interruptions there lie, on
each side of the lower caudal margin, one spot just in
front of the foremost supporting ray of the caudal tin,
two at the side of the 4th — 7th supporting rays, and
one at the middle of the base of the lower caudal lobe".

The Greater Scopelus is a rare species, of which,
wherever it has appeared, only solitary individuals have
been found. It has most frequently been met with in
Trondhjem Fjord, most often in two inlets, Orkedal
Fjord and Gulos, that run south, with a common en-
trance west of Trondhjem, from the main fjord, where
only one specimen has been taken, at Iiissen, a little
more to the west. All the five specimens secured by
Storm in the winter of 1879 — 80 and the summer of
1881 measured 1 4 x/2 cm- i*1 length, and were taken in
Herring-nets. Further south, in the Skager Rack, two

specimens have been found. Ekstrom received the one
(Malm’s Scopelus Kroyerii) in April, 1856, from a fisher-
man who had found it in the stomach of a Cod taken
on a long-line near the Skaw. The other specimen
was procured at Stromstad by Baron Ckdkrstrom in
April, 1870, and forwarded by him to the Royal Mu-
seum; it had been taken in the same manner in Roster
Fjord. On the Atlantic coast of Europe the Greater
Scopelus has never been met with further south, though
it seems to be one of the Herring’s companions, and
might consequently be expected to occur there as well.
South of the Herring’s geographical range, however, it has
been found in the Mediterranean (off Sicily, from which
locality it was first described, by Costa; and off Nice, as
described by Steindachner), off Madeira (according to
Johnson), and in the Gulf of Guinea (according to Gun-
ther); but as yet it must be reckoned everywhere among
the rarest fishes. In its manner of life, which is other-
wise unknown, it probably resembles Maurolicus Muller '/.

THE ARCTIC SCOPELUS.

M YCTOPHUM GLACIALE.

Fig. 235.

Base of the dorsal fin shorter than that of the anal , and less than lf of the length of the body. Length of the
pectoral fins greater than that of the centred , than either the preabdominal length or the postabdominal, and more
than 4/5 of the length of the maxillaries, which is perceptibly less than half of the distance between the dorsal fin
and the tip of the snout. Longitudinal diameter of the eyes more than half (> 2/3 ?) of the postorbital length of
the head, which is less than 3/5 of its entire length. Least depth of the tail less than 8 % of the length of the
body, or than Vs of the greatest depth of the latter. Scales of the lateral line perceptibly larger than the others.

Fig. 235. Myctoplmm glaciate , together with the under surface of its lower jaw. Natural size. From Hardanger Fjord.

Specimen in the possession of Bergen Museum.

R. hr. 8 ; D. : Ac b P. V.~; C. <c + 2 + 17 + 2 + *;

Lin. lat. 36; Lin. tr. —1; Vert. 36°.

4

Syn. (?) Labrus exoletus , Fabr., Fna Groenl., p. 166 (vide Jor-
gensen, apud Kroy.).

i Scopelus glacialis, Reinh., Vid. Selsk. Naturv., Math. Afh., vol.
VI, p. CX; vol. VII, p. 126; Kr. in Gaim., Voy. Scand.,

a Just in front of each eye, within the orbit, lies a light-coloured mass, with black limits, and similar to a luminous spot, though it
is perhaps simply analogous to the growth of adipose membrane common in the Salmons and Herrings. To avoid damaging the specimen,
we have refrained from examining this mass. It calls to mind, however, the large, composite luminous organ that appears on the snout of
Scopelus metopoclampus etc.

6 A. 16, according to Day.
c According to Kroyer.

942

SCANDINAVIAN FISHES.

Lap., cett., Poiss., tab. 16, fig. 2; Id., Naturh. Tidskr. Kblivn,
ser. 2, vol. II, p. 230; Nilss., Skand. Fna, Fisk., p. 483;
Gthr, Cat. Brit. Mus., Fish., vol. V, p. 407 ; Id., Deep
Sea Fish., Chall. Exped., p. 196; Ltkn, Vid. Meddel. Naturh.
For. Kbhvn 1891, p. 203“.

Scopelus Mulleri, Coll, (nec Gmel.), Forh. Vid. Selsk. Chrnia
1874, Tillsegsh., p. 152; 1878, No. 4, p. 23; No. 14, p.
104; 1879, No. 1, p. 84; Id., N. Nordh. Exped., Zool.,
Fiske, p. 158; (?) Br. -Goode, Bean, Bull. Mus. Comp. Zool.,
Harv. Coll., vol. X, p. 222; Br.-Goode, Fisher., Fisher,
lndustr. U. S., Dab. 203; Day, Nature, vol. XXXIV, Oct.
14, 1886, p. 571; (?) A. Agass., Bull. Mus. Comp Zool.,
vol. XV, p. 33, fig. 219; Lillj., Sv., Norg. Fna, Fisk.,
vol. Ill, p. 20.

The Arctic Scopelus seems never to attain the size
of the preceding species. Kroyer had specimens from
Greenland 46 — 96 mm. in length. The only specimen
we have been able to examine, kindly lent us by the
Museum of Bergen, has probably measured 77 or 78
mm., though its length cannot be stated with certainty,
the caudal tin being broken off short.

The difference from the Greater Scopelus is mani-
fested in the very form of the body. The body is
deepest at the occiput or just behind this point; and
the depth then decreases almost rectilineally to the base
of the caudal fin, the profile of the head being arcuate
both above and below, with the tip of the snout about
halfway up. The body is also thickest at the occiput
and shoulders, growing regularly thinner towards the
base of the caudal fin. The sides of the body are fiat,
and parallel, back from the eyes. The dorsal margin
is convex, as well as the top and bottom of the head,
but the under surface of the abdomen is flat. The
greatest depth of the body, in proportion to its length,
is only slightly greater than in the preceding species,
in our specimen little more than 21 % of the length
of the body excluding the caudal fin* * * * * 6; but the thick-
ness is perceptibly greater, nearly lo1/2 % of the length
of the body excluding the caudal fin, so that the great-
est, thickness is here nearly 2/3 (63 1/2 %) of the great-

est depth0. The least depth of the body, on the other
hand, is less, being only slightly more than 1/3 (in our
specimen 35'2 %d ) of the greatest depth.

Owing to the short form of the body, the length
of the head is also relatively greater, in our specimen
29*2 % of the length of the body excluding the caudal
fine, in spite of the fact, that the postorbital length of
the head is distinguished in this comparison just, for
its smallness, measuring only 15 % of the length of the
body excluding the caudal fin/, and but slightly more
than half (in our specimen 51*5 %/) of the entire length
of the head. The eye, on the other hand, is compara-
tively larger, its longitudinal and vertical diameters,
which are equal to each other, measuring Vio °f Mie
length of the body excluding the caudal fin7i. In front,
at the round slope towards the snout, the forehead is
rather deeply concave, but furnished with the same
sharp, longitudinal carina at the middle as in the pre-
ceding species. The gape is smaller and somewhat more
ascending than in the Greater Scopelus. The difference
in size is indeed hardly perceptible in proportion to the
length of the body, the length both of the upper jaw
and the lower being 1/5 of that of the body excluding
the caudal fin1; but in proportion to the length of the
head this difference is all the greater, the jaws here ex-
tending only a little behind the perpendicular from the
posterior orbital margin. The length of the snout is
more than Vs (24 %J) of that of the jaws. The lower
jaw projects slightly beyond the tip of the snout,, where
its point, fits, when the mouth is closed, into a cavity.
The maxillary on each side extends a little further back
than the intermaxillary, the narrow hind part of which
lies outside and above the lower margin of the posterior
extremity of the former, which here expands into a,
triangle, truncate behind, and thus forms the hindmost
portion of the margin of the upper jaw. The branches
of the lower jaw grow even further inwards than in

° Lutken supposes with Lilljeborg that “Collett has shown that Strom’s communication to O. F. MDller was based upon his

(Str6m’s) drawing”; whereas it appears, from Collett’s own quotation from Strom’s manuscript, that the said communication must have had

reference to Maurolicus Midlen Strom sent his figure and description (with Maurolicus characters) to Muller “more than 20 years before

1791” (see Skrivt. Naturh. Selsk. 1793, p. 15); but did not discover his first specimen of Myctoplium glaciale until 1774 (according to the

manuscript quoted by Collett).

6 Answering to 2 0 1 / 2 % 'in the preceding species.
c Answering to 57 — 58 % in the preceding species.
d Answering to about 46 % in the preceding species.
e Answering to 27 — 26 % in the preceding species.

V Answering to 18 — 16 °/0 in the preceding species.

g Answering to .62 — 66 % in the preceding species.

h Answering to 7—6 °/o in the preceding species.

1 Answering to 21 — 2lY2 % in the preceding species.

J Answering to 15 — 18 % in the preceding species.

ARCTIC SCOPELUS.

943

the preceding species, being almost entirely contiguous
when the mouth and gill-openings are closed, the only
space between them then consisting of a lanceolate gap
at the extreme front, just behind the symphysis.

The dentition of the mouth and pharynx is weaker
than in the preceding species, otherwise of similar type.
The preoperculum is vertically set, slightly curved, with
the hind margin expanded but extremely thin. The
operculum, which is half free at the upper margin, is
quadrangular with rounded corners and smaller than
the suboperculum, which lies below and inside it; but
these bones, as well as the interoperculum, are so thin
and so densely covered with large scales that it is diffi-
cult to determine the' limits between them and their
shape without damaging the solitary specimen at our
disposal. The branchiostegal membranes, with their 8
rays, are united to each other below and in front, but
only for a very short distance, to a line with the centre
of the eyes. The pseudobranchise are distinct and re-
gular. The gill-rakers resemble those of the preceding
species, and number 16 or 17 in the outer row on the
first branchial arch. The last branchial arch of this
species too is mostly coalescent with the clavicular arch,
the hindmost gill-slit being consequently very small.

In contradistinction to the preceding species, the
Arctic Scopelus belongs to a subdivision of the genus,
proposed by Gunther, in which the dorsal tin is shorter
than the anal. The former fin begins half-way along
the body minus the caudal fin, at a distance from the
tip of the snout measuring about 51 %a of the length
of the body excluding the caudal fin. Its base measures
about 15 %h of the same length. Its first three rays
are simple; the first two unarticulated. According to
Kroyer’s figure the fourth or fifth ray is the longest,
the posterior rays uniformly decreasing in length. The
adipose fin lies about half-way between the dorsal and
caudal fins, is broader (longer at the base) than in the
preceding species, and is supported by fine fibrils (flex-
ible, corneous rods) within it. The anal fin begins at
a distance from the tip of the snout measuring about

“ Answering to 41 / in the preceding species.
b Answering to 30 % in the preceding species.
c Answering to about 59 % in the preceding species.
d Answering to about 22 % in the preceding species.
e Answering to 7 — 8 % in the preceding species.
f Answering to nearly 10 / in the preceding species.

9 In the preceding species about twice this breadth.
h Answering to nearly 1 3 % in the preceding species.
i Answering to about 13 G % in the preceding species.
j Answering to nearly 18 % in the preceding species.

5472 %c of the length of the body excluding the caudal
fin, and its length is about 24 %d of the same. The
first two rays are simple and unarticulated, the first
ray being very small. The branched rays, accordingly
to Kroyer’s figure, decrease uniformly and slowly in
length behind. The shape of the caudal fin cannot be
ascertained from our specimen; but the fin has certainly
been deeply forked, and the length of its middle rays,
here as well as in the preceding species, differs but
little from the interorbital width, being nearly 67a
of' the length of the body excluding the caudal fin. In
our specimen we find 8 supporting (spinous) rays at the
upper margin of the caudal fin and 6 at the lower.
The pectoral fins, which are set obliquely and rather
high, are narrow, but very long, measuring 18 %f of
the length of the body excluding the caudal fin. The
ventral fins, which lie rather near each other, the di-
stance between them being scarcely equal to the breadth
of the base of either fin6', are shorter than in the pre-
ceding species. Their length is 7k/ °f that of the body
excluding the caudal fin. Their distance from the tip
of the snout, here as in the preceding species, is about
2/5 of the length of the body excluding the caudal fin.
The distance between them and the foremost point of
the base of the pectoral fins (the preabdominal length)
is 79\ and that between them and the beginning of
the anal fin (the postabdominal length) about 15 %j,
of the length of the body excluding the caudal fin.
The shortening of the body in this species, as opposed
to the preceding one, thus affects in the most essential
degree the abdominal region.

The scales are comparatively larger than in the
preceding species, but equally thin and of essentially
the same shape and texture. The greatest difference
meets us in the scales of the lateral line, which are
considerably higher (broader). A scale from the middle
of the lateral line in our specimen is 2’1 mm. long and
5'3 mm. broad. Its anterior (inserted) margin is rather
evenly rounded (not triangularly pointed at the middle,
as in the preceding species), with only faint sinuses, and

Scandinavian Fishes.

119

944

SCANDINAVIAN FISHES.

its hind margin is without the deep, narrow incision,
though according to Kroyer’s figure of this species the
incision is sometimes present in the lateral scales. On
the head the scales advance over at least the greater
part of the interorbital space, and large scales, as we
have mentioned above, cover the whole opercular appa-
ratus. On the posterior, expanded part of each maxil-
lary lies an obliquely linguiform scale (triangular with
rounded corners), entirely covering this part; and the
bases both of the pectoral and ventral fins are covered
with several scales.

The true luminous spots are comparatively small
and have a yellowish lustre. The hind margin of the
preoperculum, at the limit between the operculum and
suboperculum, is furnished with a luminous spot, and
exactly below this point, at the upper anterior angle
of the interoperculum lies another. Through each branch
of the lower jaw there shine three spots. On each of
the clavicles there are two spots. One spot is situated
at the lower angle of the insertion of each pectoral fin.
From the isthmus to the ventral fins there runs on each
side of the ventral margin a row of five spots. Between
the insertion of the ventral fins and the beginning of
the anal fin this row is continued by four spots, and
along each side of the base of the anal fin by seven
more. Between the anal and caudal fins, which part is
damaged in our specimen, both Krgyer and Collett
found six spots in a row on each side of the ventral
margin. An upper row on each side of the body, in
the abdominal region and on a level with the spot at
the base of the pectoral fin, contains three spots, all
belonging to the row of scales next below the lateral
line, the foremost of them situated at the end of the
preabdominal region, the other two in the posterior part
of the postabdominal region. On each side of the tail
there also runs a row of three spots, but this lies higher
up, each spot being situated on the lower part of a
scale in the lateral line, the first spot above the space
between the first pair in the row along the base of the
anal fin, the second in a similar relation to the last
pair in the said row, and the third at the extreme end
of the peduncle of the tail. At the dorsal margin of
the tail, just in front of the upper supporting rays of

° See Die Augenahnliche Organe der Fische, p. 52, and Len
or “stern-chasers” of Scopelus-.

This species is similar to Myctophum gJaciale in most respe
gards the tail, the body thus being relatively deeper, and the head la
in the caudal part of the lateral line are wanting.

c See Day, 1. c.

the caudal fin, traces appear in our specimen of a large,
composite, luminous spot, Leydig’s Perlfleck or Perl-
grauer Fleck". At the middle of the interorbital space
lies a spot (an epiphyseal body or a parietal organ?),
which in Scopelus Bissoib, a nearly related Mediterranean
form, has even more of the appearance of a luminous spot.

The coloration of the body, according to Ivrgyer,
is of a brownish green or an olive hue on the back, of
a brassy lustre on the sides (with a. bright golden lustre,
according to Grayc), of a darker olive-green under the
belly. The cavity of the mouth and the branchial ca-
vity, as well as the peritoneum, are black.

The internal organs have been described by Krgyeil
The liver is short but thick, with three lobes, the left
one being the largest; the gall-bladder is small. The
oesophagus is short but wide, the stomach small but
somewhat saccate. Behind the pylorus hang 8 rather
large ciecal appendages. The intestine is straight but
rather wide. The saccate air-bladder extends about half-
way along the abdominal cavity, and is connected by its
pneumatic duct with the oesophagus. The generative
organs lie as usual in the posterior half of the abdo-
minal cavity, and have complete efferent ducts.

The Arctic Scopelus has long been known in the
Scandinavian fauna, but is still rarer than the preceding
species. In 1774 a specimen was taken by Strom at
Sondmor; but since that occasion only 3 specimens have
been met with, to the best of our knowledge, on the
coast of Norway, and not a single specimen has ever
been found on the Swedish or the Danish coast. One
of the said 3 specimens — the very specimen which has
been kindly lent us from the Museum of Bergen — was
taken by Governor Christie about 1830, probably in
Hardanger Fjord; the second was sent from Haswig,
near Hammerfest, to Christiania Museum by Mr. Bull;
and the third 'was found by the Norwegian North At-
lantic Expedition in June, 1876, floating at the surface
on the fishing-bank of Storegg off Aalesund. The spe-
cies would seem to be commoner on the coast of Green-
land, from which locality the Museum of Copenhagen
has received several specimens, both from the northern
and the more southern colonies, and in the Arctic Ocean,
where the Norwegian North Atlantic Expedition took it

eneeld, Deep Sea Fish., Challeng. Exped., p. 302, The dorsal organs

:s; but the form of the body is still more shortened, especially as re-

•ger, while the above-mentioned luminous spots belonging to the scales

SALMON- L A U N C E S .

945

while trawling in 1,100 fathoms of water (71° 59 N. lat. ;
11° 40' E. long.), and Captain Gray found it floating at
the surface, but still alive (73° 12' N. lat.; 14° 28’ W.
long.). Whether it was the same species that was met
•with in 1880 — 83 by the United States’ Fish Commis-
sion “off the southern shores of New England” and by
the Blake Expedition (34° — 41° N. lat.; 65° — 75° W.
long., see Beown-Goode and Bean, 1. c.), is doubtful.
No description is extant of the specimens found on these
occasions, and the figures show important differences
from the Arctic Scopelus as described above".

The large eyes of the Arctic Scopelus are unmis-
takable tokens of a life in deep water; and most of the
specimens have consisted of unlucky individuals, borne
with too great rapidity to the surface of the ocean, and

there left to the mercy of wind and wave. But this
species is, no doubt, a nocturnal surface-fish as well,
voluntarily repairing in the darkness to the upper strata
of the sea, for it has frequently been found off the
coast of Greenland in the stomach of seals, which can-
not live in the true deep-sea regions. All that is known
besides of its manner of life is that it feeds on small
crustaceans, though not exclusively on the most minute
kinds, for Collett found in its stomach fragments of
an Amphipod, Themisto lib ell ul a , which attains a length
of 20 — 60 mm. The remaining food, as far as could
be determined, consisted of several specimens of an
Ostracod, Conccecia borealis , which, according to Saks,
has never been found at a depth of less than 300
fathoms.

Subfamily P A IS A L h V I D I N Jf.

Dorsal fin situated considerably behind the middle of the body. Snout longer than the postorbital part of the head.
Pseudobranchice present; but air-bladder and pyloric appendages wanting, or the latter present only in a rudi-
mentary form.

Within this subfamily Gunther6 includes two very
closely related genera of Ammodytes- like fishes, first
known from the Mediterranean, that were united into
one genus by CuvieiU, and ranged among the Acantho-
pterygians beside Spliyrcena, to which they also sIioav
resemblances in form of body. The Salmon-Launces are
elongated and compressed (in general a foot long or less),
silvery, more or less transparent, pelagic fishes, of pre-

datory habits, as testified by the long and pointed
canines, with which their intermaxillaries, lower jaw,
and palatines are generally armed. Ten species have
been described or at least mentioned by name: five from
the Mediterranean and the Atlantic outside, three from
the North Atlantic and the Arctic Ocean, and two from
the Pacific. All these species, however, may well be
contained within the limits of a single genus.

o o

Genus SUDISN

Of this genus one species is indeed known from the
high North, from Greenland and Iceland, namely Sadis
( Paralepis ) borealis e, which is described at length by
Kroyer in the “Naturhistorisk Tidskrift”, 2:den Raekke,
2:det Bind, p. 241, and figured in Gaimaiid’s Voyage

en Scandinavie etc., Poissons , pi. 16, B, fig. 1. But in
Scandinavia this species has not yet been found. On
the other hand, Kroyer has given, in Fiedler and
Feddersen’s “Tidskrift for Fiskeri”, 2:den Aargang
(1868), p. 70, a brief account of the discovery of a

° Both in Todd s figure (Buown-Goode) and in Agassiz the dorsal fin begins much further forward than in the Arctic Scopelus, and
the adipose fin is high and short as in the Greater Scopelus. In Todd s figure the base of the anal fin is considerably longer, in Agassiz
considerably shorter, than in the Arctic Scopelus. Along the base of the anal fin Todd’s figure shows a row of 15 luminous spots, Agassiz’
only 6. The lateral line contains in Todd’s figure 46 scales, in Agassiz 35.
b Cat. Brit. Mils., Fish., vol. V, p. 418.
c Cuv., Val., Hist. Nat. Poiss., vol. Ill, p. 356.

d Rafinesque, Car. Ale. N. Gen., p. 60. Paralepis, Risso, Hist. Nat. Fur. Merid. tom. Ill, p. 472.

e Reinhardt, D. Vid. Selsk. Naturh., Math. Afh., vol. V, p. LXXV; vol. VII, p. 125. Probably Clupea encrasicolus , Fabr., Fn.
Groenl., p. 183.

946

SCANDINAVIAN FISHES.

SUDIS ATLANTICUS,

which was found dead in May, 1865, where it had
been cast ashore near the Skaw. Two specimens are
said to have been observed, after a violent storm from
the west; but only one was secured, and this reached
Iyroyer’s hands in a greatly damaged condition. It
was a large fish, half a metre long, Avith much deeper
body than the other known species of the genus. The
greatest depth was contained only 7 times in the length
of the body. A manuscript published in part by Lut-
ken° after Kroner's death further contains the remarks

that the length of the head was 21 '7 % of that of the
body, that the dentition of the mouth was compara-
tively feeble and uniform, that the articulation of the
lower jaw lay nearer to the perpendicular from the
anterior margin of the eye than in the Greenland form,
Sudis borealis , that the ventral fins were situated below
the true dorsal fin, that the adipose fin was compara-
tively remote from the caudal, that the pectoral fins
contained 15, the anal fin 20 rays, and that the scales
were small, numbering about 20 in a transverse row.

Fam. C L U P E 1 1) M.

Body of the well-known Herring form, but more or less deep or elongated, more or less terete or compressed. Scales
middle-sized or small, thin, and generally deciduous. No luminous spots on the sides of the body. No adipose
fin. Margin of the upper jaw formed by the intermaxillaries in front and by the maxillaries behind. No bar-
bels. Air-bladder simple or internally cellular; its communication ivith the cranial cavity, where such a commu-
nication exists, without mobile osseous connexion. Branchial cavity usually furnished ivith large pseudobranchice.

Ovaries furnished ivith oviducts.

No piscine family is so important to man from an
economical point of view as the Herrings ; and for the
systeinatist too this family possesses great interest.
Around it are grouped a number of families, foreign
to the Scandinavian fauna, that combine its characters
not only with preceding types, but also with Ganoid
peculiarities. Soon after Muller, on the strength of
Vogt’s investigations, had referred the genus Amia to
the Ganoids6, where Agassiz0 had previously ranged
Lepidosteus and Polypterus, which forms as well as
Amia Cuvier1* included among the Herrings, Stannius0
showed that the character upon which Muller had laid
special stress, the double row of valvules at the passage
from the ventricle of the heart to the bulbus arteriosus,
also appears in Butirinus ( Albula ), another of Cuvier’s
Herring-fishes, which is still referred by some authors
to that family. GegenbauiG showed — a thought which

had suggested itself to Muller — that the most essen-
tial difference in this respect between the Teleosts and
the Ganoids together with the true cartilaginous fishes
consists in the following circumstance. In the latter
the ventricle of the heart, with the transversely striped
muscles of its Avail, is prolongated forward to a tubu-
lar, more or less conical chamber ( conus arteriosus ),
on the inner surface of which the valvules in their
more or less numerous rotvs are situated; Avhereas in
the Teleosts the heart is destitute of this prolongation,
the beginning (base) of the branchial artery that issues
from the ventricle being instead dilated into a bulb
( bulbus arteriosus ), without muscles (with only elastic
Avail) or with unstriped muscles alone in its wall.
Stour!7 completed these observations by pointing out
the structure and distribution of the rows of valvules
in different Ganoids, Sturgeons, Chirmeras, and true

“ Vid. Meddel. Natnrh. For. Kblivn 1891, p. 231.
b Abb. Akad. Wiss. Berlin 1844, p. 204.

0 Rech. Poiss. Foss., tom. II, part. II, p. 1.
d R'egne Animal, ed. 2, tom. II, pp. 327 — 329.

e Bemerkungen iiber das Verhaltniss der . Ganoiden zu den Olupeiden, Rostock 1846.
f Jen. Zeitschr., Bd. 2 (1866), p. 365.
g Morphol. Jahrb., Bd. 2 (1876), p. 197.

HERRING-FISHES.

947

cartilaginous fishes, representing different stages of re-
duction. Boas'* followed the same method with the
Teleosts, showing that in Butirinus there is a distinct
rudiment of a conus arteriosus , with transversely
striped muscles and two rows of valvules, and that in
Osteoglossum — now the type of a separate family of
hard-scaled fishes which in other respects come near to
the Herrings — and Notopterus — belonging to a family
also nearly related to the Herrings, but with scaly
head and without oviducts — the said rudiment is con-
siderably smaller and possesses only one row of val-
vules, answering to the front row in the previous fishes.
In the true Herrings the rudiment has almost entirely
disappeared, and is replaced merely by a layer of con-
nective tissue, without striped muscles; and this, as
we have already mentioned, is the typical condition
in the Teleosts.

The essential resemblances between the skeleton
of the Herrings and that of the Salmons were remarked
by L. Agassiz. But in the former the skeleton is, as
a rule, more firmly ossified, and shows some remark-
able peculiarities in systematic and morphological re-
spects.

From a comparison between the skeleton of a Ven-
dace and that of a common Herring, it appears that
the same looseness prevails in the union of the lateral
parts both of the neural arches belonging to most of
the anterior abdominal vertebral and the haemal arches
of the posterior abdominal vertebrae. The latter are
indeed closed throughout a considerable part of the
trunk, an osseous bridge joining the right shaft of the
arch to the left; but lower down the shafts again se-
parate, and the ribs are attached to each part of these
divided haemal spines. The vertebrae of the Herring
are further marked — as anyone who lias eaten a Her-
ring knows — by the number of scleral bones deve-
loped in the tendinous walls (aponeurotic septa) between
the transverse divisions (myocommata) of the great
lateral muscles. A perfect abdominal vertebra (fig. 236)
has no less than three pairs of such bones, the first
(na = epineural bones) attached to the base of the neural
arch or at the limit between that arch and the body
of the vertebra, the second (pa ----- epicentral bones) to
the body of the vertebra, and the third (pta — epipleural
bones) to the base of the ribs or at their insertion. In

the Salmonoids we find only one or two pairs of these
scleral bones, the uppermost alone or in association with
the lowest. In most Clupeoid forms the lower extre-
mities of the ribs in general are applied to the inner
surface of osseous growths from the skin, to which they
are, however, but loosely united. These growths belong,
as we have seen above in the case of the Sternopty-
choids, to the median scales of the carinated ventral mar-
gin, the so-called ventral plates, with their spiniform
processes in a backward direction (dh).

In the Herring too a great part of the chondro-
cranium is continuous. The long frontal bones (fig. 237,
fr) send out obliquely backwards and downwards, on
each side behind the orbits, a large process, which runs
into the squamosal bone ( squ and gos in the figure, os
pteroticum ), while the frontal bone itself behind touches
the parietal bone (par). Between the said three bones
on each side of the head the cranial wall is pierced by

Fig. 236. Abdominal vertebra of a Herring. After Brandt and Rat-
zeburg. ns, upper spinal process (neural spine); n, upper (neural)
arch; c, corpus; p, transverse process; pi, rib; na, upper extracostal ;
pa, middle extracostal; pla, lower extracostal; dh, spiniferous plate
at the ventral margin.

the large and oblong temporal aperture (ap)b charac-
teristic of the Herrings. This aperture lies in front of
the mastoid groove (temporal cavity), which is rather
large in these fishes as well as in the Salmons and Carps,
its bottom being composed of the squamosal bone, car-
tilage, and the mastoid bone (mi in the figure, os epio-
ticum), while its walls belong partly to these bones,
partly to the lateral portions of the squamosal part of
the occipital bone (ocs). It is besides furnished with a
roof formed by a backward process from the parietal
bone (par), meeting a forward process from the mastoid
bone (mt). The greater part of the roof of the cranial
cavity has lost its cartilage in the Herring, only a lon-

“ Vid. Meddel. Naturh. For. Kbhvn, 1879 — BO, p. 333.

6 The temporal aperture sometimes occurs, it is true, in the Salmon (cf. Bruch, Osteol. Rheinlachs, tab. IV, fig. 3, between 0 and z/), and
appears in the Smelt as a vestige of the great frontal fontanel in the cartilaginous cranium of young fishes. But it is there covered by the frontal bone.

948

SCANDINAVIAN FISHES.

gituclinal bridge thereof running under the frontal suture
from the anterior extremity of the triangular squamosal
(upper) part of the occipital bone forward to the trans-
verse epiphyseal bridge of cartilage. In front of the
latter bridge too the cartilaginous roof under the frontal
bones is wanting above the olfactory lobes; but further
forward, above the orbitosphenoid bone (obsp), begins
the compact ethmoidal cartilage, extending forward to
the tip of the snout. The last-mentioned cartilage, with
its two terete transverse processes on each side, one in
front of ( pcna ) and one behind ( pcnp ) the nasal cavity,
resembles that of the Salmons; but it rises anteriorly
in a carina, and its covering bones, the true ethmoid
bones, the vomer, and the lateral ethmoid bones ( etl ) are
more developed. The ethmoid bones proper extend over
the anterior transverse process of the cartilage {pcna,
the palatine process, on which the anterior extremities
of the palatines and, partly, of the maxillaries have
their articular surfaces), and each of them sends out
backwards, on the upper surface of the cartilage, below
and between the frontal bones, a long process, extend-
ing to about a line with the middle of the orbits. The
lateral ethmoid bones (etl), covering the hind transverse
process (pcnp) of the ethmoidal cartilage, spread like
wings on either side. The vomer ( vorn ) is narrow and
long, being pointed behind and somewhat longer than
the upper ethmoid bones. The parasphenoid bone (psp)
is principally characterized by its complete division into
two vertical plates behind the orbits, under the petrosal
bone (the bone round goa, os prooticum) and the basilar
part ( ocb ) of the occipital bone, and by its great elonga-
tion in a backward direction, where it projects beyond
the last-mentioned bone. In the Salmons Ave found, it
is true, that the hind orbito-muscular canal Avas rather
large and open behind; but here the canal is still larger
and entirely open, even beloAv, its sides being formed by
the vertical posterior plates of the parasphenoid bone
and correspondent lamelliform processes descending from
the petrosal and occipital bones, Avhich in their turn
form the roof of the canal (see tig. 237, C).

As the temporal aperture is a characteristic of the
upper part of the cranium in the Clupeoids, so Ave find
beloAv a large and characteristic foramen (fen), covered
only by the mucous membrane of the branchial cavity
(upper pharynx). It occupies the point Avhere the pet-
rosal bone and the basilar and lateral ( ocl ) parts of the
occipital bone Avould otherAvise meet, and is separated

by a special osseous septum (the bone round gop) from
the foramen behind it (in the lateral part of the occi-
pital bone), through Avhich the nervas vagus passes out
of the cerebral cavity. Hassk" explains this foramen
(fen) as “the first appearance in the animal kingdom of
that most important organ, the vestibular AvindoAv” (fene-
stra ovalis), the communication between the vestibule
and the tympanic cavity in the higher animals; and on
the inner side thereof lies the sacculus, belonging to the
nervous labyrinth, Avith the large otolith. The difference
from the Salmonoids stands out most sharply on a com-
parison of this part of the Clupeoid cranium Avith the
analogous part in the Smelt for example, Avhere the
foramen is replaced on the under side of the cranium
by a large swelling, including the sacculus, and formed
by the above-mentioned bones.

In the bones surrounding the auditory apparatus of
the Herring avc meet, hoAvever, Avith an appendage of
which there is no vestige in the Salmonoids, but Avhich
is probably analogous to the several lymphatic chambers
in the neighbourhood of the labyrinth that have been
noticed in the skull and the front of the spinal canal
of the Cyprinoids, and Avhich there communicate Avith
processes of the air-bladder. This appendage consists,
on each side of the skull, of three osseous globules (goa,
gos, and gop), filled with air and lined Avith a thin, sil-
very membrane. Their structure is more compact than
that of the neighbouring bones, and their Avhite colour
renders them easily distinguishable. They all lie in
different planes, one above another; but from the loAver
posterior globule (gop) runs an osseous duct, which rami-
fies and sends out a branch to each of the other tAvo.

The foremost and largest of these globules (goa) lies in the pet-
rosal bone, just behind the passage for the nervus trigeminus (ftr),
in the lateral part of the cranial floor, where the said bone sends out
its leaf-shaped process, down towards the parasphenoid bone, to share
in the structure of the side-wall bounding the anterior part of the hind
orbito-muscular canal. The outer posterior, uppermost globule (gos),
which is skirted by the horizontal (outer), semicircular canal of the
labyrinth, lies in the squamosal bone, just within the hind part of the
articular cavity for the hyomandibular bone and outside the opisthotic
bone (st, os opisthoticum or intercdlare) . The lower posterior globule (gop),
the smallest and most elongated (clavate), is situated in the lateral part
of the occipital bone, between the foramen of the nervus vagus (fv)
and the fenestra ovalis (fen), forming the very osseous septum between
these holes that has just been mentioned. The osseous shells of these
globules and its cavity are indeed continuous; but whether the cavity
of the membranous lining (sac) also affords at all times a free com-
munication between them, is a question which we must leave open,
for in one specimen we have succeeded in preparing the sac of the
lower posterior globule whole and, as far as we could see, without
any opening in front. The anterior globule (goa, the petrosal globule)

Anat. Stud., Bd. I, p. 600.

HERRING-FISHES.

949

is partly (somewhat more than half) divided within by an incomplete,
horizontal septum, only the sac of the lower part being thus continuous
with those of the other globules, while the upper part (the part nearest
to the cranial cavity), which is pierced behind by a transversely-set,
oblong aperture with a raised rim at the upper margin, is filled by an
extension of the lymph-space surrounding the auditory apparatus. In
the anterior part of the globule, however, this extension lies close to
and above the said sac. The osseous wall of the lower posterior glo-
bule ( gop ), on the other hand, forms a tubular prolongation backwards

and downwards to a line with the middle of the occipital bone, where
its inner sac is continued by a duct, with a more or less cartilaginous
coating, that joins the anterior extremity of the air-bladder, though
without free communication with the cavity thereof. All our attempts
— like those of Valenciennes'* — to inject liquid either from the air-
bladder to the said globule or in the reverse direction, have failed;
but a fine bristle may be passed with ease from the auterior osseous
globule ( goci ) and through the lower posterior (gop) into the duct and
down to the wall of the air-bladder.

vom

fen

B

eiv

vom

nit

gos-
goci -

■ - - gos
ocl

st

ocb

C

Fig. 237. Skull of a Herring X 2. A from the left; B from below; C from behind.

et , ethmoid bone; vom, vomer with its teeth; etl, lateral ethmoid bone; pcna, anterior nasal process of the ethmoidal cartilage; pcnp , posterior
nasal process of the same; fr, fr, frontal bone; psp, psp , parasphenoid bone; obsp , orbitosphenoid bone; alsp, alispbenoid bone; sp/io , posterior
frontal bone (os sphenoticum) ; ap, temporal aperture in front of the parietal bone; par, parietal bone; mt, mastoid or epiotic bone; ocs.
squamosal part of the occipital bone; ocl, lateral part of the occipital bone; ocb, basilar part of the occipital bone; st, styloid, intercalary,
or opisthotic bone; fv, foramen of the tenth cranial nerve (nervus vagus'); ftr, foramen of the fifth cranial nerve (nervus trigeminus)-, fen,
labyrinthine fenestra; goci, auditory globule of the petrosal bone (globulus osseus anterior ); gos, auditory globule of the squamosal bone (glo-
bulus osseus superior); gop, auditory globule of the occipital bone (globulus osseus posterior).

Cuv., Val., Hist. Nat. Poiss., tome XX, p. 41.

950

SCANDINAVIAN FISHES.

By means of these globules and the ducts between
them the wall of the air-bladder (but not its cavity, at
least in full-grown Herrings") is connected with the
perilymphatic spaces surrounding the auditory apparatus,
and variations of pressure may thus be transmitted from
the former to the latter. The connexion is, however,
comparatively simple, simpler at all events than in the
Cyprinoids, and, as we have mentioned above (p. 826),
this structure in the Clupeoids has consequently been
regarded from a morphological point of view as a pri-
mitive stage of development of the corresponding appa-
ratus in the Carp-fishes. Another characteristic of the
Clupeoids is the connexion of the air-bladder not only
with the stomach, into whose posterior, pointed extre-
mity the pneumatic duct opens in the Herring, but also

A

Fig. 238. Left scapular arch and pectoral fin of a Herring.

behind the vent, may help us to explain the circum-
stance that the air-bladder is sometimes found filled to
distension with water0.

The structure of the mouth is essentially the same
as in the Gwyniads. The shape and position of the
inter m ax illaries in particular show striking similarities
between the Herring and the Vendace. Here, however,
these bones are not applied so closely to the maxillaries,
which are distinguished by the accession of a further
supplementary bone, situated in front of and (partly)
outside the posterior one, which, as well as the maxillary
itself, resembles in shape and position that of the
Gwyniads.

Among the remaining peculiarities of the skeleton
we will only touch upon the shoulder-girdle, which has

pt -\

Natural size. A, from within ; B, from without.

pt, posttemporal bone; scl, supraclavicular bone; cl, clavicle; clp, ascending process from the anterior margin of the clavicle; sc, scapula;
cr, coracoid bone; per, prrecoracoid bone; pclx — 3, the several postclavicular bones; br, brachial bones (inner row); bre, outer (distal) row

of brachials.

with the surrounding water, the hind part of the air-
bladder being continued by a duct which opens on the
posterior side of the triangular dermal flap forming a
prolongation of the hind margin of the anal aperture* * 6.
The connexions of the air-bladder with the nearest en-
vironments of the labyrinth, and the large temporal
aperture, which interposes only a thin and soft wall be-
tween those environments and the surrounding water,
may well be the cause of the sensitiveness to noise so
highly characteristic of the Clupeoids. The communi-
cation with the surrounding water, through the opening

a That the globules and ducts are excrescences of the air-bla

than probable; but we know of no direct observation on this head.

6 Cf. Weber, De aure et auditu, p. 73; Bennett, Journ. An
vol. VI (1883, Jan.), p. 24.

c Cf. Guv., Val., 1. c., tom. XX, p. 70.
d Shoulder-Girdle and Sternum, Roy Soc. 1868, p. 57.

guided Parker1* to some morphological conclusions
highly worthy of attention. The postclavicle is not only
divided, as in the Salmon, into a row of separate bones
(in the Herring three, tig. 238, B, pclt, 3), but also
situated outside the clavicle (cl), an arrangement known
only in the Herrings. How closely the shoulder-girdle
of the Herring is connected with the dermal growths,
a, sign of its primitive rank from a morphological point
of view (cf. above, p. 635), further appears both in the
thin, squamoid texture of the upper postclavicular bones
at the hind margin, and in the still more squamoid,

ler and in an embryonic state freely communicate therewith, seems more
Physiol., vol. XIV (1879 — 80), p. 405; Day, The Zoologist, ser. 3,

HERRING -FISHES.

951

leaf-shaped extension of the posterior part of the supra-
clavicle ( scl ), on which too the posttemporal canal of
the lateral line is sometimes continued. In the appa-
ratus of the pectoral tins we find a peculiarity other-
wise unknown among the Teleosts, but characteristic of
the Ganoids, namely the outer row (bye) of brachials
(basal bones of the pectoral fins) possessed by the
Herring®.

The organs of respiration are exceedingly well de-
veloped in the Clupeoids; and they are protected as a
rule by numerous and long gill-rakers. The pseudo-
branchne are large, and in many forms'' we find on each
side a separate, accessory branchial cavity, resting on
the hindmost (fourth) epibranchial bone, which is flat-
tened out into a plate for that purpose".

The digestive organs are as a rule comparatively
simple, as in the Salmons. In all the Scandinavian
Herrings the short oesophagus passes without external
distinction into the stomach, which is prolonged into a
sac and pointed behind; and from the anterior part of
this sac issues in a forward direction the more or less
muscular pyloric portion, bending back quite as sharply
to pass into the intestine, which is fairly straight and
runs without any convolutions to the vent. In exotic
forms however — as for example in the Brazilian Clu-
pea anchovia — the wall of the pyloric portion is some-
times thickened, in the same way as in the Mugiloids
(see above, p. 329), into a sort of ingluvies, and the
prolongated intestine forms several convolutions. Other
Clupeoid forms show no backward prolongation of the
stomach. The pyloric appendages are generally well de-
veloped, but quite as variable as in the Salmonoids.
Even among the Scandinavian species the variations in
the number of ca^cal appendages behind (below) the
pylorus are very great, the Sprat for example having
6 — 13, the common Herring 20 — 24, and the Shads
about 80. In some exotic Clupeoids d the anterior por-
tion of the intestine (the duodenum), into which the
pyloric appendages open, is so greatly prolonged that
bunches of these appendages occur almost throughout
the length of the abdominal cavity. The mucous mem-
brane lies, as usual, in longitudinal folds within the
oesophagus and stomach; but in the intestine the folds,

which are most developed, among the Scandinavian Clu-
peoids, in the Shads, are transverse. The liver is gene-
rally small, sometimes exceedingly small; the spleen
commonly long and narrow. The saccate organs of
reproduction ■with their complete excretory ducts are
so large during the time of propagation that they fill
the greater part of the abdominal cavity; but at other
times, as in the generality of fishes, they are contracted,
sometimes scarcely visible. The connexions of the air-
bladder we have already mentioned, and we will simply
add that the pneumatic duct sometimes opens into the
posterior end of the stomach, sometimes further forward
or into the oesophagus. In some cases too the wall
of the air-bladder lies so close to the dorsal side of
the stomach that the said duct is merely a very short
funnel.

The scales are generally thin, deciduous, and cy-
cloid, smooth-margined, or, not unfrequently, notched or
even fringed with sharp spines at the hind margin. The
head is always scaleless, as well as the fins in general;
but the scales extend rather often over a great part of
the caudal fin and the base of the dorsal, more rarely
over that of the anal, and the axillary scales of the
pectoral and ventral fins are in most cases well developed.
Especially characteristic of most Herring forms are the
angular, spinigerous scales at the abdominal margin,
which send out lateral processes to meet the lower ends
of most of the ribs (fig. 236, dh; fig. 240, p. 958).

The system of the lateral line shows great deve-
lopment, accompanied by a wide extension of the so-
called adipose membrane, on the head; but the lateral
line proper is wanting on the sides of the body, with
the exception that in the common Herring we some-
times find the first (3 — 5) scales in one of the rows
pierced in the ordinary manner. In certain forms, as
for instance the Shads, the lateral line again crops up in
some scales of singular shape, situated on the caudal fin".

The fins of the Clupeoids display the same wealth
of variations as those of the Carp-fishes, and here too
the variations principally affect the vertical (unpaired)
fins. The dorsal, which is never very long, sometimes
suffers reduction to the vanishing point; the anal, on
the other hand, is sometimes exceedingly long, occa-

a Cf. Davidoff on the ventral fins of the Cyprinoids; see above p. 718. These bones appear in a cartilaginous form in several other Physostoms.
6 See Hyrtl, Uber die accessorischen Kiemenorgane der Clupeaceen, Denkschr. Akad. Wiss. Wien, Math. Naturw. Classe, Bd X
(1885), p. 47.

c This spreading of the bone may be seen in the common Herring, though the accessory branchial cavity is not present.
d See Hyktl, 1. c., p. 51.

e See Heckel und Kner, Susswasserfische der Ostreichisch'en Monarchic, pp. 230, 231.

Scandinavian Fishes.

120

952

SCANDINAVIAN FISHES.

sionally so long that it joins the caudal fin, as in the
Sheatfish.

The Herring family is rich in forms. Even after
the sifting to which they have been subjected by Valen-
ciennes", Gunther6, and Bleeker0, there remain about
200 named species, a large number of which have only
a dubious claim to that rank. The family consists
mainly of salt-water fishes, but many species are ana-
dromons, and some are true fresh-water forms. It is
in the temperate and frigid seas of the Northern Hemi-
sphere that the family has attracted most attention and
played the most important part in the economy of man;
but gregariousness is a trait common to the whole fa-
mily and shared by its members in the Southern He-
misphere. Great shoals of certain species d rove in the
South Pacific, along the coasts of New Zealand and
Australia, as the common Herring does in our latitudes.
But owing to the less advanced development of indus-
trial enterprise in those southern regions the fisheries
have not N et attained such dimensions as in the earlier
seats of civilization.

To the ancient Greeks and Romans the Clupeoid
family was by no means unknown, though the Herring
itself does not occur in the Mediterranean ; and even in
the time of Aristophanes they had learnt to cure their
take of Herrings (principally Anchovies and Sardines)
by salting. Halec originally meant brine. Which spe-
cies they designated by their names, it is, however,
difficult to determine. Among the modern Greeks Tlirissa
is the name applied to the Mediterranean Clupea ( Alosa )
anrita. Chalcis (in Aristotle a fish that roved in shoals)
was, according to Callimachus, the same species as
Tricliis , and Aristotle gives c Membras, Trichisf, and
Trichias as different ages of the same species, probably
the Shad. The Anchovy was known by Aristotle as
Enkraulos, by /Elian as Engraulis, Enkrasicholus, and
Lycostomus. Clupea was a classical Latin name. Alosa
( alausa , the Germanic Alse ) occurs first in Ausonius.

Of the four subfamilies under which the Clupeoids,
after the above-mentioned reductions in the number of

the species, naturally fall, only two are represented in
the Scandinavian fauna, by the following species of
more or less common occurrence:

I: Length of the upper jaw from the
tip of the snout less than 3/4 of
that of the head ; intermaxillaries
shorter than the maxillaries5'; ven-
tral margin carinated7' — Subfa-
mily Clupeince.

A: Postorbital length of the head
less than x/4 of the distance be-
tween the dorsal fin and the tip
of the snout. —Subgenus Clupea.
a: Length of the base of the
anal fin less than x/4 of the
distance between the ventral
fins and the tip of the snout Clupea harengus.
b: Length of the base of the
anal fin more than x/4 of the
distance between the ventral
fins and the tip of the snout Clupea sprattus.

B: Postorbital length of the head4
more than x/4 of the distance
between the dorsal fin and the
tip of the snout. — Subgenus
Alosa.

a: Least depth of the tail
more than 4/5 of the length

of the maxillaries __ Clupea pilchardus.

b: Least depth of the tail less
than V- of the leno-th of

the maxillaries Clupea alosa.

a: Gill-rakers on the first
branchial arch about 40 —

60: var. Clupea finta.

(3: Gill-rakers on the first
branchial arch about 60 —

120: var. Clupea alosa.

II: Length of the upper jaw from the
tip of the snout more than 3/4 of
the length of the head. — Sub-
family Engraulince Stolephorus encrasicholus.

Several indications, as for example the more or less
advanced position of the ventral fins and the gradual
prolongation of the anal fin, suggest that in the Clupe-
oid family the course of development has been from
the small-mouthecl Herrings to the large-mouthed An-
chovies. But our materials are not sufficient to enable
us to pass a decisive opinion on this point.

a Cuv., Val., Hist. Nat. Poiss., vol. XIX.
b Cat. Brit. Mus ., Fish., vol. VII.

c Atlas Ichtlnjologique des Indes Orientates Ne'erlandaises, tome VI.
d Clupea sagax , Clupea sundaica, Spratelloides delicatulas.
e De Anim. Hist., lib. VI, cap. 15.

f From hair , an allusion to the hair-like bones in the flesh.

g Intermaxillaries longer than the maxillaries in the subfamily Dorosomatince ( Cliatoessince ).
h Ventral margin not carinated in the subfamily Dussumieriince.

1 At all events when measured obliquely from the middle of the posterior orbital margin to the lower posterior angle of the operculum.

HERRINGS.

953

Genus CLUPEA.

Mouth of moderate size ( in comparison ivith that of the Anchovies small), the length of the maxillaries being less than
half (in Scandinavian forms 40 — 48 %) of that of the head. Tip of the lower jaw projecting to a greater or less
extent beyond that of the snout. Dentition of the mouth feeble, more or less rudimentary. Anal fin not much
longer ( less than twice as long) or even shorter than the dorsal; its rays less than 30.

The greater number of the Clupeoids belong to the
genus of the true Herrings; and the number of species"
contained therein is great enough to have occasioned
several attempts to divide it into subgenera. Valen-
ciennes even distributed its members among several
genera6, which he based principally on the differences
in the dentition. But Nilsson arrived long ago “at the
conclusion c that the teeth are highly variable in dif-

ferent specimens, even of the same species, the bones
which are furnished with teeth in one specimen being
toothless in another”. But the Shads — whose dentition
is as a rule the feeblest — differ so greatly in other re-
spects as well from the remaining Herrings that they
may with reason be placed in a separate subgenus
bearing the name of Alosa.

Subgenus CLUPEA.

Body without spots. Lower anterior portion of the outer surface of the operculum smooth (without grooves).
Dorsal fin set so far bach that the distance between it and the tip of the snout is more than 4 times the post-
orbital length of the head. No large scales of singular form on the caudal fin.

Of this subgenus the Scandinavian fauna possesses
only two species, the Herring and the Sprat, both of
great economical importance, and so like each other
that only a practised eye can distinguish between them,
where they occur, as they often do, in company. In
modern times the most painstaking student of their va-
rieties has been the eminent German naturalist, Pro-
fessor F. Heincke, who has come to the conclusion
that the difference between them is only gradual, in the

a About 90 species have been described.
h Cuv., Val., Hist. Nat. Poiss., vol. XX, pp. 14, cett.
c SJcancl. Fna , Fisk., p. 489.

d Die Varietaten des Herings , II, p. 56 (VII Ber. Comm.

sense that the Herring exhibits races differing from one
another in the same manner as, but to a less degree
than, the two species. We have above seen the same
relation obtain between the Salmons and Charrs, and
in the present instance too we find that the smaller
species attains its place in the form-series of the genus
more rapidly (at a smaller relative size) and more fully
(with more prominent characters). The larger of the
two species is the well-known fish,

rs. deutsch. M.).

954

SCANDINAVIAN FISHES.

THE HERRING (sw. sillen.)

CLUPEA HARENGUS.

Plates XLIII, fig. 1 and XLIV, fig. 1.

Length of the base of the anal fin less than 1/i of the distance between the ventral fins and the tip of the snouta.
Number of spines at the ventral margin ( between the isthmus and the vent ) 36 — 48 b.

R. hr. 8 ; D.

3—5
(13)14—16(17)

(18—200; A.

2—3(4)
(12)13—16(17)

(16— 18d);

P. — liT„; V. 7J_8-p C..v + 1 + 17 + 1 +*; L.lat. (56)60-66(70);
L. tr. 15 — 16; Vert. 51 — 58?.

Syn. Hareng (Chalcidis .species) p. p., Belon., Nat., Divers. Poiss.,
p. 168; Harengus , Rondel., De Pise., lib. VII, cap. XVI;
Gesn., Hist. Anim., lib. IV, p. 408; Schonev., Iclithyol.
Slesv. Holsat., p. 36; Neucrantz, De Harengo, Liibecas
1654; Willugtib. , Hist. Pise., p. 219, tab. P, 1, fig. 2.
Clupea maxilla inferiore longiore, maculis nigris carens, Art.,
Iclithyol., Gen., p. 7; Syn. p. 14; Spec., p. 31; Lin., Fna
Suec., ed. I, png. 120; Gissl., Vet.-Akad. Handl. 1748, p.
107. Duiiam., Tr. Gen. Peches, part. II, sect. Ill, p. 335.
Clupea Harengus, Lin., Syst. Nat., ed. X, tom. I, p. 317;
Penn., Brit. Zool., tom. Ill, p. 294, tab. LXVIII, N:r 160
ed. London, (1776) ( + White-bait., p. 325, tab. LXIX, N:r
176, quam speciem Cyprini generis censuit) ; Fabr., Fna
Groenl ., p. 182; Bl., Fisch. Deutschl., part. I, p. 186,
tab. XXIX, fig. 1; Retz., Fna Suec. Lin., p. 352; Lacep.,
Hist. Nat. Poiss., tom. V, p. 427; Sw., Palmstr., Sv. Zool..
Bd. I, N:r 22; Pall., Zoogr. Ross. Asiat., tom. Ill, p.
209; Nilss., Prodr. Iclitli. Scand., p. 23; Ekstr., Vet.-
Akad. Handl. 1834, p. 20; Ofvers. Vet.-Akad. Fork. 1844,
pp. 82 et 119; Yarr., Brit. Fish., ed. I, vol. II, p. 110
(+ Clupea Leachii, p. 117 + Cl. alba, p. 126); Parn.,
Mem. Wern. Soc., vol. VII, p. 315, tab. XXXV; Cuv.,
Val., Hist. Nat. Poiss., vol. XX, p. 30, tab. 591 (+ Clupea
Leachii, ex Yarr., p. 243 + Clupea elongafa, ex Lesdeur,
p. 247 + Clupea Pallasii , p. 253 + Rogenia alba, ex
Yarr., p. 341); Kr., Damn. Fish., vol. Ill, p. 138; Nilss.,
Slcand. Fna, Fisk., p. 491; Sundev., Stockh. L. Hush.
Sallsk. Handl. 1855, pp. 81, 96, 187; Mgrn , Finl. Fiskfna,
p. 67 ; Lindstr., Gotl. Fisk., Gotl. L. Hush. Sallsk. Arsber.
1866, p. 19 (sep.); v. Bemm. in Herkl., Bouwst. Fna Ne-

derl., part. Ill, p. 380; Gthr, Cat. Brit. Mus., Fish., vol.
VII, p. 415; Boeck, Silden, Sildef., I, Chrnia 1871; Sars,
Jndber. Dep. Indre, 1872, p. 5; Coll., Forh. Vid. S'elsk.
Chrnia 1874, TiUaigsh., p. 185; 1879, N:r 1, p. 97; N.
Mag. Nature. Chrnia, Bd. 29 (1884), p. Ill; Malm, Gbgs
Boh. Fna, p. 570; Lundb., Bih. Vet.-Akad. Handl., Bd. 3,
No. 4; IIcke, Var. Her., Kiel. Commiss., IV — VI Jahrg.,

p. 37; VII Jahrg., p. 59; Winth., Naturh. Tidskr. Kbhvn,

ser. 3, vol. XII, p. 47 ; Bncke, Fisch., Fischer., Fischz.

O., W. Preuss., p. 169; Mor., Hist. Nat. Poiss. Fr., tom.

Ill, p. 443; Smitt, Arch. Biol., vol. Ill (1882), p. 259,

tab. XI; Mela, Vert. Fenn., p. 353, tab. X; Jord., Gilb.,
Bull. U. S. Nat. Mus., No. 16, p. 265 (+ Clupea mirabilis,
ex Gir., ibid.); Day, Fish. Gt. Brit., Irel., vol. II, p. 208,
tab. CXXXVIII, fig. 2; Mob., Hcke, Fisch. Osts., p. 135;
Br. -Goode, Fisher., Fish. Industr. U. S., Sect. I, p. 549,
tab. 204 ( + Jord., Cl. mirabilis, p. 568); Smitt, Bih. Vet.-
Akad. Handl., Bd. 14, Afd. IV, No. 12 (1888); Lillj., Sv.,
Norg. Fisk., vol. Ill, p. 45; Ljgn, Nord. Familjebok, Bd.
14, col. 1079.

Obs. Pallas (1. c.) united the Atlantic and Pacific forms into
one species. Girard (Proc. Acad. Nat. Sc. Philad., 1854, p. 138)
ranged the latter as a distinct species, Clupea mirabilis', but the char-
acters hitherto assigned to it — at most 15 rays in the anal fin
and at most 51 vertebrae — may also be found in the Atlantic
Herring.

The well-known Herring comes nearest both in
form and coloration to the Vendace, but its body com-
monly shows strong compression from the terete back
downwards. But when the Herring is full (i. e. when
the reproductive organs are ripe), the belly is also
comparatively terete, though the downward compression
is always marked at the head and isthmus. The fish
attains a size of 43 cm. or more, but varies consider-
ably in this respect, according to the conditions under
which it lives. The greatest depth, just in front of
the dorsal fin, measures on an average 18 or 19 % of
the length from the tip of the lower jaw to the ex-
treme end of the caudal fin, but sometimes rises, in
gravid females, to 21 % thereof, and, on the other
hand, in young Herrings, even at a length of 125 mm.,
measured as above, sometimes sinks to only 15 % there-

° Here, as in the Gwyniads, we have reckoned the tip of the snout from the articular knobs of the maxillaries.
h The limits of variation according to Heincke.
c In exceptional instances 17 and 21.

a In exceptional instances 14 and 19; 15 and 20 according to Heincke.

e In exceptional instances 14.

f In exceptional instances 9.

9 According to Heincke.

HERRING.

955

ofa. The least depth of the body (of the tail) mea-
sures on an average 673 — 672 0//°h °f the said length.
According to the method of measuring the length of
the body employed above, in the case of the Gwyniads,
i. e. from the articular knobs of the inaxillaries to the
end of the middle caudal rays, the least depth of the
body varies between 8 and 7 % of its length. The
variations in our specimens from Bohuslan are such
that the averages regularly decrease from 8' I % to 7'3 %
as the length of the body increases from 1G to 37 cm.
The body as a whole is of the most beautiful fusiform
shape, the dorsal and ventral contours forming regular
and similar curves; but the lateral compression is so
marked that the greatest thickness, which, as we have
mentioned, lies nearer to the back, measures, in the line
of the greatest depth, about half — in young specimens
and after the spawning about 2/5 — of the latter.

The length of the head varies between 22 72 and
1 9 1/2 % of that of the body". It is of the well-known,
three-sided, pyramidal shape that we have already seen
in the Flying-fishes, but with narrower upper surface
and with the snout constricted above, in front of the
eyes. The most characteristic points are the prominent
tip of the lower jaw and the coat of adipose membrane
concealing a great portion of the eyes and extending
both over the sides of the snout and the upper part
of the shoulder-girdle. This membrane forms two folds
in front, one composing the anterior limit of the
orbit, and another the anterior eyelid, which passes
above without a break into the posterior, growing
narrower like the latter, but is somewhat broader be-
low, lying in a fold outside the lower, pointed corner
of the posterior eyelid, which gradually passes behind
and below into the adipose membrane of the opercular
apparatus and the cheeks (the suborbital ring). In this
latter membrane, which is continuous both with the
covering of the sides of the snout (the preorbital bones)
and with the skin of the forehead and occiput, we find
an abundance of the ordinary ramifications of the lateral
line, which also occur on the shoulders, though every-
where less developed than Ave shall find them in the
Shads. The top of the head, from the nostrils right

to the occipital crest, as far as the frontal bones extend,
is coursed on each side by a well-marked lateral ridge,
bounding a depressed, lanceolate area, dotted with rnu-
ciferous pores, the tip of the area lying at the occipital
crest itself. But at the middle of this area runs an
elevation, in front terete or even mesially depressed,
behind sharper, Avhicli is formed anteriorly by the above-
mentioned ridge on the ethmoidal cartilage, and dis-
appears posteriorly, behind the int-erorbital space. The
eyes are round, or slightly longer than high, but the
part free of the eyelids is elliptical, with the upper and
loAver ends pointed. They are rather large, their longi-
tudinal diameter varying on an average betAveen 74 and
7 5 of the length of the head, and being about equal
in adult specimens to the length of the snout from the
middle of its tip to the anterior corner of the eye, but
measuring only about half the postorbital length of the
head. They are set so high that their upper margin
approaches close to the frontal plane. The nostrils lie
on a level Avith the upper margin of the pupil and
about half as far from the anterior corner of the eye
as from the middle of the tip of the snout. The an-
terior nostril of each pair is round and distinct, though
comparatively small; the posterior more crack-like and
usually overhung by the adipose membrane belonging
to the sides of the snout. The two pairs are separated
by a distance measuring on an average about 2/5 of
the breadth of the snout at the articulations of the
inaxillaries. The sharp tip of the snout is formed by
the thin-lipped intermaxillaries, Avhicli strongly remind
us, both in position and form, of those of the Vendace.
They are flat but someAvhat arched, triangular, Avith
the posterior (outer) corner elongated to a point and
armed on the inner surface of the inferior margin Avith
a toav of about 6 pointed, incurved (recurved) teeth.
They are Avithout nasal processes — the tip of the snout
being consequently not protrusile — and their anterior
loA\:er corners are rounded to such a degree that they
are distinctly parted from each other at the middle of
the tip of the snout. But the sinus thus caused is filled
by the labial skin, the tip of the snout being thus en-
tire and truncate, Avithout indentation or groove. In

“ In the largest Herrings we often find retrogressions to the characters of youlh. In a full female, with the length of the body
measuring 42 cm. and with eggs 0’8 — 1 mm. in diameter, taken in February, 1887, off Helso (Northern Bohuslan), the greatest depth is
only 1 6 ' 1 % of the said length.

6 Varying, according to Lilljeborg’s measurements, between 5 and 7 f.
c According to our ordinary method of measurement.

956

SCANDINAVIAN FISHES.

the maxillaries we have to distinguish, as among the
Vendaces, between the more terete, introrse, articular
shaft, which supports the intermaxillaries on its up-
turned, grooved, and proximally broader, front surface,
and the lateral part, set almost at right angles to the
former, and indeed terete in front, but soon expanding
into the flat, sabre-shaped disk that forms the exter-
nally visible maxillary. This part is obtusely rounded
behind and toothed with spines at the thin lower mar-
gin, and by the addition of the supplementary bone is
rendered fairly uniform in breadth throughout the
greater part of its length. The posterior supplementary
(jugal) bone is elliptical, and in front pointed, the an-
terior narrower and of uniform breadth. The length
of the lateral maxillary disk, measured from the arti-
cular knob, varies on an average between 10 and 872 %
of that of the body or between 47 and 4472 % °f that
of the head, and the bone extends back to a line with
the anterior margin of the pupil or even with its centre,
though always so that the length of the cheek (between
the maxillary and the vertical anterior margin of the
preoperculum) is greater than its height exactly below
the eye. The cheek is covered by the large, third
suborbital bone, which also forms the greater part of
the posterior limit of the orbit, the fourth (hindmost)
suborbital bone being comparatively short. The lower
jaw is characterized by the rather great height of the
dental part — about twice the breadth of the maxillaries
or 75 °f the length of the lower jaw — and by the ter-
ete ridges on the outside of this part, which diverge
backwards from each side of the chin, one of them run-
ning along the upper margin, the other parallel to the
lower margin. The symphysis is also fairly high, and
is furnished below with a small, prominent chin-protu-
berance, but channelled underneath at its angle. Only
at the extreme front of the upper margin do we find
an arcuate row of small teeth (6 on each side of the
symphysis), in well-marked, but closely approximated
sockets. The length of the lower jaw varies between 13
and 12 % (in the oldest specimens 11 %) of that of the
body, or 60 and 57 % (exceptionally 61 % or, in the
oldest specimens 55 %) of that of the head, and it is
greater, as a rule, than the length of the base of the
anal fin, though in exceptional cases this relation too
is reversed. On the symphysis and the sides of the
lower jaw hangs a fairly broad labial fold. Internally
the mouth is furnished with the ordinary transverse
folds (vela), the lower being the more developed. The

vomer has in front a longitudinal, lanceolate elevation,
bearing a double row of small, pointed teeth, somewhat
larger, however, than the jaw-teeth ; and a shorter row
of smaller teeth usually occupies the extreme front of
each palatine. The tongue is free and fleshy, but thin-
margined. On the glosso-hyoid bone is an almond-
shaped card of small teeth. The gill-rakers are set in
a single row on most of the branchial arches, long and
setiform, with numerous small spines on their sides.
On the first branchial arch there are about 70, as a
rule 45 — 47 (exceptionally 44 or 50) on the lower part
thereof, and the longest (at the middle of the arch) are
about equal in length to the suboperculum at its suture
with the operculum. Only on the upper part of the
third arch and on the fourth arch as far as it forms
the anterior margin of the short, hindmost gill-slit, do
we find an inner (posterior) row of short, lamellate
rakers. About 20 spines, similar in form to the latter,
occupy the outside of the lower pharyngeals, which re-
semble branchial arches, and are furnished with only
one row of small teeth, whereas the upper pharyngeals
bear a triangular card of similar teeth. The outer gill-
openings are large, extending above the upper margin
of the operculum, from a level with the superior mar-
gin of the eye, down to a line with the anterior margin
of the latter. The operculum is obliquely quadrilateral,
with the upper posterior corner rounded and the lower
anterior corner prolongated to an acute angle, so that
the faint S-shape of the subopercular suture is directed
obliquely backwards and upwards. The length of this
suture varies between 23 (exceptionally 2 1 1/2) and 26
(exceptionally 27) % of that of the head. On the sur-
face of the operculum we can distinguish three slightly
convex patches, separated by two shallow grooves, one
running obliquely backwards and upwards, the other
obliquely backwards and downwards, from the articular
head of the operculum. But these patches are smooth
or marked only with faint concentric s trial, and the an-
terior lower patch in particular is destitute of radiating
grooves or ridges. The suboperculum resembles a nar-
row, eccentric sector of a circle, varying in breadth;
sometimes its margin shows an irregular, shallow in-
dentation. The preoperculum is obtuse at the inner
angle, but the lower posterior corner forms a rounded
right angle. Its posterior disk is so broad that its length
(the lower arm together with the disk) is about equal
to its height (the vertical arm together 'with the disk).
The interoperculum is triangular and equal in length

HERRING.

957

to the preoperculum. Its externally visible part (below
the preoperculum) is, however, sword-shaped and of
uniform breadth. The hind margins of the preoperculum
and interoperculum together form a slightly curved line,
continued below the latter in a contour forming an
obtuse angle with the inferior margin of the suboper-
culum. In the Herring, as in the other Clupeoids,
we find a bone that seems at first sight to be a lower
suboperculum, forming an angular break at the lower
posterior angle of the opercular apparatus. In order
that the apparatus may be more firmly closed at this
angle, the anterior margin of the shoulder-girdle rises
at this point in the form of an elongated, sharp-edged
knob, the substratum of which consists of an upright
process {dp, fig. 238, p. 950) from the upper margin
of the horizontal arm of the clavicle. On closer exa-
mination, however, it soon appears that the apparent
lower suboperculum is really tire hindmost of the three
broad branchiostegal rays set on the epihyoid bone.
The first five rays, which are situated on the cerato-
hyoid bone, are subulate and more curved. The branchio-
stegal membranes are free both from each other and the
isthmus, and overlap each other only to a very small
extent in the extreme front, the left membrane lying
outside the right.

The dorsal fin, which is trapezoidal in shape, begins
at a distance from the tip of the snout (the articular
heads of the maxillaries) measuring 47 — 51 % of the
length of the body. The length of its base is 13 — 10 %,
and its greatest height (the length of the first or second
branched ray) 12 — 10 % (less frequently 9 %), of the
length of the body. The anal fin is of a similar shape
as the dorsal but much lower. The distance between
its beginning and the tip of the snout is 71 — 75 %, the
length of its base 11 — 10 % (exceptionally 12 or 9 %),
and its greatest height 5V2 — 4V2 % (exceptionally 6 or
4 %), of the length of the body. The caudal fin is
deeply forked, the lower lobe being somewhat longer
and more pointed than the upper. Its middle rays
occupy about 5 — 4 % of the length of the body, and
measure only V3 — 1/i of the length of the longest rays
in the fin.

The pectoral fins are set low, as in the Salmons,
and of an oval, obliquely pointed shape, the second
branched (third) ray being usually the longest. Their
length varies between 16“ and 13 % of that of the body,

and during youth is generally somewhat more, in old
specimens perceptibly less, than half the preabdominal
length, which increases even relatively with age, from
30 to 35 or nearly 36 % of the length of the body.
The ventral fins are broader and shorter, but triangular.
Their length varies between 10 and 8 % of that of the
body. Their insertion always lies some distance, though
in young specimens this is sometimes very small, behind
the perpendicular from the beginning of the dorsal fin,
and at a distance from the tip of the mouth measuring
51 b — 55 % of the length of the body. The postabdo-
minal length is about 2/s of the preabdominal, and
varies between about 20 and 23 % (exceptionally 24 %)
of the length of the body.

The thin and deciduous scales (fig. 239) are round
or quadrilateral with rounded angles. The hind (free)
part is indented at the margin with small, rounded
sinuses, and shows only faint traces of radiating grooves.
The anterior part (covered and about twice as large) is

Fig. 239. Scale from the left side of a Clupea liarengus
3 dm. long. X 3.

smooth-margined and marked with extremely fine and
numerous, irregularly undulating and confluent, trans-
verse striae, without radiating grooves or with only ru-
diments thereof, in the form of a few streaks (5 — 7
upwards and downwards) obliquely crossing the others.
In this striature the growth-rings of the scale also
appear as concentric lines. The lateral line does not
pierce any scales — only in exceptional instances are
there a few perforated scales in front — but under the
scales it runs straight from the upper clavicle to the
middle of the base of the caudal fin. The singular
marginal scales of the belly (fig. 240) are of an oblique
diamond-shape, with a strengthening ridge in the me-
dian line of the belly, ending in the backward spine,
and another on each side, continued by the upward
process. They are remarkable in the Herring for their
number, which is greater than in the following species,
being as a rule, according to Heincke, 27 — 30 (excep-

a In young Herrings sometimes 17.
b In the fry sometimes 5 0 1 /2 .

958

SCANDINAVIAN FISHES.

tionally 24 — 32) from the isthmus to the ventral tins,
and 13 — 15 (exceptionally 11 — 20) between these tins
and the vent, in all 36 — 48. Just in front of the ven-
tral tins lies a double plate, the upward processes being
bifid, or shoAving indications that they Avere originally
composed of as many as three processes. The other
preabdominal plates, on the other hand, Avith the ex-
ception of the foremost ones, are distinguished from those
of the folloAving species by their long lateral processes.

The colour of the back is steel-blue Avith a lustre
of green and gold; the sides are silvery. The limit be-
tween the colours of the back and belly is often marked
by a tinge of lustrous green. The neAv-caught Herring
glitters with a Avealth of hues, and in the darkness it
gleams Avith phosphorescent light. The snout and the
upper parts of the head, as Avell as the superior part of
the opercular margin, are sometimes pure black. The

Fig. 240. Spiniferous scales from the ventral margin of a Vlupea
harengus 25 cm. long. Twice the natural size.
a, the foremost spiniferous scale; b, the fifth, counting forward from
the ventral fins; c, the double scale just in front of the ventral fins;
d, the fifth scale, counting forward from the vent; e, the hindmost
spiniferous scale.

gill-covers are of a golden lustre, more and more suf-
fused Avith blood, the longer the fish has been dead.
The ventral and anal tins are Avhite but transparent, as
Avell as the pectorals; but the upper margin of the last-
mentioned tins is of a grayish or greenish black, which
colour extends, in greater or less intensity, some Avay
doAvn the fins. The dorsal and caudal tins are darker,
though the membrane is transparent, more densely speck-

led Avith grayish black, shading on the rays into green.
The iris is sometimes silvery Avith a brassy lustre; in
other cases the latter colour is predominant.

With coloration of a lighter or darker tone, and Avith
body more or less transparent, though this transparency
is ahvays faint, the Herring varies according to the
greater or less intensity of light in its surroundings.

In the temperament of the whole Herring family
gregariousness and a love of migration are the most pro-
minent traits. The common Herring, an inhabitant of
the North Atlantic and probably, as we have mentioned
above, of the corresponding regions in the Pacific, roves
in huge shoals Avithin the said geographical range. Hoav
far its wanderings extend, is still an undecided question,
though numerous investigators, from the beginning of
the last century until iioav, have devoted their attention
to its solution. In an Atlas maritimus commercialism
printed in London in 1728, reference is made to the
annual course of the Herring-fishery off the coasts of
Great Britain and Ireland, then essentially the same as
it is to the present day. The most important fishery
in those Avaters begins in summer to the extreme north,
off the Shetland Islands and then on the Scotch coast,
but afterAvards advances further and further south on
both sides of Great Britain, to England, Ireland, and
the Channel. Hence it Avas inferred that the Herring-
shoals must come from the far North; they Avere sup-
posed at first to be so large and densely packed that
they had difficulty in forcing a passage “betAveen the
coasts of Greenland and North Cape”; but the Herring
“army” divided as it came further south. On these as-
sumptions the learned and clever Burgomaster of Ham-
burg, Johan Andersson, d. 1743“, based his theory of
the annual migrations of the Herring. The Herring, he
opined, had its true home under the ice of the Arctic
Circle. There lay its cradle, and thither it retreated
year after year to escape its merciless pursuers, and to
recruit its thinned numbers. But soon the home became
overcroAvded; large multitudes Avere yearly compelled to
emigrate southwards, and to divide into separate armies,
Avhich marched partly to the right, to America, and
partly to the left, to the Avest coast of Europe and into
the Baltic. This theory Avas still accepted in the days
of Pennant6; but Bloch6' and Noel'6 shoAved it to be

a See his Nachrichten von Island , Gronland und tier Strasse Davis, Hamburg 1746, pp. 51 — 72.
b Brit. Zoology , 17 76.
c Fische Deutschl., part. 1, p. 189.
d Magazin Encyciopedique, tome VI, p. 5 (Paris 1795).

HERRING.

959

untenable, the latter stating that, in his opinion, “each
marine region as a rule has its native Herring, which
is easily distinguished from other forms by its migrat-
ing at a different time of year, when it forsakes its first
home in the depths, and approaches land to deposit its
roe in more suitable localities”. Mac Culloch'1 pur-
sued the criticism of Andersson’s theory, and in direct
opposition to the regularity assumed by Andersson, he
ranged the unaccountable variations in the Herring’s
approach to the coast. It was hence that Yar-
eell b drew his well-known and often misused expres-
sion, “the Herring is a most capricious fish.” But Mac
Culloch also endeavoured to find an explanation of
this fickleness in the different motives that induce the
Herring to migrate from deep water to the surface and
the shallows. Among the causes hereof he adduces the
desire of spawning, a point already touched upon by
Noel, the search for food, consisting of animals known
in his time as “medusEe and other analogous marine
vermes, which are produced in such abundance during
the summer, in all the shallow seas,” and the fear of
enemies. In Sweden Nilsson has been the most emi-
nent exponent of this theory, and he attributed thereto
a great economical importance, for he based on it his
assertion that a Herring-fishery may be utterly de-
stroyed, or at all events ruined for many years, by
the manner in which it is carried on. The Bohusl&n
fishery was in his opinion a case in point; but this
conclusion has been opposed by many.

The pith of the question evidently lies in the elu-
cidation by natural history of the varieties of the Her-
ring, if such varieties be really in existence. Nilsson
originally6 adopted eight constant, local varieties:

1) The Ocean Herring (Forma oceanica ): Rabo''
Herring (var. oeresundica), Kulla6 Herring (var.
schelderensis), B oh u si an Grass-Herring (var.

majalis), Bohuslan Breeding-Herring or Great-
Herring (var. bahusica), Norwegian Winter-
Herring (var. hiemalis), Norwegian Autumn-
Herring (var. autumnalis ) ;

2) The Shore Herring (Forma tceniensis ): KivilG
Herring (var. cimbrica), Baltic Herring or
“Stromming” (var. menibras).

Subsequently 9 he reduced the number of these
varieties to six, by uniting the Rabo Herring with the
Kulla Herring and the Bohuslan Grass-Herring with the
Great Herring. The names given above state where
the varieties were supposed to occur, and the distinctive
characters were derived from the differences in the size
of the head and eyes, in the depth of the body, in the
position of the fins, and in the number of marginal scales
on the belly. Merely a hasty observation is enough to
show that these characters hold good in the main, or,
to employ Nilsson’s words, when the Herring appears
in large numbers together. And yet they are only signs
of different developmental stages. It is but natural,
however, that among innumerable individuals, as when
the Herring appears in multitudes sometimes so densely
packed within inlets of the sea that the name of moun-
tains has been conferred upon them, individual diffe-
rences should be comparatively easy to trace. But if
we eliminate these individual variations by calculating
a sufficient number of averages from an adequate num-
ber of individuals, we can make remarkable discoveries.

Among the factors necessary to an estimation of the
significance of form-characters, or to a determination of
the natural relations obtaining between different varie-
ties or species, the differences of sex and the alterations
of growth occupy the foremost rank. In order to find
expressions for the former, I have caused the averages
to be calculated for 75 Baltic Herrings, measured by
Air. Lundberg, Inspector of Fisheries'1.

“ Quart. Journ. Sc., Lit., Arts, vol. XVI, p. 210, London 1823 — 24.
b Hist. Brit. Fish., ed. 1, vol. II, p. 112.
c Prodromns Ichthyologice Scandinavian , p. 23.

d Raa is a large fishing- village on the Sound, south of Helsingborg.
e Kullen is a promontory north of Helsingborg, south of Schelder Bay.
f A large fishing-village near Kristianstad.
g Skandinavisk Fauna, Fiskarne, p. 493.
h Bih. Vet. Akad. Handl., Bd. 3, No. 4.

Scandinavian Fishes.

121

960

SCANDINAVIAN FISHES.

Averages in the Baltic Herring {Stromming ; Clupea harengus, var. membras ).

c?

9

Number of specimens measured

Length of the body from the tip of the lower jaw to the end of the caudal lobes, expressed in millimetres

35

212.4

40

219.9

Distance between the dorsal fin and the tip of the lower jaw .

„ „ „ ventral fins ,, „ „ „ ,, „ „ .

„ „ „ pectoral fins „ „ „ „ „ „ „ .

Depth of the body behind the tips of the pectoral fins

Thickness ,, ,, ,, ,, ,, ,» - , ,, ,, ,,

Depth „ „ ,, ,, „ „ „ ,, ventral „

Length of the head

Horizontal diameter of the eye

Vertical „ „ „ „

Interorbital width ...

Length of the snout

,, „ upper jaw

„ „ „ lower

„ „ „ base of the dorsal fin

„ „ ,, pectoral fins

„ „ „ ventral „

„ ,, „ base of the anal fin

% of the length of the body

77 77 77 77

77 77 77 77

7? 77 77 77

77 77

77 77

77 77 77

77 77 77

77 77 77 77

44.75

44.67

48.71

48.7 9

20.64

20.71

18.2 7

17.93

9.12

8.90

15.20

15.27

20.81

20.73

5.56

5.42

5.09

5.06

3.54

3.52

7.15

7.21

8.69

8.69

11.25

11.15

10.60

10.71

13.02

13.03

8.39

8.32

9.53

9.56

In vain do we seek here for any noticeable dif-
ference between the sexes. All the percentages are the
same in both columns, except in two instances. The
depth of the body behind the tips of the pectoral tins
and its thickness at the same point are somewhat greater
in the males than in the females. This difference, how-
ever insignificant it may appear, is of economical im-
portance, and had in Nilsson’s opinion a zoological
importance as well. Greater depth and thickness com-

bined involve greater fleshiness and greater value even
to the epicure. The most important difference adduced
by Nilsson between the Norwegian Summer (Autumn
or Fat) Herring and the Norwegian Spring (Gray-boned")
Herring, also consisted in the relative depth and thick-
ness. The same difference appears too between the so-
called Autumn Stromming and Spring Stromming, on a
calculation of the averages for these two forms, as far
as they have been distinguished, in Lundberg’s tables:

Averages in

Spring

Stromming.

Autumn

Stromming.

Number of specimens measured

23

29

Length of the bodv expressed in millimetres ■.

210.9

215.5

Depth of the body behind the tips of the pectoral fins in % of its length

17.9

17.7

thickness ,, ,, ,, ,, ,, ?? 77 77 7? 77 77 77 77 77

9.0

8.8

The difference is indeed slight; but here as in the
preceding table, its course is opposite to that of the
changes of growth in the Herring, and it is therefore
not without importance. The Autumn Stromming too
generally finds a better market than the Spring Strom-
ming. Nilsson remarked this difference between the
Kivilt Herring and his Stromming .

Exactly the same distinction may be traced, how-
ever, between different takes of the same Herring-

shoal, netted at the same time, in the one instance
farther from land and in deeper water, in the other
nearer shore and in the shallows. This I have shown
in my report to the Swedish Home Secretary on the
experiments in drift-net fishing carried out under my
supervision in Bolnislan during the winter of 1880 —
81. From the measurements taken by Mr. Trybom,
Assistant Inspector of Fisheries, I calculated the follow-
ing averages.

The word in the original ( grabenssill ) really means, according

to Ljungman, grofbenssill , i. e. coarse-boned Herring.

HERRING.

961

Drift-net take, Dec. 20th, 1880, in Koster Fjord

Seine take, Dec. 21st, 1880, ,, „ „

Drift-net take, Jan. 3rd, 1881, „ „ „

,, „ Jan. 5th, ,, „ the Skager Rack, off Koster

Seine take, „ „ „ Koster Fjord

,, „ Dec. 13th, 1880, at Ottero, off Grebbestad

Number of
specimens
measured.

A

v e r a g e

S.

Length of the
body expressed
in mm.

Greatest depth
of the body in
% of the length
of the body.

Greatest cir-
cumference of
the body, in %
of the length
of the body.

6

314

20

46

5

280

17

41

5

315

18

45

5

299

19

44

5

259

17

39

12

284

17

42

Here it appears distinctly that the drift-net takes
(from deeper water) and the seine takes (made close
in shore) differ from each other in the same way as
the Fat Herring from the Gray-boned Herring and
the Kivik Herring from the Stromming.

It is the difference between the Ocean Herring
( oceansillen ). and the Baltic Herring (stromming en),

however, that has been set forth most clearly and
most reasonably, ever since Linnaeus adopted the
latter as a separate variety. The relations that obtain
between these forms, I have endeavoured to deduce
by another method from measurements and tables of
averages'*.

Averages in Herrings from

Norway.

The Baltic.

Bohusliin.

Scotland.

Norway.

Scotland.

Number of specimens measured

8

6

32

10

5

3

5

5

Len

gth of the body expressed in millimetres

170.4

203.2

240.6

240.9

152.2

200.7

221

260.8

1.

Length of the head in % of the length

of the body

21.7

20.6

20.5

20.3

21.9

21.3

20.3

20.2

2.

,, ,, ,, maxillaries... ,, ., ,, ,,

„

„

10.2

9.3

9.2

9.2-

10.3

lO.o

9.3

9.i

3.'

„ „ „ lower jaw „ „ „ „ „

„

55

13.o

12.i

12.i

12.o

13.2

12.8

12.i

11.9

4.

Height of the dorsal fin „ „ „ „ „

„

„

11.0

10.3

9.9

9.9

11.5

10.i

lO.i

9.7

5.

Distance between the dorsal fin and the

tip of the snout ,, ,, ., „

5J

?>

48.9

48.8

48.8

49.7

48.5

49.5

48.3

51.0

6.

Length of the pectoral fins.. ,, ,, ,, ,, .,

55

55

15.5

14.8

14.3

14.5

15.8

15.0

14.6

14.4

7.

Preahdominal length _ ,, ,, „ ,, „

55

55

31.2

31.9

32.7

36.o

30.4

32.4

33.4

38.7

8.

Distance between the ventral fins and the

tip of the snout ,, ,, ,, ., ,,

55

52.2

52.2

52.9

53.7

51.5

53.5

53.2

54.2

9.

Length of the ventral fins ,, ,, ., ,, .,

J?

55

,,

9.8

9.3

9.i

8.8

9.9

9.6

8.8

8.8

10.

Postabdominal length... „ „ „ „ - „

„

55

„

20.8

21.9

21.9

22.7

20.6

22.0

22.o

23.3

11.

Distance between the anal fin and the

tip of the snout „ „ „ „ „

?>

55

72.4

73.6

73.7

74.5’

72.1

72.9

73.7

75.2

12.

Height of the anal fin ,, ., ,, ,, ,,

55

5.5

4.9

4.9

4.6

5.6

5.2

4.8

4.4

13.

Length of the middle caudal rays „ „ ,, „ „

55

55

4.9

4.6

4.2

3.6

5.3

4.2

3.9

3.4

14.

„ „ „ pectoral fins in % of the preabdominal length

50. o

46.6

44.o

42.6

51.7

47.2

43.8

41.5

Flere the averages run with very great uniformity,
falling or rising with age; and the form-series thus ar-
rived at in the first four columns is controlled by the
last four columns, which contain the alterations of growth
in the youngest and the oldest among these races, the
former consisting of Fat Herrings ( matjes ) from Sta-
vanger, the latter of Scotch Herrings from Peterhead.
Such an unbroken continuity in the form-series — if we

choose to regard the Herring-races as separate forms —
and such a parallelism in the alterations of growth could
hardly be traced, were there no genetic affinity between
all the above Herring-forms, from the Baltic and the
west coast of Sweden, from Norway and Scotland. Thus,
it appears, we can scarcely find any ground whatever
for the assumption of a form-distinction or even of a
racial difference. We see, however, that in the relations

“ These measurements deviate in two respects from the preceding ones: the dimensions measured from the snout have been taken from
the articular knobs of the maxillaries, and the length of the body has been measured to the end of the middle caudal rays. The percen-
tages are consequently different from those already given, which can have no effect, however, upon the ultimate results.

962

SCANDINAVIAN FISHES.

which show the most marked alterations of growth — 7,
8, 10, and 13 in the table — the oldest Fat Herrings
from Stavanger (Column 6), though smaller on an aver-
age than the Baltic Herrings (Column 2), have advanced
further than the latter. Their development has pro-
ceeded more rapidly. Similarly the youngest Scotch
Herrings (Column 7), though inferior in average size

Averages in Herrings from

Bohuslan.

Scotland.

Bohuslan.

Scotland.

Number of specimens measured

11

5

8

5

Length of the body expressed in millimetres

221

221

263.4

260

Length of the head in % of the length of the body

20.5 ’

20.3

20.3

20. a

Preabdominal length „ „ „ ,, „ „ „ „

32.7

33.4

33.i

33.8

Distance between the anal fin and the tip of the snout „ ,, ,, „ „ „ „ „

72.8

73.7

74.3

75.2

to the Herrings from Bohuslan (Column 3), have out-
stripped the latter in the alterations of growth. The
more oceanic its home, the more rapid and more vi-
gorous is the development of the Herring type. This
is still more clearly shown by comparing with the last
two columns Bohuslan Herrings more nearly approach-
ing to them in a.o;e:

The same results may be deduced from Lundberg’s Ocean Herrings with the organs of generation equally,

tables by thence selecting for comparison Baltic and or at all events not too unequally developed:

Averages in Herrings from

The Baltic.

Bohuslan.

Number of specimens measured

11

42

19

6

7

Length of the body to the end of the caudal lobes, expressed in mm

186.1

205. g

223.3

294.2

247.9

,, „ ,, head in % of the above length of the body

21.4

20.9

20.8

20.3

18.6

Number of specimens measured

11

13

6

7

Length of the body to the end of the caudal lobes, expressed in mm

186.i

201.8

294.2

247.9

Vertical diameter of the eyes in % of the above length of the body

5.5

5.2

4.7

4.5

The Baltic form (the Stromming) thus fails, even
at its most advanced age, to attain averages equal to
those shown by the much smaller Herrings from Bo-
huslan in the last column. The Strommings, as Nils-
son has pointed out, have on an average a larger head
and larger eyes; but this is only a general rule, for
exceptions are not uncommon.

Between the Stromming and the Herring, however,
the averages obtainable from Lundberg’s tables show
another distinction, the significance of which is all the
greater, since it indicates a different direction of deve-
lopment or at least a divergent tendency, and seems
most naturally to admit of explanation, as we have
above stated of the analogous relations in the Salmon-
oids, on the assumption either that the forms are still
in process of differentiation from each other, or that

one of them is undergoing a degeneration expressed by
reversions to the earlier stages of development. A rule
of general validity, which I stated in 1882" in a de-
scription of an hermaphroditic example of this species,
tells us that, after the dorsal fin has passed through the
forward removal which, as Sundevall'" first showed,
attends the development of the Herring from the larval
stage, there ensues, during the subsequent growth, a
backward removal of this fin. In a preceding table
(p. 961) we have also seen that the percentages for
the distance between the dorsal fin and the tip of the
snout (relation no. 5) increase with age. The ventral
fins share in this backward motion (relation no. 8).
Let us now examine these relations in the Stromming,
according to Lundberg’s tables. For this purpose we
divide the specimens measured by him into four groups,

a Arch. Biol., vol. Ill, p. 269. It should, however, be remarked that I was prevented from reading the proofs of this paper, which
was printed at Brussels, and that a number of misprints, especially in the numerals, have consequently been overlooked.

6 Om fislcyngels utveclcling, Vet.-Akad. Handl. 1855, p. 20.

HERRING.

963

the first measuring 67 — 111 mm. in length of body,
the second 136 — 201 mm., the third 203 — 240 mm.,
and the fourth containing all the large ^specimens, be-

tween 242 and 345 mm. in length. The first three
groups roughly answer, according to other calculations
of mine, to the age-classes of the first three years.

Averages in Strommings.

-

Number of specimens measured ...

12

47

47

20

Length of the body to the end of the caudal lobes, expressed in millimetres

86

178.9

218.5

253.5

Distance between the dorsal fin and the tip of the lower jaw in % of the length of the body

43.8

44.7 5

44.07

44.32

?? ventral fins ,7 ,, ,, ,, ,, 77 77 ,, ,, 77 ,, ,, ,, ,, ,,

47.1

48.93

48.58

48.39

After the second year there thus appears, on an
average, by no means a backward removal, but a re-
gular advancement both of the dorsal and the ventral
fins. The normal development of the Stromming, in
accordance with the rules for the Herring-type, termi-
nates in general at- the end of the second year. And
it then shows, as we have seen above (the table on
p. 961, relation no. 8), at an average length of 203
mm., the same position of the ventral fins as the Nor-
wegian Herring does at a length of 170 mm. As re-
gards the position of the dorsal fin (relation no. 5), the
same rule applies to the Bohuslan Herrings.

That a racial difference exists here, seems, in a
certain sense, undeniable. But this difference evidently
agrees, as Petersen" has stated, with the geographical
separation, and Winther’s0 observations go to show
that this separation is sometimes evanescent. “The
entire alteration in the Herring inhabiting the Sound”,
he writes, “may be summed up as follows. The little
Baltic Herring (Kivik Herring, Nilss.), which flocks
together at the beginning of autumn at the south end
of the Sound, advanced in 1867 right to the middle
of the strait, north of the Flint Channel, where it found
the great basin known as the “ Hulkene ’ untenanted by
the Bottom Herring c, which had already departed. After
the spawning it followed the northward current out
of the Sound into the south of the Cattegat, whence
it returned year after year to its old spawning-place,
constantly increasing, under the favourable influence
of more congenial environments (the greater saltness
of the water?), in size and fecundity. It continued to

“improve” in this manner until 1873, when its deve-
lopment had advanced so far that it could no longer
content itself within the confined limits of the Sound,
but as it gradually attained the size and form of the
“ Kulla Herring ”, began to repair to the spawning-
places of the latter variety. Only few returned in 1875
to the old spawning-places in the Sound, where a new
stock of Baltic Herring this year (1875) replaced the
old one in the Hulkene, north of the Flint Channel”.
The Stromming can thus become Kulla Herring, an
alteration which, as we have seen, involves nothing
more than a higher degree of the typical development
of the species. But it by no means follows that all
Strommings become Herrings, nor does it seem possible
for anyone to show that all young Bohuslan Herrings,
loddsillar as they are called, in course of time become
Ocean Herrings.

That a particular stage of development under per-
manent circumstances may become fixed as the termi-
nation of the development of a species under those
circumstances, is nothing unusual, and the characters
of youth, especially in fishes that breed before they have
attained their full specific size and development, may
become hereditary and, under certain conditions, remain
unaltered. This is, no doubt, the explanation both of
the actual difference between the average Stromming and
the average Herring, and of the fact that the young of
the so-called Spring Herring and Coast Herring differ
from the fry of the Autumn Herring and Ocean Her-
ring''. But the difference appears, to the best of our
knowledge, only in the averages and, probably, only

° Kritilc etc., Vid. Meddel. Naturh. For. Kbhvn, 1888, p. 1.
b Nord. Tidskr. Fisk., Aarg. 3 (1876), p. 12.

c The name given by the fishermen to the shoals of larger Herring in the Sound.

d “II faut necessairement admettre la residence de ces poissons sur des fonds differents, ou la diversite de la grandeur et de la gros-
seur constitue autant de varietes on de races qui se perpetuent par voie de generation’ ; Cuv., Val., 1. c., XX, p. 49.

964

SCANDINAVIAN FISHES.

so long as the geographical separation is maintained.
The three Herring varieties adopted by Heincke (1. c.),
which he called A, B , and C , are also distinct expres-
sions of different ages. The development has started
from C, the characters of which, as Heincke says, are
a blending of those of A and B; and the last-mentioned
variety represents that stage of development in which
the dorsal and ventral tins have moved furthest for-
ward, again to recede, in accordance with the above-
mentioned rule for the development, to the position
which they occupy in Heincke’s variety A.

Such is the manner in which the distinctions ad-
duced between different kinds of Herring may be ex-
plained away. Practically expressed, the result is that
no constant character has been discovered for the as-
sumed varieties or races. It must, therefore, be assumed
that the geographical separation is not marked enough
to effect a complete severance of variety or race. The
Herring is a migratory fish, no more restricted to fixed
localities within its range than other such fishes. On
endeavouring to find a centre for that range in the
basin of the Atlantic, we arrive at a conclusion which
in a certain degree recalls Andeksson’s antiquated
theory. A line including the White Sea, the extreme
north of Norway, Iceland, Southern Greenland, New-
foundland, the west coast of Canada and of the North-
ern States, and further south on the European side
meeting the Bay of Biscay, forms the approximate limit
of the Herring’s extension in the Atlantic. That the
Herring besides is really an oceanic fish, is clearly
shown by the fact that it attains its maximum size and
its highest development of form in the ocean. The
largest Herrings in the market come from Iceland,
Norwegian Nordland, and Norwegian Finmark. Their
average size (to the end of the caudal lobes) is 33 —
37 cm. Exceptionally large specimens are indeed known
from other localities. From England we are told of
Herrings 39 and 43 cm. long; but the most trustworthy
and the latest accounts in Day (1. c.) give 32 cm.

as the maximum length. The Bohuslan Herring of the
present day is also in general of the same size, though
exceptions are mot unknown. We have mentioned above
a Herring from Helso (Northern Bohuslan) which, though
shrunk by the spirit in which it is preserved, measures
42 cm. to the end of the caudal lobes. It is not ab-
solutely impossible that such Herrings may be native
to our waters; but exceptions make no rule, and it
must be regarded as most probable that the Herring
attains its highest development, and has the centre of
its range, in more northern regions, in the North At-
lantic between Iceland and Norway. It is, beyond
doubt, from this source that the Herring-fisheries of
the North Sea and Norway, as well as of Bohuslan,
derive their fluctuating supply".

The experience of many years teaches that the
Herring comes to Southern Norway from the north-
west. The Scotch Herring-fishery also begins yearly
off the Shetland Islands and gradually extends further
and further south. The Herring thus makes from the
north an annual ascent of the plateau, offering a depth
of at most 100 fathoms, on which Great Britain and
Ireland are situated. If this plateau (see the map) were
raised to the surface of the ocean, it would form an
unbroken stretch of dry land between the said islands
and the Continent from the Skaw south to about the
middle of the Bay of Biscay. The west coast of the
territory thus produced would coincide to the south
with the present limit of the Herring’s range in this
part of the Atlantic. Between this plateau and Nor-
way, along the west coast of Southern Norway, runs
the deep, so-called Norwegian Channel into the Skager
Rack, its depth falling short of 200 fathoms from the
neighbourhood of Bergen south to Lindesnas, i. e. out-
side the region where the Norwegian fishery for Spring
Herring is carried on. The channel is rather narrow
off Lindesnas, but widens and grows deeper in the
Skager Rack, so that between Arendal and the Skatv,
somewhat nearer to Norway than to Denmark, there

a This opinion I advanced in 1878, as an explanation of the revival that had just begun in the Bohuslan Herring-fishery and of

the prospects for its continuance, of which the authorities were then very dubious. About 1870 the fishery for Spring Herrings on the

south coast of Norway had begun to decline. In 1872 the Royal Museum had obtained quite typical Spring Herrings from Stromstad. Dur-

ing the first years of the decade the Museum received from Bohuslan several pelagic species, Bonitos and other rovers of the open Atlantic,
notorious as pursuers of the Herring. In 1856 too A. V. Malm had taken off Kalfsund Herring-fry 46 — 49 mm. long, which, as was then
assumed, could not possibly be the young of the Grass Herring (a smaller kind, spawning in spring) that is always to be found on the coast of
Bohuslan. It therefore seemed beyond dispute that the Great Herring had been present for several years in the North Cattegat and the Ska-
ger Rack, without being directly observed in those waters, until a part of the Herring army penetrated, after they had spawned, into the
island-belt. I found every reason to believe, as has since been confirmed, that a new so-called Herring-period had set in on the coast of
Bohuslan, i. e. that the Herring had made its way from the North Sea into the Skager Rack, deserting for this locality its spawning-places
off the south-west of Norway.

HERRING.

965

are nearly 400 fathoms of water. The continuation of
the channel, up to. the 40-fathoms line, follows the
west coast of Sweden south to about the latitude of
Kongsbacka, i. e. as far as the true coast fishery for
Great Herring has ever extended to the south in
Sweden.

To trace the Herring to its ocean home is a hope-
less task. Conjecture must still play a prominent part
in all opinions of its life in the depths. But so much
is apparently clear, that its wanderings follow the de-
pressions and steeps of the bottom. How deep it can
descend, we know not; but the advanced development

that it can descend to considerable depths, of some
hundred fathoms at least, though probably not to the
deepest parts of the Atlantic.

As Mac Cullocii has remarked, the rovings of
the Herring have three motive causes, the quest of a
spawning-place, the chase of food, and the dread of
its pursuing enemies.

The spawning-season of the Herring lasts all the
year round; but only a part of the wandering multi-
tudes, in general those of the same age, are in breed-
ing condition at the same time. Different spawning-
seasons may hence characterize different localities, and

KChrisUa/iia

\herdect

fscox

^EXGLAISTD /

rN^Loridon 0( ,

0 100 fathoms
100 -1000 -
WOO -2000'-

fesliMva over <000
Scale 1 22,000000

Fig. 241. The North-eastern Atlantic together with the North Sea and the Baltic. After 0. Torell.

of the adipose membrane covering the head ranges it
nearer than the Mackerel to the true deep-sea fishes.
The pressure need not greatly distress the Herring, for
only few fishes can so easily adapt the distension of
the air-bladder to varying degrees of pressure. The
lower temperature of the depths need not deter it, for
fishes with so extensive a geographical range must by
nature be accustomed to widely different temperatures,
though the Herring seems sensitive to sudden varia-
tions. The same may be said of the greater or less
salinity of different layers of water. All that we know
of the Herring’s nature well admits of the assumption

in the same locality it is not uncommon to find two
separate spawnings every year. In the Atlantic the
two spawnings, where they occur, take place before and
after the winter, in the Baltic before and after the
summer. This applies to the main body of the shoals;
but during the intervening periods isolated instances of
breeding fish may be observed almost everywhere. As
a rule the oldest Herrings spawn first, in the autumn,
the younger ones later, when the spring has set in or
in the summer. From the Gulf of Bothnia Gisler ad-
duces the Spring or Ice Stromming, which spawns within
the island-belt when the ice breaks up, the Net Strom-

966

SCANDINAVIAN FISHES.

ming, which spawns outside the island-belt from the
middle of May to the end of June, and the Autumn
Stromming , which spawns within the island-belt at the
end of August and beginning of September". In the
island-belts of Stockholm and Morko* * * 6 the true spring
spawning occurs at the end of May and beginning of
June, the autumn spawning at the end of August and
beginning of September. In the south-east of the Bal-
tic, near Dantzic and Konigsberg each of the two
spawning-seasons (varying from March — June and Au-
gust— September) is of about the same importance to the
fishery. Off Rtigen and on the coast of Scania the
autumn spawning is the more important; in the extreme
west of the Baltic, off Trave and Schleswig, the spring
spawning, takes the upper hand. “The further south
Ave advance along the Scanian coast”, writes Lundbeeg0,
“the more seldom do Ave meet with sprmg-s pawning
Herring. The fishermen state, one and all, that several
years may elapse Avithout a single one being seen ....
The spring-spaAvning Herring of the Sound, considered
by G. Winther to be a distinct race, peculiar to the
Sound, Avould also appear to consist of small specimens
of the common Herring. Our Stvedish fishermen there
do not knoAv of any spring-spawning Herring in the
Sound, nor is there any regular fishery for Spring
Herring on the SAvedish side. A little Herring is
indeed taken all the year round, mainly to serve
as bait. But the Herring-fishery proper of the Sound
and Cattegat does not commence until the middle or end
of August or the beginning of September, and lasts till
the middle or end of October. Most of the Herrings
taken, however, are not in full breeding condition, i. e.
though they are full of roe and milt, it is not quite ripe
or running. At Kullen I Avas assured that Herrings in
this last condition or spaAvning Herrings are very seldom
caught during the true season for the Herring-fishery;
but that, after the Herring-fishery proper is over, later
in the autumn, at the beginning of November or end
of October, shoals of spaAvning Herrings seek shelter,
Avhen the Avind is in the north, under the lee of Kulla-
berg, and are then taken close in shore.” In the Cat-

tegat the true spawning-season begins in the middle of
September. The Herrings taken in drift-nets off the
coast of Bohuslan during the latter half of August, 1882,
Avere for the most part not yet in breeding condition;
but on the 26th of September I found spaAvning Herrings
among the takes made 3 — 5 miles off Marstrand. That
Avas the first time, during the present Bohuslan fishery
for Great Herring, Avhen it Avas conclusively shoAvn that
the Herring spawns, at least toAvards the end of September,
off the said coast, and that a very profitable fishery with
drift-nets might be carried on for at least a month be-
fore. That, later in the year, it continues its spaAvning
Avithin the island-belt as Avell, Avas proved by Mr. C. A.
Hansson, avIio found Herring-roe ready for hatching in
Stromstad Fjord on the 5th of March, 1885. More re-
cently (in 1888) Trybom observed the September spaAvn-
ing of the Herring in the North Cattegat off Fladen,
Groves-Flak, Lilia Middelgrund, and the shelving banks
north of Anholt, and outside the Cattegat at. the edge of
the shalloAvs Avest of the SkawJ Farther out in the Skager
Rack, betAveen Hanst.holm (Jutland) and Christiansand
(Nonvay), somewhat nearer to Denmark than to Norway,
Heincke met Avith “spawning Bohuslan Herring” in the
middle of September, 1889 b The younger Herring,
the Grass Herring of the Bohuslan island-belt, spawns
in spring, during March, April and May.

The Herring, like most fishes, chooses its spawning-
places in Avater shalloiver than its ordinary home, and
during youth at least, it also seeks for this purpose
water of less salinity7 *. The large Herring can spawn
in 60 — 100 fathoms of water 9, though it usually comes
nearer to the coast; the smaller Herrings ascend into
water sometimes no more than a fathom deep. In the
neighbourhood of Stockholm the spawning is performed,
according to Sundevall, outside the island-belt or in
the larger fjords, on rises of the bottom or reefs Avith
5 — 10 fathoms of water, close to land sometimes in
only 3 fathoms. In the Baltic fishery the rule applies,
that the males are the earlier, both in age and season,
to attain breeding condition — during the spaAvning-
fishery the males are usually taken first. But Boeck

“ He further adduces from these regions sill, large males, with soft and running milt at midsummer or earlier, Seine Stromming,

shotten Spring Stromming which is resting during the early summer and is then taken within the island-belt, and Red-bellied, Stromming,

answering to the Fat Herring of the Atlantic, with sexual organs not yet tumid.

6 Cf. Sundevall and Ekstr&m.

c Meddelanden rdrande Sveriges Fiskerier, Haft. I, p. 40.

d Sillundersokningar vid Sveriges vestlcust hosten 1888, Kongl. Civildepartementet, III, p. 16.
e Mitth. Sect. Kirsten-, Hochs. Fischerei, Deutsch. Fisch. Ver., Jan. — Febr. 1890, p. 23.

1 Meyer, Jahrber. Comm. Unters. Deutsch. M., Kiel, 1874 — 76, p. 232.
o Boeck, Nord. Tidskr. Fiskeri, Kbhvn, Aarg. 2, 1875, p. 263.

HERRING.

967

states quite the opposite of Norway". When the spawn-
ing begins, however, the sexes mingle promiscuously,
and the operation is performed much in the same way
as we have above described when treating of the Gwy-
niads. Gisler tells in animated language how the Baltic
Herring then approaches land in large and densely pack-
ed shoals, often extending more than three-quarters of
a mile along the coast, and always containing many
times more females than males. The fish tumble about,
and lash with their tails so violently that the scales drop
off and float to the surface, in company with small air-
bubbles which the Herrings emit. The sea is dyed gray,
and a powerful and rank smell, appreciable at a great
distance, fills the air. The spawning multitude does
not shun the net, but rather presses willingly into its
meshes. The operation does not last long, however, at
the same spot, “probably no more than five or six hours”,
according to Sundevall, when the shoal withdraws.
Out at sea the spawning presents a similar sight: in a
confused mass, gleaming with phosphorescent light, the
fish toss about, near the surface when the night is dark
and the 'weather mild; deeper in the water when there
is moonlight and in frosty weather* 6 * 8. The impregnated
eggs sink to the bottom, and attach themselves to sea-
weed, stones, shells, and other firm objects, or some-
times cake together on the gravel or sand, or even on
a clayey bottom.

The number and size of the eggs vary, as usual,
according to the size of the mother fish and their own
degree of ripeness. Their number may be estimated at
about 20,000 — 40,000 in different females; their size,
when they are ripe and ready to be deposited, varies
in the Baltic Herring0, generally speaking, between 0‘92
(exceptionally 0'85) and 1 mm., in the North-Sea Her-
ring between 1 and l'o mm/ Of the development we
learn from Sundevall’s notes0 that, “after the embryo
had been formed, it was seen (in August) to turn in the
egg seven or eight times a minute, subsequently more

seldom, and latterly only once every two or three mi-
nutes. The hatching generally takes place in a fort-
night or a little more, but in water of a higher tem-
perature, over + 20° C. (+ 68° Fahr.) for example, only
three days are required/ As long as the fry retain the
yolk (fig. 242), they move in a peculiar manner. By
violently bending or tossing the body, an operation
repeated every second, or at somewhat greater or less
intervals, they work their way upwards, to the surface
(at least when they are confined in vessels 3 — 6 dm.
deep), and as soon as they have touched the surface,
they keep still and sink again to the bottom, where
they lie for a while, and then resume this upward mo-
tion. — As soon as the yolk is absorbed, which takes a
week’s time, they commence swimming in dense shoals,
with a serpentine movement. The tins and the general
shape of the body seemed to have attained their full
development in the course of two or three months,
when the fish is about 36 mm. long.” These remarks
apply to the Spring Stromming. But so great may be

Fig. 242. Newly hatched Baltic Herring, 7 nun. long, taken on the
12th of May, 1854. C. J. Sundevall.

the difference between the summer and winter growth
that the Autumn Stromming, which develops during the
colder months, requires in the Baltic, according to Mobius
and Heincke, 7 — 9 months to attain the said degree of
development6', and has then grown to a length of more
than 60 mm. The ventral tins are developed in the
Spring Herring of the Baltic, according to Heincke6,
when the fish is only 251/ 2 mm. long; but in the Autumn
Herring they do not appear until the fish measures
33 V2 mm. The following comparison has been drawn
by Meyer between the growth of the Spring Herring
in the Baltic and that of young Trout:

° 1. c., p. 26.

6 For a description of the noisy spawning at sea, and how the Herring immediately after the operation quits the spawning-place, see
Cuv., Val., 1. c., XX, p. 87.

c Kupffer, Jahresb. Comm. Unt. D. Meere, Kiel 1874' — 76, p. 177.

d According to Boeck, 1. e., 1*1 mm.

e Om Fiskyngels utveckling , Vet.-Akad. Hand!., Bd. I (1855), p. 17.

/ The- hatching takes place, according to Kupffer (1. c., p. 29), in water of a temperature between + 14° and + 19° C. within 6 —

8 days; according to M&bius and Heincke ( Fisch Osts., p. 137), at a temperature of + 10° or + 11° C. in 11 days and at a temperature
of + 7° or + 8° C. in 15 days.

g According to Meyer (Comm. Deutsch. Meer. 1874 — 76, p. 248), however, small Autumn Herrings 45 — 60 mm. long and of almost
perfect shape are found in February.

h Comm., 1. c., p 128.

Scandinavian Fishes.

122

968

SCANDINAVIAN FISHES.

Length im-

Length

Length

Length

Length

mediately

1 month

3 months

6 months

12 months

after exclu-

afterwards,

afterwards,

afterwards,

afterwards,

si on, mm.

mm.

mm.

mm.

mm.

Herring

5-8

17—18

45—50

75—80

130—140

Trout

15

20

30

64

125

When it leaves the egg, the young Trout is thus
about twice as large as the young Herring; but three
months afterwards it has been outstripped by the latter.

The larviB of the Herring are distinguished by an
elongated (Eel-shaped), somewhat terete, and highly
transparent body, with the vent situated far back, espe-
cially during the earlier stages, with the dorsal fin be-
hind the middle of the body, but with the ventral fins
set farther forward. The paired fins are lobate (with
brachiate base), the pectorals appearing before the ven-
trals. In the Spring Herring of the Baltic the scales
are developed at a length of only 41 mm.

When two years old, the Baltic Herring is about
20 — 21 cm. long; when three years old, 22 — 23 cm.;
when four years old, 24 — 25 cm. In Norway the Spring
Herring, it. is assumed, requires 7 or 8 years to reach
its full size. The males attain maturity at a length of
16 — 20 cm., the females at a length of about 21 cm.

After passing through the larval stage, the Herrings
gather in shoals and commence their rovings along the
coast, often ascending into a foot of water, and pursued
by Garpike and other predatory fishes. On the shelv-
ing beach near Halmstad, though the Herring is else
a stranger to Laholm Bay, the shoals of young Herrings
were so dense in my childhood that during summer we
boys used to catch numbers by wading out and netting
them in our handkerchiefs. In July, 1834, about five
miles west of the Skaw, within the innermost sand-bar,
Krdyer*2 observed a shoal of Herring-fry that were about
5 cm. long. The shoal extended for a distance of about
200 yards and was more than 20 yards broad. It was
so dense that the water seemed quite black. Herring-fry
one year old are of rarer occurrence in the open sea6;
but on the 12th and 13th of August, 1889, Heincke0
found several specimens 15 — 20 cm. long among larger
Herrings in the Skager Rack, on the seaward side of
the Jutland Bank, where the depth was about 100 m.
The Herring can thus endure the hardships of a life

in the troubled ocean, and accompany its elders on
their wanderings, at an early age.

The Herrings seem to seek strength, or at least to
find a sense of security*2, in the close companionship of
numbers, however little this avails. Out at sea they
probably swim in more open order and mingle indis-
criminately. The drift-net fishermen of Northern Scot-
land, who sometimes sail 80 miles from land, take full
Herrings and shotten Herrings, ripe Herrings and maid-
■en Herrings (matjes), in the same haul. But the nearer
the Herring-shoal approaches to the shore, at different
seasons of the year, the denser and the more exclusive-
ly do the different sorts congregate, in summer those
which are most eager in the pursuit of food, in autumn
those with the ripest organs of reproduction. Large
shoals of maiden Herrings (fat Herrings) — with a
thick coat of fat round the intestine, but with small,
filamentous or ribbon-like organs of generation — re-
pair in summer to the west coast of Norway north of
Stavanger, and feast on their favourite food, the so-
called Aat. The spawning Herrings, on the other hand,
resort in still greater multitudes during autumn and
winter, alternately, as it appears, for a series of years
to the southern part of the west coast of Norway, and
for another, shorter period to the Skager Rack and the
Northern Cattegat, as well as to the fjords of Norwegian
Nordland. In the last-mentioned region the Herring
sometimes advances within the island-belt before the
spawning, but goes out again to breed. To the south
of Norway and off the coast of Bohuslan the greater
part of the Herring-shoal spawns before its appearance
within the island-belt, at least in the beginning of the
said period. In the Baltic the Herring goes out to sea
after spawning; but on the west coast of Scandinavia it
makes its way into the island-belt and the fjords, and
rests there for a while, before withdrawing to the open
sea. At these times it even enters fresh water. Moreau
states that it ascends the mouth of the Seine up to
Quillebeufh After resting it again begins to feed, puts
on flesh, and becomes what is known in Norway as
slosill (with thread-like organs of generation). But the
Herrings of the shoal are not all in the same condition,
although most of them are ready to spawn. The main

a Danm. Fiske, vol. Ill, p. 159, note.
b Boeck, 1. c., p. 18.

c Mith. Sect. Rust., Hochseef. (Deutsch. Fisch. Ver.) 1890, p. 17.
d Cf. the above-mentioned observations of the Three-spined Stickleback (p. 657).
e Cf. also Cuv., Val., 1. c., p. 66.

HERRING.

969

body sweeps along, and in the press swim Herrings
small (so-called lottsillar), middle-sized, and large —
shotten Herrings, spawning Herrings, and Herrings not
yet in breeding condition.

The Herring is, no doubt, impelled, like other mig-
ratory tishes, by a well-known instinct, to retrace its
way to the waters which first afforded it a home; and
when the spawning Herring has found a locality where
the circumstances are congenial to reproduction, it re-
turns there, as a rule, during the following years, so
long as these circumstances continue. On the coast of
Bohuslan it has been observed, both in former times"
and recent years, that during the commencement of a
so-called Herring-fishery period the fish as a rule come
earlier year by year, but afterwards later and later,
until, tired of the place, they neither return in such
numbers, nor approach so near to the coast. They have
found more favourable circumstances in another locality
or farther out to sea — a more tranquil spawning-place,
a more plentiful supply of food, and greater security
both for themselves and their offspring. The Scotch
fishermen too have learnt that they may sometimes shoot
their drift-nets for Herring quite close in shore, but in
other years must sail far out into the North Sea through-
out the fishing-season. These variations, which have
been called Herring-fishery periods, cannot be determined
beforehand. Our historical knowledge thereof, defective
as it is, by no means points to definite periods in fixed
succession. That they may depend on meteorological
and hydrographic, not to say cosmic, alterations, we
cannot positively deny; but the periodicity of these al-
terations is as yet unknown*. Century after century,
ever since the eleventh, we have at least indications that
the Herring has “come in” on the coast of Bohuslan.
From the end of the twelfth century, at least down to
1537, the Herring-fishery of the Sound, with Skanor and
Falsterbo as its headquarters, was far famed; but Lund-
berg has shown c that this was probably due, less to the
greater abundance of the fish then, as compared with
the present time, than to the commercial relations then
obtaining, to the part taken in the fishery by the
powerful Hanse Towns, and to the business-like manner
in which they turned its resources to account. The

Baltic Herring (Stromming) never appears in such vast
numbers, and in this respect, as well as in its moderate
size, more closely resembles the Shore Herring of Bo-
huslan. But Herring-fishery periods of abundance and
scarcity occur in the Baltic too, as Sundevall has
shown of the island-belt of Stockholm d. He adduces
the instances that the seine-fishery for Stromming was

successful .
unsuccessful
successful .
unsuccessful

at Nynas
up to 1825 incl.,
. 1826 — 1839,

. 1840—1849,

. 1850 et seqq.,

Arsta

. . . 1828,
1829—1841,
1842—1851,
1852 . . . ,

Forsvik
. . . 1834,
1835 — 1843,
1844—1851,
1852 . . . ,

“so that each new period of plenty or scarcity sets in
some years later, the farther the locality is situated
within the island-belt.”

Ekstrom gives the following description of the Baltic
Herring’s life in the island-belt of Morko: “In spring,
as soon as the ice has broken up, and cleared away in
some degree, the Stromming ascends from the depths
where it has wintered. It rises so near the surface as
to be carried along by any storm or gale. When it has
come so near the coast that it does not wish to advance
any further, a halt is made, the whole shoal turning
round with military precision, and facing the wind.
Thus it remains, almost motionless, until the wind
changes and blows from the coast. It now turns again
and faces the wind and the shore, but draws nearer the
land to seek suitable stations. If it has drifted to a
strange coast, it roves along the shore until it comes to
places that seem convenient for the spawning and as a
home during the summer. It now stays in their neigh-
bourhood, roving about, now farther from land, now
closer in shore, according to the direction of the wind,
for on its wanderings from place to place it invariably
swims against the wind or current, except, as mentioned
above, when it drifts in spring to the coast. This mig-
ration, which depends on the direction of the wind and
the set of the current when the ffsh ascends from the
depths, renders the fishery more or less productive on
different coasts .... Among these islands at any rate,
such has always been the case. The experience of many
years has taught the fisherman that, if the ice breaks
up during a south-westerly gale, which is usually of
long duration at this time of year, and which sets into

a See Fagr^us, Trangrumsacten (1784), p. 129.

b “Bruckner assumes a periodic variation in the heat radiatecj by the sun, and thus explains the variations of temperature, of atmo-
spheric pressure, and of the rainfall. The thirty-five years’ period has nothing to do with the number of sunspots.” Kremser, Meteorolo-
gische Zeitschrift (Wien), 1891, p. 228.

c Det stora sillfishet i Slcane under medeltiden ocli nyare tidens borjan , Autiqv. Tidskr. f. Sverige, Del. 11, No. 2.
d Stockholms Lans Kongl. Hushallningssallskaps Handlingar, 6:te Haftet (1855), p. 187.

970

SCANDINAVIAN FISHES.

the island-belt, he may expect a successful fishing-season.
But if the wind is north-east at the beginning of the
spring, the fishery is always a failure. These migrations
extend, however, over no great distance, being restricted
to limits of some few leagues. The islander has a rough
knowledge of the coast-line to a distance of at least
some leagues from his home, and consequently can tell
against which promontories or into which inlets the
Stromming is driven by the wind prevailing for the time.
So the islanders of Morko say, “If this wind holds, they
will have Herring in the islands of Oster Gothland;”
“With this wind the Herring will come to the island-
belt of Stockholm;” etc. The places most affected by the
Stromming are shallows with a level bottom in the large
fjords, or the shores that abut on deep water, but do not
fall sheer into the depths, having a fairly level bottom
between the outer edge and the land. Such shores are
generally to be found in these islands off promontories.
They are all the better if there is a current. The bottom
should be sandy or stony, and overgrown, at least in
patches, with weeds. About midsummer, at the middle
or end of June, the spawning is over, and the Strom-
ming retires to deeper water. Towards autumn, in
August, it again ascends, but never visits at this season
the places where it has passed the spring or spawned,
repairing instead to much deeper spots. In December
or even earlier the greater number withdraw to their
winter-quarters, which they choose in some deeper part
of the sea. These places are not the same . year after
year, for, when the Stromming is taken in winter with
the ice-seine, it is found standing now at one spot, now
at another; but it keeps, generally speaking, to the same
neighbourhood. The islanders of Morko have certain
strowimingsvarp, i. e. certain sheets of water where
Strfimming may be taken with the ice-seine, but only
the tract is known, not the exact spot.”

The gregariousness of the Herring is bound up with
its timidity. That it is easily alarmed by noise, we
have already remarked, and Ekstrom adduces evidence
to prove that the passing of steamers may frighten it
away from the navigable channels of the island-belt, and
also that the mere setting of gill-nets is sometimes
enough to disturb its spawning and drive it away from
a fishing-station of ascertained value. In Bohuslan too
the firing of guns is now prohibited during the fishing-
season. The Herring is not tenacious of life. The ra-
pidity of its death is notorious, and has given rise in

many places to the proverbial expression, “as dead as
a Herring.” But, according to Ekstrom®, “the infor-
mation we possess on this head is exaggerated. It is
generally believed that the fish dies the very moment
it is lifted above the surface of the water. I have per-
sonally made numbers of experiments to test this state-
ment, and have found that the time varies with the
temperature of the atmosphere. In spring, at the end
of April, when the air is still cool and usually cold,
the Stromming lives 18—20 minutes after it has been
taken out of the water. If caught late in the evening or
at night, it sustains life for fully half an hour. But it
must lie handled carefully and not exposed to any ex-
ternal violence. As the summer approaches, at the
middle of May for example, it never lives more than
6 — 10 minutes, and at midsummer, when the air is
quite warm, the duration of its life out of the water
seldom exceeds 4 minutes. It should be remarked, how-
ever, that the individuals on which I made the above
experiments had not been entangled in the meshes of a
net, but were taken, quite uninjured, in a vessel out of
the water and deposited on the beach or in the boat.
If the Stromming has been caught in the seine, it dies
almost at the moment it leaves the water, and those taken
with gill-nets are dead before they are drawn up.”

In spite of its feeble dentition the Herring ranks
among the predatory fishes, though its victims are
usually of small size. In its earliest youth it lives on the
most minute marine animals. At a length of 11 mm.
a Herring larva had begun to feed on the larvte of
worms (Lindstrom), and a young Herring 17 mm. long
had its intestine “full of food, amongst which small
species of Cyclopidce might easily be recognised” (Sun-
devall). In its later youth and during the rest of its
life the Herring, no doubt, lives principally on Entom-
ostraca, Schizopods, and Pteropods. Certain parts of
the North Atlantic teem with these animals, which are
so plentiful as to afford a sufficiency of food to the very
largest whales. The Norwegian fishermen class the
Herring’s ordinary food ( Aaten ) under three heads, which
they call jRodaat, Gulaat , and Svartaat or Krutaat. The
first consists chiefly of Copepods, either extremely small
or (e. g. Calanus finmarchicus ) as much as 8 mm. long.
“It seems incredible,” writes Boeck, “that creatures so
small can play so important a part in the economy of
a whole country; but the Mackerel and the Autumn
Herring owe their fatness almost entirely to these ani-

Cf also Cuv., Val., 1. c., p. 63.

HERRING

971

mals, and under the microscope we may see the layers
of fat between the muscles and viscera within the frail
shell of these minute organisms.” The Gulaat is made
up principally of Annelid larvae, the Svartaat of small
mollusks, the larvae of larger mollusks, and Pteropods.
When the Herring has gorged itself with this food, and
not had time for digestion or to excrete the remains,
it is worthless as an article of trade. The fish decom-
poses rapidly, will not absorb the salt, and emits a
disgusting stench. In Norway it has therefore been
prescribed that such fish must be left three days in the
seine before being drawn up and salted. The Svartaat

271) enter into the diet of the Herring; and it is no
doubt partial to all other kinds of small fishes.

The Herring has enemies in numbers. Among fishes
its most destructive foes are the Cod, the Coalfish, the
Hake, large Scombroids, the Salmon, and Sharks. Among
whales the lesser rorqual ( Balcenoptera rostrata), the
great northern rorqual ( Balcenopt . I ait ceps) and the fin-
whale {Balcenopt. musculus ) feed on Herrings. The por-
poises are also Herring eaters; and among the most
troublesome of all are the seals, which often leave no-
thing in the net but the heads of the fish. Numerous
birds (gulls, terns, the gannet, etc.) swell the tale of

Pig. 243. A Yarmouth drift boat out fishing. After Holdsworth.

is the worst, and almost hopelessly spoils the fish. But
animals of a relatively small size are not the Herring’s
only food: it can also devour fishes of comparatively
great dimensions. In the stomach of a large Baltic
Herring Ekstrom found three good-sized specimens of
Gobius minutus; and the stomach of a female Herring
3 dm. long, which was caught in the island-belt of
Stockholm in January, 1893, and in which the ovaries
were swollen to about a third of their full size, con-
tained portions of five Herring-fry that had been almost
equal in size, and the largest of which had measured
6 V2 cm. We have remarked above that the Doubly-
spotted Goby (p. 251) and Cry st alio gobius Nilssonii (p.

its persecutors. But in the war of extermination against
the Herring the most prominent part is taken by man.

In the open sea and in the great arms of the sea
the Herring is taken in drift-nets, gill-nets, and purse-
seines. Nearer land the gill-net and purse-seine are
also used, but most commonly the seine (Sw. vad“) or,
as in Denmark, the band-yarn , a sort of stake-net.

The drift-net is a continuous train of nets (Sw.
lank) seized to a stout rope or warp (Sw. drift-rep ), to
which large tloats (clumps of wood — Sw. Jclabbar, kob-
bar — or kegs — Sw. dunkar , brillar — or bags of skin,
usually dogskin — Sw. sdckbojar) are attached with
stronger seizings, so as to keep the whole ‘drift’ afloat

° The word vcid is difficult of explanation. Some write vada, and derive it from the verb vada (to wade). The Danes write v&d ,
and it is a question whether the word is not the same as that we find in tclci dning svad (a breadth of cloth) etc.

972

SCANDINAVIAN FISHES.

in a vertical position0. In the North-Sea fisheries one
of these ‘fleets’ of nets frequently extends to a length
of a mile and a quarter or more. The net is shot in
the evening and hauled up in the morning. During the
night it is allowed to drift with the boat, the warp
being made fast in the bows. Drift-nets may also be
used as gill-nets , the two ends being anchored; but the
common gill-net, called skote in the Baltic, ndring in
Halland, is of simpler construction and resembles the
net used in fresh-water fishing. In recent times the
North American* * 6 purse-seine has been introduced into
Europe and employed in the Bohuslan fishery. It is a
deep seine, with stout and closely corked head-rope but,
strictly speaking, without foot-line, which is replaced
by a purse-line running free through rings. The seine
is shot in a circle round a shoal of fish swimming at
the surface; and when the circle is complete, and the

net walls in the shoal, the purse-line is hauled in, till
the seine assumes the shape of a bag. The land-seine-
is an engine as common and well-known as any kind
of net; but in. the Herring-fishery it attains its maxi-

mum dimensions, being so long and deep that whole
arms of the sea or great portions thereof may be en-
closed so as to bar the Herring’s retreat. The fish are
then secured in smaller seines. The construction of
the stake-net ( bundgarn , fig. 244) we have described
above (p. 352).

The voracity of the Herring is such that it may
also be taken with hook and line, and sometimes no
bait is necessary, the mere glitter of the dancing hook
being sufficient to entice the fish.

The annual take of Herrings can hardly be esti-
mated; it must amount to thousands of millions. In
the Baltic and the Sound more than 200,000 barrels of
Strommings and Herrings, valued at about £166,600,
are taken yearly by Swedish fishermen. A barrel of
Strommings contains on an average more than 2,200
fish. It is estimated that 1,900 — 2,000 Scanian Herrings
go to the barrel. According to these estimates the
average annual catch of the Swedish fishermen alone
exceeds four hundred million Strommings and Herrings,
and we may surely assume that the Finnish, Russian,
German, and Danish fishermen together take at least
three times as many. To judge from Dr. A. H. Malm’s
latest reports on the Bohuslan fishery, the average an-
nual take of Herrings for the fishing-seasons 1888 — 92
was about five or six hundred millions, a hectolitre
containing at least 300 Herrings. The average annual
value of this fishery for the said four seasons was about
£100,000, but in 1891 — 92 the value rose to £130,000.

The several takes are given in the following extracts
from Dr. A. H. Malm’s “Reports on the Sea- fisheries of
the Province of Gothenburg and Bobus”0:

Herr

i n g - f i

s li e r y

with

s e i

n e s.

gill-

nets.

drift

-nets.

Hectolitres.

Value.

Hectolitres.

Value.

No. of fish.

Value.

1891—92

1,403,438

£59,493

228,708

£65,453

5,254,560

£5,373

1890—91

1,094,185

£65,115

196,821

£49,288

9,651,360

£5,676

1889—90....

1,538,046

£30,334

223,102

£39,394

—

£7,706

1888—89..

1,443,390

£28,535

279,089

£24,380

—

£4,788

The value of the Halland Herring-fishery for the I spector of Fisheries, at £3,272, and for the year 1891
year 1890 is estimated by Mr. Tkybom, Assistant In- | at £3,660. The Herring-fishery of Denmark yielded in

a For a more minute description of the drift-net see Holdswokth, Deep-Sea Fishing and Fishing Boats , p. 100; Lundberg, Meddel.

rorande Sveriges Fiskerier, Haft. I, p. 33, with illustr.

6 At the Fisheries Exhibition (London, 1883) a net of this kind, coining from Cornwall, was catalogued as original.
c These reports do not take into account that portion of the fishery which falls to the share of fishermen from Halland.

HERRING.

973

1890 nearly £45,000 (£3,457 from Liim Fjord"). Nor-
way’s income from this source6 in 1891 was £360,600'’.
The results of the Herring-fisheries of Great Britain
and Ireland were in 1887 and 1 8 8 8 d :

1887.

1888.

Cwts.

£.

Cwts.

£.

England and Wales..

1,605,140

442,288

1,729,641

485,806

Scotland

3,217,361

641,572

2,846,796

614,838

Ireland

—

—

75,548

30,728

The Herring-fishery of Holland yielded in 1 880 e about
227,000,000 Herrings and in 1881 about 197,500,000,
with a market value of between three and four million
guldens (£250,000 — £333,000). The French fishermen
of the North Sea and the Channel took in 1 88 1 f 39,000,000
kilo, of Herrings, valued at about 9,000,000 francs
(£360,000). The Iceland Herring-fishery of 1882 9
yielded 50,000 barrels of a total value of £72,000.

On the European side of the Atlantic the annual
take of Herrings thus amounts to between 500,000,000
and 600,000,000 kilo., and commands a market price
of about £2,700,000. On the east coast of North Ame-
rica, according to Hind6, Herrings were taken in 1874
to a weight of about 91,000,000 kilo. No great accu-
racy can be expected of all these calculations and esti-
mates, but they give at least a notion of the value
represented by the Herring, for all its cheapness. The
welfare of nations has depended on the Herring- fishery ;
and none need be surprised that this fishery has always
been an apple of discord.

Many are the culinary forms in which the Herring
appears. In order to fit this cheap article of food for
transportation from the place of its catch to the world’s
ernporia, it has been the custom from time immemorial

to dry, smoke, and salt the Herring; and the kitchen
has taken measures accordingly. The Herring-fishery
became an El Dorado to the Dutch when they learnt
to gut the fish1 — to remove the gill-arches and in-
testine — before salting. But the Herring, like other
fishes, is best and cheapest when fresh; and with the
speedy means of transit and improved methods of pre-
servation — borate or, still better, ice — now available,
it may be conveyed to a considerable distance, and
kept fresh for at least a fortnight or three weeks. In
this condition it is excellent either boiled or fried. The
English whitebait , the principal course at the fashionable
dinners at Greenwich or Blackwall on the Thames, has
a world-wide reputation. Whitebait consist chiefly of
Herring-fry about two or three inches long, but also
of Sprats, Sticklebacks, Gobies, and other small fishes.
They are taken at flood-tide in spring and summer
with a special kind of net, which is dipped a few feet
below the surface from a boat anchored in 30 — 40 feet
of water. They should, above all, be procured quite
fresh and fried as soon as possible. They answer to
the fiskakaga (fish-cake) of Scania and Halland, only
that the latter is made into a cake by the addition of
a greater quantity of butter or lard. In France White-
bait are known as blanches ( blaquets ) or menis (me-
nus ses).

Herrings and Herring-fry are also much used as
bait for other fishes.

The numerous names by which the Herring is
known in trade, and the details of the numerous me-
thods employed in curing it for the market, cannot
be given here. The reader who is interested in these
questions will find an able treatment of them in
L.tungman. 1. c.

a Drechsel, according to the “Dansk Fiskeriforenings Medlemsblad” 1892, pp. 74 and 75.

h For the Norwegian Herring-fisheries and their fluctuations see H. Baars, Die Fisehereiindustrie Norwegens , Bergen 1873, p. 36.
c The Central Bureau of Statistics at Christiania, according to the “Dansk Fiskeriforenings Medlemsblad” 1892, p. 490.
d Fish Trades Gazette 1889, Jan. 12th and 26th.

e According to an official statement, in the Catal. Gt. Intern. Fish. Exhib. London 1883, 1st ed., p. 422.

J ibid., p. 384.

‘J ibid., p. 382.

h Br. -Goode, Fisher., Fisher. Industr. U. S., sect. I, p. 549.

1 The discovery is ascribed by tradition to Willem Beukelzoon, a skipper from Biervliet in Flanders, d. 1387.

974

SCANDINAVIAN FISHES.

THE SPRAT (sw. SKARPSILLEN OR HVASSBUKEN).

CLUPEA SPRATTUS.

Plate XLIV, fig. 2.

Length of the base of the anal fin more than 1 4 of the distance between the ventral fins and the tip of the snout.

Number of spines at the ventral margin 3.2 — 35°.

R. br. 6 — 7*; D. ^^(17 — 19); A. ^-^-(18-20);

P. — ; V. -V-1; O. so + 1 + 17 4- 1 + «;■ Lin. lot. 47—50;

15 — lb bc

L. tr. 11 — 13; Vert. 46 — 49.

Syn. Sprattns 1. Sparlingus (Harengorum foetura), Willughb., Hist.

Pise., p. 221; Bay, Syn. Pise., p. 105; Latulus, Schonev.,
Ichthyol. Slesv. IJolsat., p. 41. Clupea quadrinncialis, ma-
xilla iuferiore longiore, ventre acutissimo ( Spratt Anglis,
Hvassbuk Suecis), Art. Icht., Gen., p. 7; Syn., p. 17;
Spec., p. 33; Lin., Fna Snec., ed. I, p. 120. Brisling,
Strom, Sondm. Beskr., p. 271.

Clupea Sprattus, Lin., Syst. Nat., ed. X, tom. I, p. 318;
Penn., Brit. Zool. (ed. Warrington, 1776), vol. Ill, p. 303;
Retz., Fna Suec. Lin., p. 353; Swartz, Suensk Zoologi,
vol. I, No. 28; Nilss., Prodr. Ichthyol. Scand., p. 22;
Yarr., Brit. Fish., ed. I, vol. II, p. 121 ; Parn., Mem.
Wern. Nat. Hist. Soc., vol. VII, p. 322, tab. XXXV; Kr.,
Voy. Scandin., Lappon., (Gaim.), Zool., Poiss., tab. 18;
Damn. Fisk., vol. Ill, p. 177; Cov.. Val. ( Harengula ),
Hist. Nat. Poiss., vol. XX, p. 285; Sund'ev. {Clupea),
Stockh. L. Hush. Sails. Handl. 1855, pp. 81, 108, 185; Nilss.,
Skand. Fna, Fisk., p. 516; Mgrn, Finl. Fiskfna, p. 69;
Lindstr., Gotl. Fisk., Gotl. L. Hush. Sails. Arsber. 1866,
p. 20 (sep.); v. Bemm. {Harengula) in Herkl., Bouiust.
Fna Nederl., part. Ill, p. 381; Gthr {Clupea), Cat. Brit.
Mus., Fish., vol. VII, p. 419; Cole., Forli. Vid. Selsk.
Chrnia 1874, Tillaagsb., p. 193; 1879, No. 1, p. 98;

N. Mag. Naturv. Chrnia, Bd. 29 (1884), p. 112; Seidl.,
Fna Balt., p. 98; Malm, Gbgs, Boh. Fna, p. 582;
Winth., Naturh. Tidskr. Kbhvn, ser. 3, vol. XII, p. 48;
Mor. {Meletta), Hist. Nat. Poiss., Fr., tom. Ill, p. 447 ;
Bncke {Clupea), Fisch., Fischer., Fischz. O., W. Preicss.,
p. 172; Hcke, Var. Her., VII Ber. Comm. Deutsch. M.,
pp. 45 et 60; Mela, Vert. Fenn., p. 354, tab. X; Day,
Fish. Gt. Brit., Irel., vol. II, p. 231, tab. CXXXIX, fig. 2;
M6b., Hcke, Fisch. Osts., p. 139; Lillj., Sv., Norg. Fislc.,
vol. Ill, p. 97.

Clupea Schoneveldii, Kr., Voy. Gaim., 1. c., Damn. Fiske, vol.
Ill, p. 193.

Spratella pumila, Cuv., Val., 1. c., p. 357 + Meletta vul-
garis, p. 366; tabb. 600 et 603.

The Sprat is in most respects so like the Herring
that a detailed description is unnecessary. It is, how-

a According to Heincke.

6 Sometimes 8, according to KrGyer.
c Sometimes 7, according to Heincke.
d With 88 mm. as the average length.
e With 120 mm. as the average length.

ever, a much smaller fish, measuring at most about 17
cm. to the tips of the caudal lobes; but at this smaller
size it has undergone changes of growth in many re-
spects more complete: it is, in a word, a more fully
developed Herring form. This appears, for instance, in
the length of the lower jaw, a measurement which we
have never found to exceed 53 % of the length of the
head, whereas in the Herring it has never proved less
than 55 % of the same. In this character the altera-
tions of growth run in decreasing percentages from the
young to the older specimens.

In Swedish the Sprat has been distinguished from
the preceding species by the name of sharp sill (Sharp
Herring), a reference to its most palpable characteristic
in most cases, the sharper ventral margin, with more
prominent spines on the ventral plates; but this char-
acter is often illusory. A more trustworthy distinction
is the difference in the number of the ventral plates,
which in the Sprat does not exceed 35, 20 — 24 (com-
monly 22 — 23) in front of the ventral fins and 9 — 13.
(commonly 11) behind them.

The form of the body is the same and shows the
same variations as in the Herring, being deeper or
shallower, but in the deeper form of Sprat ( Clupea
Schoneveldii) the ventral profile is still more curved in
comparison with the dorsal. During the growth of the
body from a length of 66 mm. to one of 104 mm/
(from the tip of the snout to the end of the middle
caudal rays), the least depth of the body has proved
to be on an average 7'1 % of the length thereof; and
during growth from 108 — 147 mm.e we have found
this average to be 8*1.

The head shows the peculiarity that the eyes are
generally somewhat larger than in the Herring. During
growth from a length of 73 mm. to one of 147 mm.

SPRAT.

975

the longitudinal diameter of the eye has proved to vary
between 31 and 26 % of the length of the head. The
branchiostegal membrane does not form so distinct an
angle with the inferior margin of the suboperculum;
this angle is sometimes almost imperceptible, the poste-
rior (upper) branchiostegal rays extending further back
than the interoperculum, and the last of them being
obliquely truncate at the extremity or produced upwards
to a blunt point, which is applied to the lower margin
of the suboperculum. The number of gill-rakers on the
first branchial arch we have found to vary between 46’
and 51, thus fewer than in the Herring. The dentition
resembles that of the Herring, but is feebler, the teeth
being often imperceptible, and usually wanting on the
palatines and vomer, but often present on the ento-
pterygoid and ectopterygoid bones. The most important
difference from the head of the Herring consists, how-
ever, in the comparatively short lower jaw, which is
always shorter than the base of the anal fin, and the
length of which varies between about 47 and 5 2* 1/2 %
of that of the head. The length of the maxillaries too
varies in the Sprat between 89 and 84 %, in the Her-
ring between 75 % (in exceptional cases 73 %) and 79 %
(in exceptional cases 81 %), of that of the lower jaw,
which is besides perceptibly shorter in the Sprat than
the distance between its hind extremity and the upper
angle of the base of the pectoral fin, in the Herring
equal to this distance or a little longer.

The dorsal fin is of the same form and occupies
the same position as in the most developed Herrings.
At a length of body averaging 88 mm. the average
distance between the dorsal fin and the tip of the snout
proved to be 49'9 % of the length of the body, and in
specimens averaging 120 mm. in length the correspond-
ing percentage was 50’7. The distance between the anal
fin and the tip of the snout is somewhat less than in
the Herring, and varies between about 65 and 71 %
of the length of the body; but the base of the fin is
longer, its length varying between 13 and 15 % of that
of the body. The caudal fin is usually not quite so
deeply forked as in- the Herring. The length of its

middle rays is about 6V2 % of that of the body and
rather more than V8 — 2/5 of that of the longest rays
in the inferior caudal lobe.

The pectoral fins resemble those of the Herring in
their length as well as in other respects. The ventral
fins are in general shorter and have fewer rays than in
the Herring. Their length varies between 9 7 3 and 772 %
of that of the body. Their position affords one of the
most important characters of the Sprat, the distance
from the tip of the snout to the anterior (upper) angle
of the base of these fins extremely seldom — and then
only slightly — exceeding the length of the body,
and being less on an average than that between the tip
of the snout and the beginning of the dorsal fin, rarely,
and then but slightly, greater than the latter distance.
It is consequently the rule that the ventral fins are in-
serted vertically below the beginning of the dorsal fin
or further forward. The preabdominal length varies
between 28 and 3072 the postabdominal between
about 17 and 22 %, of the length of the body.

The scales are exactly similar to those of the Her-
ring, and the most essential difference in the abdominal
plates has been remarked above.

The coloration too is so like that of the Herring
that no constant difference can be adduced. The line
of lustrous green between the dorsal and ventral colours
is sometimes ribbon-shaped, rather sharply defined. Ac-
cording to Valenciennes* this line is sometimes a gold-
en band; and Collett mentions0 “a half-grown spe-
cimen” that had lost its scales, but was marked along
this part of the sides with two, almost black stripes.

The geographical range of the Sprat does not ex-
tend so far north as that of the Herring, but else co-
incides therewith in the East Atlantic. On the Ame-
rican side the Sprat has never been found; and Faber’s
statement*2 that it occurs off the coast of Iceland is
not convincing, for he describes its operculum as stri-
ated, in distinction from the smooth operculum of the
Herring. But on Gunther’s authority0 the Sprat has
been included, together with several other European
fishes7, in the fauna of Tasmania.

“ 48'3 — 50'4 /, according to our measurements.

4 1. c., p. 288.
c 1879, 1. c.

d Fische Islands, p. 178.
e Proc. Zool. Soc. Lond. 1871, p. 672.

I Scicena aquila, Zeus faber, Caranx trachurus, Stolephorus encrasicholus , Conger vulgaris , Orthagoriscus mola, Rliina squatina , Galeus

Cfinis, Squalus ( Aeanthias ) Blaivvillei, etc.

Scandinavian Fishes.

123

976

SCANDINAVIAN FISHES.

In the Mediterranean the Sprat is replaced by a
very closely allied species, the Melette of the French,
Clupea phalerica0. Judging from a specimen 90 mm.
long, taken off Genoa, and sent to ns by Professor
Giglioli, the Melette is distinguished from the Sprat

by the greater depth of the tail, the more advanced
position of the dorsal tin, and the somewhat greater
size of the head. Its relations to the Sprat in these
respects are most readily indicated by the following
table of percentages:

Averages

One specimen

in

of

Clupea sprattus.

Clupea phalerica

Length of the body from the articular knobs of the maxillaries to the end of the middle caudal rays, mm _

87.7

120

90

Length of the head in % of the length of the body

21.2 b

20.8

22.2

Distance between the dorsal fin and the articular knobs of the maxillaries ,, ,, „ „ „ „ „ „

49.9 c

50.7

47.8

Least depth of the tail - „ „ „ „ „ „ „ „

7.1

8.1 d

8.8

Leugth of the head __ in % of the distance between the dorsal fin and the articular knobs of the maxillaries

42.5 e

41.0

46.5

Least depth of the tail ,, n u 55 •? ?? ?? ?? ??

14.3

16.0/

I8.1

The Sprat is common from Trondhjem Fjord (Col-
lett) to La Rochelle (Valenciennes). It is said to oc-
cur, though scantily, northwards to the Lofoden Islands,
as G. 0. Sars was informed by the fishermen, and
southwards in the Bay of Biscay beyond the Gironde (Mo-
reau). In Scandinavia it is most common off the south-
west of Norway and the coast of Bohuslan; but it is
taken in numbers within the Baltic, and is common,
according to Mela, throughout the Gulf of Finland, as
well as in the Gulf of Bothnia up to lat. 64° N., rarer
to lat. 65°. On the Baltic coast of Russia, according
to Seidlitz (1. c.), only solitary specimens are met with
that have attained full growth and sexual maturity,
though the young are plentiful. Off Memel and Dant-
zic spawning Sprats are common in May and September;
and in Putzinger Wiek (the eastern part of the Gulf of
Dantzic), says Benecke, countless spent Sprats may be
seen in the May spawning- season floating dead at the
surface. In Kiel Bay the Sprat is also common. On
all the coasts of Denmark the Sprat occurs, though only
at a few points in any great number (Wintiier). In the
Zuyder Zee and all round Great Britain it is as plen-
tiful as anywhere in Scandinavian waters.

The Sprat-shoals live and feed in the same man-
ner as the Herring-shoals, and often intermingle with
the latter. It is, we may say, a rare occurrence to
take any quantity of Strommings or small Herrings

without finding a number of Sprats in the catch; and
in the same ivay small Herrings are found interspersed
among a haul of Sprats. The Sprat-fishery cannot in-
deed be so productive in most cases as the Herring-
fishery, the Sprat being a much smaller fish; but con-
sidering the size of the fish the Sprat-shoals are hardly
inferior in point of numbers or density to the Herring-
shoals. The same gregariousness, the same manner of
life, and the same diet conjoin the two species with
each other. They are also so similar in appearance
that only a practised eye can distinguish between them,
unless the observer has experience enough to feel the
difference in the sharpness of the ventral margin.

The spawning-season of the Sprat occurs within
the island-belt of Stockholm, according to the joint
observations of Sundevall and Cederstrom, in June.
In the west of the Baltic, according to Heincke, the
Sprat spawns both in spring and autumn. He observed
in Kiel Bay during the months of July and August
Sprat-fry 25 — 35 mm. long, which he assumed to date
from the spawning in April and May, and during the
latter part of October and till December other speci-
mens 23 V2 — 33 V2 mm. long, which could consequently
not have been spawned, in all probability, earlier than
in July. Malm found in July and August at Strom-
stad and off Gaso (Bohuslan) young Sprats 20 — 57 mm.
long, which would thus seem to be the offspring of a

“ Melet'tes , Belon, p. 208. Apliya phalerica , Rond., p. 212. Glupanodon phalerica , Risso, Ear. Mer., tom. Ill, p. 452; Mor.
(Clupea), III, p. 445. Meletta mediterranea, Cuv., Val., XX, p. 369. Alosa papalina , Canestr., Fna D'ltal., Peso., p. 135.

6 Maximum in the specimens measured by us 21
c Minimum ,, „ ,, ,, „ „ 48

d Maximum „ ,, ,, ,, ,, ,, 8

e 4a

11 11 11 11 11 11 11

11 ii 11 'I 11 11

/

SPRAT.

977

later or earlier spring-spawning, perhaps even of a
winter-spawning, the same year. From the South of
England we are told by Holdsworth" that the true
spawning-month of the Sprat in the neighbouring wa-
ters is January, but that this fish also spawns during
the summer; and he assumes, with reference both to
the Sprat and the Pilchard, that the winter-spawning
is performed near land, the summer-spawning at the
surface in deeper water. But off the coast of Bohuslan
the Sprat seems to be in a breeding condition in July or
August as well, for at Stromstad Malm took “females
almost ready to spawn”, on the 15th of July, 1 865*.

Of the hatching of the Sprat’s eggs we are igno-
rant; we do not even know whether the ova float at
the surface, or sink, like those of the Herring, to the
bottom. But the fry have been described both by Malm
and Heincke, and differ, according to the latter, from
Herring-fry of the same size in the orange tone of
their coloration, the greater depth and thickness of the
body, the earlier development of the ventral fins, the
earlier prolongation of the air-bladder, and the smaller
number of the vertebrae

After spawning the Sprat makes its way into the
island-belt, where it soon recruits its strength and puts
on flesh, in this respect easily surpassing the young
Herrings. In the inner island-belt of Stockholm the
mass of Sprats begin to appear in August. “Till Oc-
tober Sprats are occasionally taken, after which time
they again disappear. When they come, they are ra-
ther lean; but towards autumn they grow fatter” (Sun-
devall). During the same season, however, Sprats are
caught in the outer shallows and in the outer fringe of
the island-belt. Even in winter, as at the time of
writing (March, 1893), Sprats are plentiful among the
consignments of Stromming exposed for sale in Stock-
holm. A successful Sprat-fishery is also carried on
among the Smaland Islands off Krakvik0, but we have
no information of any such fishery farther south on
the east coast of Sweden. The chief Sprat-fishery of this
country belongs to Bohuslan, especially to the neigh-
bourhood of Fjellbacka. “Here the Sprat puts in an
appearance at Michaelmas, and remains until some time

after Twelfth Day. Up to Twelfth Day, or at least
during the weeks just before Twelfth Day, it swims in
shoals by itself; but afterwards it is accompanied by
young Herrings, generally two years old” (A. W. Malm).
It is ‘ Said to come in both north and south of the
Weather Islands, straight from the sea (the Skager
Rack). But shoals of Sprats sometimes arrive later in
the winter, as in 1892, when this fish was taken on the
south coast of Bohuslan during February and March
(A. H. Malm). In England too the main body of the
Sprat army lies nearer land during winter.

The Sprat-fishery has much in common with the
Herring-fishery, and is carried on both with skotar (fine-
meshed nets) and seines. In Bohuslan the annual take
was officially stated during recent years as follows:

1888 .. 1,900 hectolitres, valued at A 1,373,

1889 4,270 „ „ „ £2,176,

1890 2,550 „ „ „ £2,077,

1891 3,803 „ „ „ £4,062.

From these Sprats, which they can procure fresh,
the inhabitants of Bohuslan prepare the best quality of
their Anchovies, a kind of Pickled Flerring, not to be
confounded with the true Anchovy, which is a rare fish
in Sweden. But a considerable proportion of the Sprats
thus preserved in Bohuslan comes salted from Norway,
where the annual take, according to Collett, sometimes
exceeds 100,000 hectolitres. In Norway and England
too Sprats are prepared for food in the same way; and
according to Day, three million tins of Sprats, pickled in
a similar manner as the anchovies of the west coast of
France, are annually imported into England. In Ger-
many and in certain parts of Scotland the Sprat is
smoked. This is the method employed in curing the
well-known Kieler- Sprotten. Mild smoked Sprats in

oil are not inferior to Sardines.

The Sprat suffers chiefly from the ravages of the
same enemies as the Herring. But its eyes are attacked
by a parasitic crustacean, Lernceonema monillaris. This
Penellid shines at night, and to this origin Day ascribes
a supposition common among English fishermen, that
the Sprat-shoal is often guided and lighted on its way
by “lanthorn Sprats”.

a Deep Sea Fishing and Fishing-Boats, pp. 133 — 135.
b Gt. Intern. Fish. Exhib. London 1883, Sweden, Spec. Catal., p. 170.
c Intern. Fischerei-Ausst. Berlin 1880, Schwed. Catal., Spec. Th. , p. 25.

978

SCANDINAVIAN FISHES.

Subgenus ALOSA.

Sides of the body marked , at least in front , with dark spots. Lower anterior surface of the operculum coursed
by more or less distinct grooves and ridges. Dorsal fin set so far forward that the distance between it and the
tip of thj, snout is less than four times the postorbital length of the head. Inner part of the caudal lobes fur-
nished with foliate scales , larger than the others.

Now that all attempts to employ the dentition as
a means of defining subgenera within the genus Clupea
have proved futile, no better prospect of attaining this
object is offered, it appears, than by the different de-
velopment and position of the fins. But the subgenus
Alosa, which is further distinguished by the extremely
feeble dentition of the mouth, is so well defined within
the Scandinavian fauna that no difficulties meet us here
in its characterization. In exotic forms, however, as

a b

Fig. 245. Scales from the left side of a Pilchard 20 cm. long (a)
and from a ■ Shad 30 cm. in length ( b ). X 3.

for instance in the Japanese Clupea zunasi , the striation
of the opercula is hardly perceptible, and the lateral
markings of the body are confined to a single small
spot on the adipose membrane of the upper scapular
region; and in the Mediterranean Clupea aurita, where
the dark markings are also restricted to the upper part
of the tract round the gill-openings, the said striation
consists merely of a ridge crossing the anterior part of
the operculum in an obliquely downward direction. But
the other characters given in the above diagnosis range
these forms beside our Shads, which may be regarded
as the type of the subgenus, and which are usually to
be recognised at the first glance by the striation of the
opercula and the spots on the sides of the body, though

the latter vary both in size and distinctness, being in
some individuals wanting.

The scales of this subgenus are more strongly
developed at the basal angles of the fins and their bases
than is the case in the preceding members of the genus.
The caudal fin is covered to a great extent with scales,
and bears on each side two patches of these, one on
the upper and one on the lower lobe, among which the
hindmost is the largest, and is also 'venous. Sometimes,
however, these patches of scales are so thin and trans-
parent that they easily escape observation. At the base
of the dorsal fin, the upper angle of the pectoral fin,
and fhe outer angle of the ventral fin, we find pointed
scales, which partially conceal these fins; and rounded
scales overlap the loiver margin of the pectoral fins,
covering the lowest rays, while the base of the anal
fins is also scale-clad. In texture too the scales of
this subgenus differ from those of the preceding one,
the radiating grooves, which are there extremely faint
streaks (see above, p. 957), being here far more sharply
defined (-fig. 245).

Both the anal and the dorsal fins show a more
pointed prolongation of the outer posterior corner than
in the preceding subgenus.

The most constant character, hoAvever, is the for-
ward position of the dorsal fin, a character which is
expressed in the above diagnosis of the subgenus by a
comparison with the postorbital length of the head.
This last measurement is relatively greater in the Shads
than in the true Herrings, the validity of the character
being thus unimpaired even when the trunk is shor-
tened, as in the American Menhaden, which would else
be referred by the position of the dorsal fin to the-
preceding subgenus.

HERRING-FISHES.

979

THE PILCHARD OR SARDINE.

CLUPEA PILCHARDUS.

Fig. 246.

Length of the maxillaries at most about 40 % of that of the head. Longitudinal diameter of the eges at least
about 60 %, least depth of the tail at least about 80 %, length of the suboperculum at the suture with the oper-
culum at least about 55 %, of the length of the maxillaries. Scales comparatively large , their number in a lon-
gitudinal row on the sides of the body at most, about 30.

9 o 9 3 i

R. hr. 7 “; D. — — —(17—18); A. (17 — 186); P. -;

14— 15v 14— 16v 15—16’

V. — ; G. x + 1 + 17 +A+x\ L. lat. 28 — 30; L. tr. 8 — 10; Vert.
50—53.

Clupea sprattus, Brunn., Ichthyol. Massil., p. 82.

Ulupea sardina, Cuv., R'egne Atiim., ed. 2, tom. II, p. 319
(+ Cl. pilchardus, ibid.); Mob. ( Alosa ), Hist. Nat. Poiss.
Fr., tom. Ill, p. 458.

Syn. Sardines ou Celerins , Belon, Nat., Dio. Poiss., p. 167.

Harengus minor sive Pilchardus, Willughb., Hist. Pise., p.
223, tab. P, 1, fig. 1.

Clupea Pilchardus, Walb., Ichthyol. Art., pt. Ill {Gen. Pise.),
p. 38 (varietatem Clupece harengi dixit); Bl., Naturg. Ausl.
Fisch., pt. IX, p. 40, tab. CCCCVI; Yarr., Hist. Brit.
Fish., ed. 1, vol. II, p. 96; Cuv., Val. ( Alausa ), Hist.
Nat. Poiss., vol. XX, p. 445, tab. 605; Nilss. {Clupea),
Skand. Fna, Fisk., p. 522; Kr., Tidskr. Fisk. Kbhvn,
Aarg. 2 (1868), p. 71; Steind., Stzber. Akad. Wiss. Wien,
LVII, i (1868), p. 738; Gthe, Cat. Brit. Mus., Fish., vol.
VII, p. 439; Coll., Forh. Vid. Sels. Chrnia 1874, Tillcegsh.,
p. 194; Cederstr., Ofvers. Vet. Akad. Forli. 1876, No. 4,
p. 66; 1879, No. 2, p. 62; Malm, Gbgs, Boh. Fna, p.
585; Day, Fish. Gt. Brit., Irel ., vol. II, p. 224, tab.
CXXXIX, fig. 1; Bncre {Alosa), Handb. Fischz., Fischer.
(M. v. d. Borne), p. 170; Lillj. {Clupea), Sv., Norg.
Fisk., vol. Ill, p. 106.

The Pilchard is a more terete fish than the Her-
ring, but is otherwise similar in form. It is inferior
to the Herring in size; but in this respect a distinction
has long been maintained between two forms, the Me-
diterranean, which is the smaller, being about as large
as the lott-sill (small Herring) of Bohusl&n or the com-
mon Stockholm Stromming, and which is known by
preference as the Sardine, and the Atlantic Pilchard,
which attains, according to Cornish c, a length of at
least 3 dm., according to I)uNNd, of 3 1/2 dm., including
the whole caudal tin. In Sweden only adult specimens
have been met with, the largest 'measuring 285 mm.
in length (Malm), the smallest 218 mm.

“ 6 — 8, according to Day.

* 19 — 21, according to Gunther; 21, according to Lilljeborg.
c Zoologist, 1879, p. 62.
d In Day, 1. c., p. 230.

■980

SCANDINAVIAN FISHES.

The greatest depth varies between about 19 and
22 % of the length of the body to the end of the middle
caudal rays; and the greatest thickness between about
45 % (in small specimens) and 59 % of the greatest
depth.

The length of the head (measured as above, in the
Gwyniads and Herring, from the articular knobs of the
maxillaries) varies between about 22x/2 and 21 1/2 % of
that of the body. The diameter of the eyes, which
are circular, measures (in Pilchards 81 — 211 mm. long)
between 30 and 23 %, and the interorbital width (at
the middle of the eyes) 20 — 24 %, of the length of the
head. The length of the snout is equal in small Pil-
chards to the diameter of the eyes; in adult specimens
it shows individual variations between about 26 and
30 % of the length of the head. The nostrils — the
anterior in each pair is merely a narrow slit — lie
nearer to the tip of the snout than to the eyes. The

Fig. 247. Head of a Pilchard, with feebly developed covering of
adipose membrane. Natural size.

tip of the mouth, which is formed by the margins of
the intermaxillaries, is incised at an oblique angle,
owing to the position occupied by these bones with re-
spect to each other. The length of the maxillaries,
which extend back to about a line with the anterior
margins of the pupils, decreases in the above-mentioned
specimens from about 39 lf2 to 3 7 1/'2 % of that of the
head, and their breadth varies between about 36 and
32 % of their length. The length of the cheek (from
the hind extremity of the maxillary to the posterior
margin of the preoperculum) is much greater, as in the
Herring, than its height below the eyes. The length
of the lower jaw, the symphysis of which shows hardly
any thickening, decreases in Pilchards of the above
lengths from about 55 to 48 % of that of the head.

The cheeks and opercula vary in appearance according
to the density of the adipose membrane on the head.
When this membrane is thin (shrunken, fig. 247) the

a Of. Moreau’s definition of the genus Sardinella (1. c., p. 450):

striation both of the lower anterior part of the oper-
culum and of the preoperculum and suborbital bones
consists of distinct ridges and grooves. But in other
instances, when the adipose membrane is thicker (fig.
246), its ducts with their numerous ramifications occupy
the grooves, and render these as well as the ridges less
distinct. The upper part of the scapular region is en-
tirely covered with this growth of adipose membrane,
the elevated pores granulating the upper surface of the
head. Measured in an horizontal direction the postorbital
length of the head (from the apparent hind margin of
the eye to the middle of the posterior opercular margin)
increases with age from about 41 — 46 % of the entire
length of the same; measured in an oblique direction
(from the middle of the hind margin of the eye to the
lower posterior angle of the operculum) it varies between
about 47 and 53 % of the latter length, and is some-
what more, even in young specimens, than one-fourth
of the distance between the dorsal fin and the articular
knobs of the maxillaries. The operculum forms an al-
most rectangular parallelogram, with the upper posterior
angle rounded, the posterior side rather convex, and the
inferior side straight. The last-mentioned side (the sub-
opercular suture) measures about 22 or 2 1 1/2 % of the
length of the head. The form of the suboperculum
varies, its breadth (height) being about one-third of its
length in young specimens, about one-half of the same
in adult Pilchards. The preoperculum is broad, with
rounded, rectangular corner. The interoperculum is
narrow, and has the same relation to the posterior
bran chi ostegal rays as in the Herring, these rays form-
ing a rectangular sinus with the inferior margin of the
suboperculum. On opening the operculum we find that
the black posterior limit of the pharynx and the branchial
cavity is rectangular®, corresponding to the form of the
operculum itself, so that the posterior margin of the
latter coincides with the vertical arm of the clavicular
angle, and its inferior margin as well as the suboper-
culum with the horizontal arm, the margin of which
consists of the above-mentioned process (answering to
cl p in fig. 238, p. 950). The gill-rakers are long and
fine, with dense and regular lateral spines of small size.
On the first branchial arch we have counted 110 gill-
rakers, 67 on the lower (horizontal) part of the arch,
and 43 on the upper. The dentition of the mouth is
extremely feeble, being usually confined to a few teeth
at the tip of the lower jaw and a crenelation of the

Ceiuture scapulaire a borcl anterieur vertical”.

PILCHARD.

981

maxillaries at the posterior part of their under margin.
The palate and tongue are almost toothless".

The dorsal tin begins at a distance from the tip of
the snout (the articular knobs of the maxillaries) mea-
suring in small specimens (Sardines) about 41 V2 in
large (Pilchards) about 40 %, of the length of the body,
and apparently not exceeding 90 % (84 — 87 V2 % accord-
ing to our measurements) of the distance between the
ventral tins and the same point. Its base, which is about
equal to its height, measures about 12 1/3 — 1 1 1/3 % of
the length of the body. The distance between the anal
tin and the tip of the snout is about 66 — 70 % of the
length of the body. Its outer margin, like that of the
dorsal tin, is somewhat concave. Its base occupies about
14 — 16 % of the length of the body, and is four to three
times its height. The caudal tin was damaged in all
our specimens; but, to judge by the remnants, it, seemed
to be deeply forked, the length of the middle rays being
about 74 °f that of the longest ones, which was about
17 % of that of the body.

The pectoral fins are rather more pointed than in
the Herring. Their length is about 15 — 14 %, that of
the ventral tins about 872 — 873 %, of the length of the
body. The preabdominal length is somewhat greater
than the postabdominal, but both are contained about
four times in the length of the body.

The size and texture of the scales we have men-
tioned above. At the ventral margin we have counted
20 spinigerous scales in front of and 16 behind the
ventral tins. These spines are set in a groove, which
is almost entirely concealed by the lowest scales on
the ventral sides.

The coloration of a new-caught, specimen is thus
described by Malm: “Back above bluish green, sides
and belly silvery white, with a dash of yellowish green
towards the tail. The whole opercular apparatus, and
especially the upper scapular region, as well as the
anterior part of the iris and a portion of the maxil-
laries, of a light, lustrous yellow, like that of sulphur-
pyrites, which extends some way over the shoulder-
girdle. On the shoulder-girdle, but someAvhat above
the middle of the side, a light, blood-red spot. Central
part of the operculum also blood-red. The whole

gill-cover iridescent, resembling the most beautiful
mother of pearl. Pupil rounded. Iris nacreous, tinged
with orange behind, of a greenish gold above, with a
whitish ground; finely punctated with black. Lower
jaw yellowish at the tip. Tip of the snout blackish.
Orbit, of an emerald ground-colour in front of the eye.
Below the lateral line a row of six small, blackish spots,
the first, situated in the red colour of the operculum,
the second on the upper part of the shoulder-girdle, the
third in front of the tip of the pectoral fin, separated
therefrom by a distance equal to about half the diameter
of the pupil. Distance between the third spot and the
last one equal to the length of the pectoral fin. Dia-
meter of these spots half that of the pupil. Fins greenish
gray; anal lighter; ventrals colourless. Tongue so dense-
ly punctated with dark dots that it is almost blackish”.

The Pilchard is of the same importance in Medi-
terranean countries and, to a considerable extent, in the
west of France and south of England as the Herring in
the North- Atlantic fishery. Boiled and preserved in oil,
it is known all the world over; but its geographical range
is not so wide as the Herring’s, and it can hardly vie in
numbers with the latter. To the south it is known off
Madeira and the Canary Islands6. To the north it is a
stranger c even in the North Sea above Yarmouth, still
more so in Scandinavian waters, though it has been
found as far north as Bergen, whence Lilljeborg brought
home to Upsala Museum two Pilchards in 1858. Malm
mentions three specimens from Bohuslan, one of them
taken at Kalfsund on the 15th of May, 1855, the other
two at Stromstad, respectively on the 11th of August,
1865, and the 15th of July, 1869. The Royal Mu-
seum has received from Bohuslan two specimens 23 cm.
long, the first obtained by Prof. S. Loven in 1861, the
second taken in Roster Fjord by Mr. C. A. Hansson
on the 25th of July, 1877. According to Nilsson a
female about 25 cm. in length and with ripe roe was
caught among common Herrings off Kullen on the 25th
of September, 1849. Krqyer mentions a specimen ta-
ken in a stake-net ( bundgarn ) in Kaerteminde Fjord
(Fiinen) during October, 1867.

The true Atlantic home of the Pilchard thus lies
south of England, where it appears on the coasts of

a “After the tongue has dried, a thin carina may be traced along it, with some extremely fine teeth, visible beneath the magnifying-
glass, and set in a single row, on the inner part.” Nilsson, 1. c., p. 524.
h Lowe, Trans. Zool. Soc. London, vol. II, p. 189; Steind., 1. c.

c In modern times at least. On the east coast of Scotland, according to Parnell, the case was formerly different; — see Mem. Wern.
Nat. Hist. Soc., vol. VII, p. 321.

982

SCANDINAVIAN FISHES.

the Spanish Peninsula and France and off the south
coast of England, in the same manner as the Herring
further north. In the southern part of its range, north-
wards along the coast of the Spanish Peninsula and in
the Mediterranean, it is said to be met with near land
all the year round. But where the deep water, bounded
by the 100-fathoms line, lies farther from shore, be-
tween the south of the Bay of Biscay and the British
plateau, the influence of which on the Herring’s geogra-
phical range has been noticed above, the Pilchard, a fish
that rests and winters in the depths of the ocean, ap-
proaches land in an order the reverse of that observed
by the Herring. The Herring comes, as we have seen,
from the north, and spreads round the coasts of Great
Britain. The Pilchard appears earliest in the year off
the south coast of France, and the fishery commences
later and later to the north". In deep water, however,
the Pilchard occurs all the year round off the coasts of
Cornwall and Devon, being often found, according to
Day, in the stomach of fish taken on long-lines in
January; but the Pilchard-fishery with drift-nets does
not begin until July, and with the seine not until
August.

The Pilchard’s habits are the same as the Herring’s.
It is extremely timid and gregarious, but greedily feeds
on minute animals or even on the spores of seaweed and
on Diatomacece, its sound appetite gaining it the fatness
for which it is famed. The most important Pilchard-
fishery on the French coast depends on this voracity.
Fish-roe is imported from Norway — principally Lofoden
Cod-roe, which costs the French fisherman 80 — lOOfrancs
or more a barrel — and this expensive, bait, sometimes
mixed for the sake of economy with sand or chopped
meat and pounded crustaceans, is strewn on one side
of the net floating at the surface behind the boat, when
the Pilchard is seen on the other side of the net. When
the net is as full of Pilchards as the fisherman thinks
proper — which he sees by the sinking of the cork-line —
a fresh net is shot, so long as the store lasts, and the
fishing goes well. The bait is costly, but justifies the
outlay; and in good years the French fishermen take
one or two milliards of Pilchards, worth fifteen to twenty
million francs. Of the Cornish Pilchard-fishery Fox* 6

states that the export of salted Pilchards from Penzance
and Falmouth to Genoa, Leghorn, Naples and Venice
amounted in 1879 to 11,938 hogsheads or about 35 —
36 million fish; and according to Couch0 the English
export of these fumadoes has averaged for many years
30,000 hogsheads, sometimes amounting to 60,000.

In 1879 the purse-seine was first used in the
French Pilchard-fishery, but it has not yet come into
favour. In the Cornish fishery this engine has been
employed longer, even previous to its introduction into
the American Mackerel fishery.

The Pilchard-fishery is subject to the same fluc-
tuations as the Herring-fishery, being still more de-
pendent on the temperature of the water, but naturally
too on the supply of food. According to Dunn16 it has
more than once happened during very severe winters
that- the English Channel proved too shallow to afford
the Pilchard protection from the cold. On these occa-
sions he saw “countless millions” of Pilchards perishing
or floating dead at the surface. In quest of food the
Pilchard army roves westwards from the Channel out
into the Atlantic; but Dunn adduces instances to show
that this fish sometimes stays off the Cornish coast,
feeding on the multitudes of crustacean larvae in the
Zoea-st-age. With the same object it also makes its
way up the Irish Channel; but it seldom extends its
wanderings to the North Sea. In the migrations of the
Pilchard and its approach to the coast so-called semi-
secular periods may be observed0, as in the case of
the Herring-fishery.

The spawning of the Pilchard is probably performed
in the open sea. Late in May and in June Dunn met
with great numbers of ripe Pilchards 25 — 30 miles
south of the Lizard; but the species breeds', it is stated,
in winter as well, and, according to Yarrell, as early
as in October. “I have reason to suppose,” says Couch,
“that the spawn is shed at the surface, and mingled
with it a large quantity of tenacious mucus, in which
it is kept floating while it is obtaining the vivifying
influence of the light and warmth of the sun,” an
assumption that has been corroborated by the obser-
vations of Dunn and Cunningham7. According to Day
the female Pilchard lays about 60,000 eggs; and it has

“ See Vaillant and Henneguy, Rapp. Gen. snr la pgclie de la Sardine (Gerville-Reache), Paris 1888, p. 28.

6 In Buckland, Nat. Hist. Brit. Fish., p. 165.
c Fish. Brit. 1st., vol. IV, p. 93.

(l Fish. Trades Gazette, No. 485 (3rd Sept., 1892), p. 9.
e See Couch, 1. e., p. 85.

f Mith. Sect. Kiist., Hochs. Fischer., Deutsch. Fisch. Ver., 1892, p. 4-5.

SHAD.

983

long been known that during the spawning-season the
Pilchard is so dry and tasteless as to be scarcely
eatable.

Man is not the Pilchard’s only enemy. Fishes and
Dolphins make havoc among its shoals. Among them
Sharks (such as Squalus acanthias and C archarias glau-
cus) and Scimnoids are the most destructive.

Couch “ has described and figured a Clupea squa-
mo-pinnata, which is stated to have larger scales than
the Shads, and has therefore been explained by Gun-
ther6 as a hybrid between the Pilchard and either the

THE SHAD

CLUPEA

Fig. 248 and PI

Length of the maxillaries at least about 45 % of that c
about 50 %, least depth of the tail about 75 %, length
of the maxillaries. Number of scales in a rou

Twaite or the Allice Shad, though the two last-mentioned
forms do not breed, so far as we know, in salt water.
The defective specimen, the only one known, can only
serve, which was all that Couch intended, as an in-
centive to further investigations. More noteworthy is
another form, also found in English waters, which re-
sembles the Pilchard in the striation of the opercula
and the large scales on one side of the body, but has
smaller scales, like those of the Herring, on the other
side. This form has been figured and described by Day*
as a cross between the Pilchard and the Herring.

iW. stamsillen).

ALOSA.
te XLIII, fig. 2.

f the head. Longitudinal diameter of the eyesd at most
of the sub operculum" at most about 49 %, of the length
along the sides of the body at least about 60.

Wmm

Fig. 248. Clupea alosa, var. finta , 9, * 1 2 of the natural size. Taken in a Salmon-trap off Killingholm (Stromstad Fjord) on the 18th July,

1881, by Mr. C. A. Hansson.

“ 1. c., p. 123, pi. CCVI.

1 Cat., 1. c., p. 436.

c Proc. Zool. Soc. Loud. 1887, p. 129, pi. XV.
d In specimens more than 1 dm. long.
e At the suture with the operculum.

124

Scandinavian Fishes.

984

SCANDINAVIAN FISHES.

A. Var . finta — Twaite Shad (Sw. Staksitt). Fig. 248.

Gill-rakers on the first branchial arch about- 40 — 60
(fig. 249, a).

B. Var. alosa — Allice Shad (Sw. Majfisk). Pi. xliii, tig. 2.

Gill-rakers on the first- branchial arch about 90—120
(fig. 249, b).

R. br.

-9; D. ; A.1

14 — 16

17-

— ; P-

-21 14

1 ■ V 1 •

— 16’ ' 7—8’

C. x + 1 + 17 + 1 + x; L. lat. 60—65*; L. tr. 17"; Vert. 55—59'

R. br. 7—8; D.e 5 ; AJ : P. — ; V.h -

15—16’ 22—24’ 14—15^ 8

C. \x + 1 + 17 + 1 + x ; L. lat. 70—80; L. tr. 20 — 23; Vert. 57— 58*

a

b

Fig. 249. First left branchial arch of Clupea alosa. Natural size.
a: of a Twaite Shad (var. finta); b: in the Allice Shad (var. alosa ) shown in PI. XLIII, fig. 2, together with a gill-raker ( X 2)

from the middle of the arch.

Syn. Agon , Pucelle, Alose , Belon., Nat., Divers. Poiss., pp. 267,
302, 307. Thrissa , Rondel., De Pise. Mar., p. 220. Alosa,
vel Alausa, aut Thrissa, Schonev., Ichthyol. Slesv. Hols., p.
13. Shad, Willgghb., Hist. Pise., p. 227, tab. P, 3.
Clupea apice maxillae superioris bifido, maculis nigris utrinque,
Art. Ichthyol., Gen. Pise., p. 7; Syn., p. 15; Spec., p.
34 (ubi: “Apophyses in parte concava” (branehiarum) “ossese,
albae et satis robusta?” ; varietatem fintam igitur descripsit).
Clupea Alosa , Lin., Syst. Nat., ed. X, tom. I, p. 318 (ex
Art.; finta igitur); Br,., Fisch. Deustchl., part. I, p. 209,
tab. XXX, fig. 1; Retz., Fna Saee. Din., p. 353; Flmng,
Brit. Anim., p. 183; v. Yhlen, Fiskaren, 1878, p. 38;
Mob., Hcke, Fisch. Osts., p. 141.

Alausa vulgaris, Cuv., Val., Hist. Nat. Poiss., vol. XX, p.
391, tab. 604; Hckl, Kn., Sussioasserf. Ostreich. Mon., p.
228; v. Bemm. in Herklots, Bouwstoffen voor eene fauna van
Nederland, pt. Ill, p. 382; Canestrini (Alosa), Arch. Zool.,
l’Anat., cett., vol. IV, fasc. 1 (Apr. 1866), p. 141; Steind.
(Alausa), Stzber. Akad. Wiss. Wien, Math. Naturw. Cl.,

a A. 20 — 25, according to Day.

6 60 — 75, according to Day; 63 — 71, according to Fatio.
c 16 — 17, according to Kroyer; 16 — 19, according to Fatio;
d The latter number, according to Fatio.

4 — 5

e D. , according to Fatio.

14—17(19)’ * & * * 9

fA 3-_

20—24 ’ ” ” ”

9 Sometimes 13, ,, ,, ,,

h V. sometimes 10, ,, ,, ,, and Day.

* According to Fatio.

LVII, i (1868), p. 737; Canestr. (Alosa), Fna d'ltal.,
pt. Ill, p. 22.

Var. A.

Alosa ficta aut falsa, Ddh., Tr. Pech., pt. II, sect. Ill,
p. 320, tab. I, fig. 5; tab. II, fig. 1.

Twaite Shad, Penn., Brit. Zool. 1776, vol. Ill, p. 307.
Clupea fallax, Lacep., Hist. Nat. Poiss., vol. V, p. 452.
Clupea (Alosa) finta, Cuv., Regn. Anim., ed. 2, tom. II,
p. 320; Nilss., Prodr. Ichthyol. Scand., p. 22;
Yabr. (Alosa), Brit. Fish., ed. 1, vol. II, p. 131;
Schagerstr. (Clupea), Fysiogr. Sails. Tidskr., p.
291; Kr. (Alosa), Danin. Fisk., vol. Ill, p. 202;
Troschel (Alausa), Arch. Naturg., Jahrg. 18, Bd. 1,
p. 228; Nilss. (Alosa), Skand. Fna, Fisk., p. 527;
Sieb., Sussioasserf. Mitteleur., p. 332; Gthr ( Clu-
pea), Cat. Brit. Plus., Fish., vol. VII, p. 435; Coll.
(Alosa), Forh. Vid. Selsk. Chrnia 1874, Tillsegsh.,
p. 195; Cederstr., Ofvers. Vet. Akad. Forh. 1876,
No. 4, p. 67 ; Malm, Gbgs, Boh. Fna, p. 587 ;

15 — 16, according to Lilljeborg.

SHAD.

985

Winth., Naturh. Tidskr. Kbhvu, ser. 3, vol. XII
(1879), p. 50; Feddeks., ibid., p. 81; Mor., Hist.
Nat. Poiss. Fr., tom. Ill, p. 456; Bncke, Fiscli .,
Fischer ., Fischz. 0., TV. Preuss., p. 167 ; Day ( Clu -
pea), Fish. Gt. Brit., Irel., vol. II, p. 236, tab.
CXLI; Cole., N. Mag. Naturv. Chrnia, Bd. 29 (1884),
p. 113; Lillj., Sc., Norg. Fisk., vol. Ill, p. 120;
Fatio ( Alosa ), Fne Vert. Sitisse, vol. V, part. 2,
p. 40 ; Tryb. ( Clupea ), Sv. Fiskeritidskr., Arg. 1
(1892), p. 120.

Vav. B.

Clupea major. Alosa vera Autorum, Duh., 1. c., p.
316, tab. I, fig. 1.

Allice Shad, Penn., 1. c,

Clupea alosa, Lacep., 1. c., p. 447.

Clupea (Alosa) alosa, Cdv., 1. c., p. 319; Gthr, 1. c.,
p. 433: Day, 1. c., p. 234, tab. CXL; Coll., N.
Mag., 1. c., p. 112; Lillj., 1. c., p. 113.

Alosa communis, Yarr., 1. c., p. 136.

Alosa vulgaris, Sel. Longci-i., Fne beige, p. 220;
Trosch. (Alaitsa), 1. c. ; Sieb. (Alosa), 1. c., p. 328;
Winth., 1. c., p. 49; Mor., 1. e., p. 453; Storm,
Vid. Selsk. Skr. Trondhj. 1883, p. 31; Fatio, 1. c.,
p. 29.

Alosa Cuvierii, Malm, 1. c., p. 654.

C. Var. sapidissima, forma americana, pinna anali fintce, bran-
chiospinis alosce.

Clupea sapidissima, Wilson, in Ree’s, Cyclop , Univ.
Diet. Arts, Sc. Lit., p. 461 ; Jord., Gilb., Bull.
U. S. Nat. Mns. No. 16, p. 267; McDon. in Br.
Goode, Fisher., Fisher. Industr. U. S., sect. I, p.
594, tabb. 212 et 213.

Alosa proestabilis, Dek. N. York Fna, pt. IV, p. 255,
tab. XV, fig. 41.

In form of body the Shad resembles the common
Herring even more closely than the Pilchard does, the
greatest thickness seldom rising to 1/2 of the greatest
depth, and usually measuring between 35 and 45 %
thereof. The alterations of growth in this respect are,
however, considerable, the rule being that young Shads
are most like the Herring, and that in older specimens
the hind part of the body is prolongated to such an
extent as to reduce the greatest depth (across the
middle of the preabdominal part), which is commonly
between 23 and 28 % of the length of the body, to
about 22 % thereof. The body then acquires a com-
pressed, clavate shape, quite different from the hand-
some type of the Herring. The least depth varies be-
tween about 8 and 672 % of the length. The Shad is
also far more vigorous of growth than the Herring.
It almost ranks among the large fishes, though the
above-mentioned varieties, differ in this respect. The

T waite Shad seldom attains a length of 6 dm.“ or a
weight of 1 kilo., whereas the Allice Shad, it is stated,
often measures 7 dm. in length and weighs 272 kilo.,
and the North American Shad attains in modern times,
according to McDonald, a weight of at least 32/3 kilo.,
which it formerly exceeded, sometimes turning the
scale, it is said, at 3 1/3 kilo, or more.

The length of the head in Young Shads measures
about 23 1/2 % of that of the body. During growth to
a length of 4 or 5 dm. this percentage falls to 22
or 2 1 Va? and in still larger Shads to about 19, judging
by a stuffed specimen. To the covering of adipose
membrane on the head and shoulders the same remarks
apply as in the case of the Pilchard; nor do the lateral
ridges on the top of the head coalesce behind in the
occipital crest. In old specimens the tip of the snout
is incised, more than in the Pilchard and most in the
Twaite Shad, which has the deepest sinus at the head
of the angular or S-shaped indentation in the margin
of the intermaxillaries. In younger specimens and in
the Allice Shad the tip of the snout is more even,
showing only the above-mentioned sinus at the middle
of the margin. The eyes are about as large in young
Shads as in the common Herring, in old specimens
perceptibly smaller. In a Shad 12 cm. long both the ver-
tical and the horizontal diameters of the eyes are nearly
v4 of the length of the head, in males 3 — 4 dm. long
little more than 1/5 of the same, and in females of
this size the proportion is only about 18 %. The cor-
responding proportions to the length of the maxillaries
are respectively about 1/2 (50'7 %), 45 — 41 %, and about
43 — 39 %. In the smallest specimen the diameter of
the eyes is somewhat greater than the interorbital width
at the middle of the eyes, in old males about 90 — 80 %,
and in old females about 80 — 70% thereof. The length
of the snout is somewhat less in the first-mentioned
specimen than the diameter of the eyes, in the others
equal to or a little greater than the same. The nostrils
are set almost as in the Herring; but the distance be-
tween the two pairs is comparatively somewhat greater,
being in young specimens 1/2, in old about 2/3, of the
breadth of the snout at the articulation of the maxil-
laries. The length of the maxillaries is about 1 1 72
— 93/4 % of that of the body or 49 — 45 % of that of
the head, and these bones extend back to about a line

“ A stuffed specimen from Alexandria and in the possession of the Royal Museum measures about 61 cm. to the ends of the
caudal lobes.

986

SCANDINAVIAN FISHES.

with the posterior margin of the eyes, the length of
the cheek behind them being less than its height below
the eyes. The length of the lower jaw is 14 — 13 %
of that of the body" or about 62 — 58 % of that of the
head. The height of the dental part is less than in
the Herring, measuring about 1/3 of the length of the
lower jaw. At the symphysis the lower jaw is most
like that of the Pilchard and only slightly prominent;
but in old or lean specimens the symphysis is more
truncate (deeper), and has two pairs of comparatively
small chin-protuberances above. The jaw-teeth dis-
appear more or less completely with age, most in the
Allice Shad; but even in old specimens a fairly regular
row may be found on the intermaxi llaries, a tooth or
two in the front of the lower jaw, and traces of the
dentiform crenelation on the hind part of the maxil-
laries. There are no other teeth in the mouth. The
differences in the gill-rakers of the two varieties have
been mentioned above, and may be seen in the figures,
flie opercular apparatus differs from that of the Her-
ring in the greater extension of the preoperculum back-
wards and downwards and in a corresponding reduc-
tion of the operculum below, the obliquely set inferior
margin (the suture with the suboperculum), which is
fairly straight or incurved, measuring only about 1 7 1/2
— 21 % of the length of the head, 35 — 47 of that

of the maxillaries, or 29 — 35 % of that of the lower
jaw. An equally perceptible alteration of growth ap-
pears in the form of the suboperculum, which becomes
narrower, as a rule, during growth, its breadth at the
middle decreasing from about 60 to 40 % of the length
of the said suture. As the length of the body increas-
es from 12 to 40 cm., the postorbital length of the
head rises from about 45 '/2 to 58 % of the entire
length thereof, or from about 25 1/g to 30 % of the
distance between the dorsal tin and the articular knobs
of the maxillaries. When the postorbital length is
measured in an oblique direction (to the lower posterior

angle of the operculum), these percentages increase re-
spectively from about 52 V2 to 62 V2 and from 29 V2 to 32.

The length of the pectoral tins, which are oblique-
ly pointed, is about 16 — 14 %, that of the ventral
tins about 10V2 — 8V2 %■> of the length of the body.
The preabdominal length varies between about 22 and
26 %, the postabdominal between 21 and 24 Vs of
the length of the body.

The dorsal tin begins at a distance from the ar-
ticular knobs of the maxillaries measuring about 42 —
44V2 % °f the length of the body, or 90 — 99 % of the
distance between the ventral tins and the same points.
Its height decreases with age from about 15 to 11 %c,
and the length of its base varies between about 13
and 15 %>, of the length of the body, the latter propor-
tion seeming as a rule to be somewhat less in young
specimens than the former, greater in old. The caudal
tin is deeply forked, the middle rays measuring only
V4 — V 5 of the length of the longest ones, which varies
between about 2 2 x/2 and 19 % of that of the body.
The inferior lobe of the caudal tin is, as usual, some-
what longer than the superior.

The anal tin is of special interest from a syste-
matic point of view. The form is indeed highly vari-
able, but we have failed to discover any rule for its
variations. Sometimes the tin is comparatively high,
the length of its longest ray measuring as much as
2/5 of its base. In other specimens it is as low as in
the preceding Clupeoids, the height being but little
more than V4 of the base. But the extent of the tin,
the length of its base, seems to afford a distinction
both between the sexes and the varieties. In each
variety of the Shad, so far as our observations have
been carried, the females have a, comparatively longer
anal tin than the males; and the base of the anal tin
is comparatively longer in the Allice Shad than in the
Twaite Shad. We may here adduce the following ex-
amples of the rule:

“ It sinks in the oldest Shads to 12 % thereof, judging by a stuffed specimen.
b About 55 — 58 % in the Pilchard.
c Or even to 9 °/, judging by a stuffed specimen.

SHAD.

987

Clup

e a f i n t a.

C l u p

e a a

l o s a.

9

d

d

?

c?

9

9

d

9

Length of the body expressed in millimetres. -

121

284

302

348

393

147

147

321

358

385

Base of the anal tin in % of the length of the body

14.7

16.o

13.9

14.6

16.o

16.0

18.i

17.o

17.6

17.i

„ „ ,, „ „ „ » „ „ ,, „ ,, head

62.8

71.5

60.8

64.5

72.o

61.3

75.8

85.i

74.i

79.5

„ „ ,, „ „ „ ,, „ „ distance between the anal tin and the tip of the snout

21.8

24.3

20.7

21.i

24.0

22.6

27.2

29.6

26.9

26.i

,, ,, „ ,, postabdominal length

63.6

75.8

56.7

61.8

72.4

70.3

86.4

87.6

80. o

79.6

„ ,, dorsal fin in % of the base of the anal fin

93.8

80.2

105.9

92.7

91.7

80. o

74.5

74.2

79.3

75.7

Length of the maxillaries ,, „ „ „ ,,

78.6

68.7

80.9

71.5

65.i

71.i

63.6

53.8

63.5

56.5

„ ,. „ lower jaw — „ „ „ „ „ „ „ >,

95.5

86.8

98,s

90.8

84.i

93.3

84.2

68.5

74.6

75.3

„ ., „ pectoral tins ,, ,, ,, ,, ,,

109.4

96.7

102.3

99.c

90.5

103.5

90. o

75.o

84.i

80.3

.. ,, ventral fins , „ „ ,, ,,

72.4

61.5

68.i

63.7

54.7

60. o

58.o

45.o

52.4

50. o

Least depth of the tail ,, ,, ,, ,, ,, ,,

57.8

46.i

57.i

52.o

48.5

53.3

46.8

39.9

42.8

38.8

The rule is clear: the young and the males have
the shortest anal fin, and the Allice Shad is charac-
terized by the long base of its anal fin. But in this
character the young male of the Allice Shad — a Medi-
terranean specimen 147 mm. long and sent here by
Hedenblad — ranks beside the female T waite Shads,
and is so closely approximated to them that the dif-
ference is useless as a specific character. The character
is, however, not without importance, for we find on
comparison that in most of the relations the base
of the anal fin is much shorter in male Twaite
Shads than in Allice Shads less than half as long.
A development of form, determined by the female
characters, has evidently advanced from fintaa to alosa;
but this development has not yet attained such defini-
teness that true species can be distinguished. The spe-
cific character first pointed out by Troschel, the dif-
ference in the number of the gill-rakers — greater in
alosa (fig. 249, h) than in finta (fig. 249, a) ■ — is im-
paired by the same indefiniteness. Steindachner and
Moreau showed that their number increases with age,
whereupon Lilljeborg remarked with justice that the
character can only be employed in comparisons between
specimens of about equal size. The case appears to be
the same here as in the Gwyniads (see above), where
we also failed to arrive at more than variety distinc-
tions by the aid of the differences in the gill-rakers.
And even if we could prove that the character derived
from this source agrees in our European Shads with
other characters6, the North American Shad apparently
baffles our systematic calculations, for it is fur-

nished, according to Jordan and Gilbert, with the
gill-rakers of the Allice Shad, but externally resembles,
according to Todd’s figure in Brown-Goode (1. c.), the
Twaite Shad.

The form and texture of the scales (fig. 245, b)
we have described above. They are more firmly at-
tached to the skin in most cases than in the Herring.
The irregularity of their arrangement and the absence
of a lateral line render it difficult to compute their
number with certainty. The number of spiniferous
scales at the ventral margin is 37° in all the specimens
examined by us, 23 or 22 in front of the ventral fins
and 14 or 15 behind them. Elere too these scales are
covered to the greater part of their extent by the
nearest lateral scales.

The coloration is essentially the same as that of
the Herring. The dark (blackish blue) spots charac-
teristic of the subgenus Alosa, set in a straight or an
interrupted (zigzag) row on the sides of the body, the
largest — sometimes as large as the eyes, but commonly
smaller — first, are most usual in young specimens.
They formerly ranked as a character of the Twaite
Shad, but are of very irregular occurrence even in this
variety. Our figure (Plate XLIII, fig. 2) also shows
that they may be quite numerous in the Allice Shad;
and Fatio relates that, on boiling adult Allice Shads
from the Rhine in order to prepare their skeletons, as
many as 15 spots appeared on the sides of the body.
“On the sides of the head too,’’ writes Kroyer of the
Twaite Shad, “a black spot may sometimes be seen
above the preoperculum”; and in one of our Twaite

a Or perhaps from the North American Clupea chrysochloris , with its still shorter anal fin, unless this form be an instance of retro-
gression in development.

* Lilljeborg has adduced the different form of the tip of the snout and the greater number of the scales in alosa (see above).

0 37 — 42, according to Valenciennes.

988

SCANDINAVIAN FISHES.

Shads this spot is the only one left, with the exception
of that on the shoulder. Nor is the black colour of the
tip of the snout, the point of the lower jaw, and the
tongue constant. But the scale follicles of the whole
body as well as of the head are strewn, more densely
than in the Herring, with tine dots of brownish black ;
and at the top of the dorsal tin this colour gathers in
a black spot, the hind margin of the caudal tin being
also black.

If we include, as we have reason to do, within the
limits of this species the North American Shad (and
perhaps the Skipjack, Clupea chrysochloris), the species
has an extensive range on both sides of the Atlantic.
On the eastern side it occurs from the neighbourhood
of Trondhjem" to the Mediterranean; on the western
side it lives somewhat further south, being found from
the Gulf of St. Lawrence to St. John’s River, Florida* * * 6.
In the Pacific it was unknown until the 2nd of July,
1873. On that date Livingston-Stone, instructed by
the Fisheries Commission of the United States, stocked
the Sacramento at Bahama (California) with 35,000
Shad-fry from the Hudson River, delivered to him on
the 25th of June from the Shad Hatchery of New York
State at Castleton. This proved one of the greatest
triumphs achieved by pisciculture; a productive Shad-
fishery was bestowed upon California. The enterprise
is also of general interest, as calculated to throw light
on the wanderings of other fishes0. We are told by
McDonald: “It is a common belief among fish-cultur-
ists that the mature individuals of all anadromous spe-
cies, including the Shad, are led back to the waters in
which they were spawned by a conscious wish on their
part to return to those very localities in which they spent
their young life. Important exceptions to this rule are,
however, well established by recent observations. For
instance, it is well established that the runs of Shad into
the Susquehanna and Potomac Rivers are characterized
by alternations of abundance; that is to say, an exces-
sively large yield for any given season in the one in-
volves a corresponding diminution in the yield for the
same season in the other, thus precluding the possibility
of each individual returning annually to its native stream.

° Even on the coast of Iceland the Shad is found, according

him that on the east coast of Iceland a kind of Herring was taken,

They called it Ugna-sild, i. e. Herring with eye-shaped spots.

6 The Skipjack strictly belongs to the Mississippi Valley and
the true Shad is indigenous to the Alabama, where attempts to introd
above all in the Ohio, it has been planted by the U. S. Fisheries Cor

c Cf. above, on the migrations of the Salmon, pp. 858 — 859.

Again, it was confidently expected that all the young
Atlantic Shad which were transferred to and planted in
the Sacramento River would, on their return from the
Pacific Ocean as mature fish, find their way back to
this stream. This tvas not, however, the case, for, to
the utter astonishment of many fish-culturists, a con-
siderable number of these now mature fish made their
appearance in many streams of the Pacific lying far
north of the Sacramento River — streams to which Shad
had never been indigenous and in which none had ever
been planted. These facts go a long way to disprove
the theory of instinct of locality, and indicate that the
river movements of the Shad are regulated by involun-
tary and extraneous influences. The migration and colo-
nisation of this fish northward along the Pacific coast
has been so general that at the present day new genera-
tions of a single plant are found in every stream on
the Pacific from the Sacramento River to Puget Sound.”
In Scandinavia the Shad may be considered as a
stranger. North of Germany it does not appear any-
where in numbers. But it is not rare. Several large
Allice Shads have been taken during the summer months,
according to Storm, in Salmon-nets by the fishermen ol
Trondhjem Fjord, and Collett states that an Allice Shad
was caught in the autumn of 1881 off Namsas (64V2° N.
lab). The finta variety has not been observed so often
in Norwegian waters, and, according to Collett, only
in Christiania Fjord and near Bergen. In Denmark the
Shad is fairly common, but almost without exception in
the finta form. This variety was frequently observed
by Krgyer among the fish taken on the Danish coast
of the Cattegat, and he was told by fishermen that they
sometimes caught hundreds of Twaite Shads in Liim
Fjord. But only three Allice Shads, all taken during
May (1871 and 1878) in Liim Fjord, are mentioned by
Winther. On the west coast of Sweden the case is
about the same. The Royal Museum has received from
Stromstad, through Mr. C. A. Hansson, two females of
the Allice Shad, one of them measuring about 44 cm.,
the other 36 cm., and taken respectively at the begin-
ning of June, 1887 (the-original of our coloured figure)
and the beginning of September, 1892. No other Allice

to a statement in Faber ( Fische Islands, p. 182). The fishermen told
similar to the common Herring, but with black spots along the sides

the neighbouring parts of the Gulf of Mexico. It is uncertain whether

uce it have been made since 1848. In the tributaries of the Mississippi,

omission.

SHAD.

989

Shads are known from the west coast of Sweden. The
Twaite Shad is commoner, and is taken off the coast of
Bohnslan, according to Malm, “now and then, most fre-
quently on Haddock-lines, both in spring and autumn,
but usually in October and November.” At the be-
ginning of July, 1832, according to Nilsson, about a
score of Twaite Shads were taken in Flounder-nets off
Mahno (Bohnslan). On the Halland coast the Twaite
Shad is fairly common, at least in Laholm Bay, where
it is called BlanJcsill and Stafsill, and in summer it
ascends to the lower part of the River Laga. It is found
quite often, according to Schagerstrom, in Schelder Bay
at the foot of the Kulla Cliffs. In the Sound it is rather
rare, but commoner than in the Belts (Winther). On
the Prussian coast, according to Benecke, it formerly
entered the Haffs in great numbers, but is iioav rare,
being taken only on few occasions among the hauls of
Stromming. Nilsson states that in April, 1850, the
fishermen of Abekas (Southern Scania, west of Ystad)
caught, partly in gill-nets set for other fish, partly in
Salmon-seines, 8 or 10 Twaite Shads. From Calmar
Sound off Borgholm Dr. Areen sent to the Royal Mu-
seum a male Twaite Shad about 3 dm. long that had
been taken in a trap on the 13th of October, 1892.
The Twaite Shad thus occurs in the Swedish part of
the Baltic, but the specimens are probably solitary ro-
vers; and this is, no doubt, the case with the Allice
Shad too, which has been found still higher up. The
Museum of Upsala contains a female Allice Shad 57
cm. long that was taken on the 6th July, 1864, in the
Dal Elf off Elfkarleby (Lilljeborg).

No Shad-fishery of importance can therefore be
carried on in Sweden, unless successful attempts be made
to plant this fish in suitable places within Scandinavia.
Things are different in the Shad’s true home. In the Old
World its habitat is of about the same extent as the
Pilchard’s, and there the Shad is one of the most im-
portant fishes from an economical point of view, its
manner of life reminding us strongly of the Salmons.

The Shad commonly passes the greater part of its
existence in the sea; and its life there is little known,
but it probably roves in scattered companies to seek its
food. This consists not only of small Evertebrates and
lower algm, but also of moderate-sized fishes. In the
stomach of a Twaite Shad 47 cm. long Collett found
about a score of Sprats 65 — 68 mm. in length.

When its propagative instinct begins to awake, or
even some time beforehand — this depending seemingly
on the temperature of the water — the Shad musters in
shoals to commence its migration to fresh water. Ac-
cording to McDonald the American Shad ascends the
St. John’s River (Florida) as soon as the temperature of
the water has fallen in autumn to about 60° Fain-.,
which happens at the end of November; but the spawn-
ing is not in full progress till the beginning of April.
In the Potomac, on the other hand, the Shad presses
on with the greatest eagerness when the temperature of
the river, at about the middle of April, is 56° Fa.hr.
Large and small, ripe and unripe, the fish make their
approach, but only the former continue their advance,
until they find a temperature of 65° — 70° Fahr., where-
upon the spawning is begun, the Shads not yet in
breeding condition halting at a temperature of 60° Fahr.
The further north the mouth of the river lies, the later
is the appearance of the Shad. This seems also to be
the case in Europe. According to Blanchere0 Shads
are taken in French rivers from March till July, and
according to Valenciennes* 6 the fish appears earlier in
the Loire than in the Seine. According to van Bem-
melen (1. c.) the Shad ( alosa first) ascends most of the
Dutch rivers in April and May. It is a rule too that
the Twaite Shad arrives about a month after the Allice
Shad. In England, according to Yarrell, the two
varieties have a preference for different rivers: in the
Severn the Allice Shad preponderates, in the Thames
the Twaite Shad. In Ireland both varieties are fairly
common, in Scotland somewhat rare.

At the commencement of its sojourn in fresh water
the Shad is in its best condition, and it retains its fatness
until the spawning begins. For the purpose of spawning
it roves, like the Salmon, but with less vigour, as far
as it can penetrate up the rivers and their tributaries, in
the Rhine up to Basel, in the Elbe into Bohemia, and
in the Loire into Haute Loire, a distance of more than
800 kilom. (Mon.). During the early part of its ascent
it is as timid as the Herring and cautious, retreating
at the approach of a thunderstorm or a. spring freshet ;
but as the sexual instinct gains sway, the fish lav
aside all fear, and “grumble and grunt, like a herd of
swine” (Baldner, in Willughby). Meanwhile, as the
males and females press together and thrash the sur-
face with their tails, the eggs are deposited and im-

" Nouv. Diet. Gen. Peches, p. 16.

6 Guv.. Val., 1. c., p. 412.

990

SCANDINAVIAN FISHES.

pregnated. When the spawning is over, the Shad’s
powers are exhausted, and its value lost. Many perish,
others are carried half-dead by the current back to the
sea; and by the end of the summer all the full-grown
Shads have usually quitted the rivers and streams.

Each adult female lays, it is stated, 100,000 —
200,000 eggs" l* * 6 or 2C mm. in diameter, which sink
to the bottom, and are hatched in a few days. In
October Yarrell obtained young T waite Shads 21/2 in.
(63 mm.) long; and in the following spring he found
young T waite Shads 4 in. (1 dm.) long and young
Allice Shads 6 in. (1\2 dm.) in length''. Where the
Shad spends the intervals of its existence, and how it
lives in the sea, until it attains a length of about 3—
4 dm., and appears after 2 or 3 years in fresh water
as an adult fish, is unknown to us.

As the Salmons occur in fresh-water forms that
never reach the sea, so the Shad is sometimes confined

for its whole life in lakes and their feeders. To this
category belong, according to Fatio, the so-called Agoni,
a kind of Twaite Shad, that inhabit the great lakes of
Lombardy and South-eastern Switzerland. The large
Twaite Shads, known as Cheppie, ascend the Po from
the Adriatic, and spawn during June and July in this
river and its tributaries, returning in August to the
sea. The Agoni, on the other hand, though they attain,
according to De Betta, a length of 3 — 4 dm. and
according to Fatio, a weight of 1 V2 lbs., pass the
whole year in the lakes of Ticino and Lombardy, where
they spawn in May and June.

In its best condition, after living a day or two
in fresh water, the Shad, especially the fat Allice Shad,
is said to be excellent eating, in spite of the numer-
ous bones, which are more troublesome even than in
the Herring. After the spawning the flesh is lean
and dry.

Genus STOLEPHORUS.

Mouth targe, the length of the maxillaries being more than half that of the head. Tip of the snout projecting
more or less distinctly beyond the lower jaw. Gill-openings large, the branchiostegal membranes being for the
greater part of their extent free from each other and from the isthmus. Anal fin free from the caudal.

Ranged beside the preceding genus the European
Anchovy may indeed appear to be so sharply distin-
guished therefrom that it might lay just claim to a
place at least in a separate subfamily. But on a com-
parison with the other genera of the family the dif-
ferences disappear, and even the genus Stolephorus is
difficult to define with natural limits. Valenciennes6
pointed out this circumstance, and restored to the ge-
nus those species which Cuvier7, on account of the
backward prolongation of the maxillaries (extending
behind the articulation of the lower jaw and even be-
hind the head), had removed to a separate genus Thrissa.

“ In American Shads weighing 4 — 5 lbs. McDonald found

number sometimes exceeds 100,000.

6 Benecke.
c Fatio.

d Of., however, Ehkenbaum (Sonderbeilage zu den Mittheilungen der Sektion fur Kirsten- und Hochseefischerei, JahrgaDg 1892, p. 12),
who is of opinion that young Twaite Shads of the said size are a year older, and have already paid one visit to the sea.
e Cuv., Vai„, Hist. Nut. Poiss., vol. XXI, p. 5.
f Regne Animal, ed. I, tome II, p. 176.
g Nat. Hist. Fish., Amph., Rept., vol. II, p. 292.

* Gunther’s Telara and Heterothrissa, Cat. Brit. Mus., Fish., vol. VII, p. 385.

Among the Anchovy forms typical in other respects we
find species in which the hind extremities of the ma-
xillaries are very obliquely truncate, with the lower
corner longer than the upper and even pointed — as
for example in Stolephorus heterolobus from the Red
Sea and India — so that the generic character depended
merely on a greater or less degree of prolongation.
To the genus Coilia, which in form of body calls to
mind the Macruroids, we find among the Anchovies
transition forms that Swainson5' proposed to isolate in
a genus Setipinna’1, characterized by the long anal fin,
which sometimes occupies more than half the length

only 20,000 — 40,000 ova; but lie adds that in other females the

ANCHOVIES.

991

of the ventral margin, and the filamentous elongation
of the first pectoral ray, a peculiarity that extends to
still more of the pectoral rays in Coilia. Bleeker"
indeed attempted, following a suggestion of Gunther’s6,
to find a limitation for such a genus in the number
of rays in the anal fin, more than 50. But a com-
parison0 between the two Indian species Stolephorus
( Setipinna ) telara and St. purava, the latter with 38 —
47 (or, according to Gunther, 50) anal rays and
pointed pectoral fins, is enough to show how nearly
the said characters run into each other. Another ge-
neric character recognised by Bleeker is the peculiarity
first observed by Gunther in some Anchovies'6, and
consisting in a more vigorous development of the jatv-
teeth, among which some are prominent as canines.
But in other respects these species resemble those with
small teeth. Last!}' Bleeker has directed attention to
the varying development of the spiniferous scales at

a Atl. Ichthyol. Lid. Or. Ne'erl., tome VI, p. 135.
b Cat., 1. c.

c See, for example, Day, Fish. Ind., tab. CLVIII, fig. 2
d Gunther’s subgenera Lycengraulis and Lycothrissa.
e Hist. Nat. Poiss., vol. V, p. 381. From orohrj, equipment
the sides of some species.

f Cuv., Reyn. Anim., ed. I, tom. II, p. 174.

the ventral margin. In conjunction with a more elon-
gated and terete form of body, these spiniferous scales
disappear in some forms of the genus, either in the
postabdominal region alone or throughout a great por-
tion of the preabdominal as well. These forms are
also characterized by the shortest anal fin, with at most
23 rays, and Bleeker proposed to reserve the generic
name of Stolephorus , coined by Lacepede0, for them
alone. But in these respects too there are intermediate
forms, and even in the most typical of the last-men-
tioned species — as for example in the European An-
chovy — traces of spiniferous scales may be found at
the preabdominal margin. The genus has been most
generally known under the later name of EngraulisS
and about 50 species have been described from tem-
perate and tropical regions all round the globe, most
of them purely salt-water fishes, but some visitors to
the rivers or at least to the estuaries.

and tab. CLVII, fig. 2 {purava).

, adornment , and qiegco , bear; with reference to the silvery band along

Scandinavian Fishes.

125

992

SCANDINAVIAN FISHES.

THE ANCHOVY.

STOLEPHORUS ENCRASICHOLUS.

Fig. 250.

MaxiUaries rounded at the hind extremity , and not extending behind the articulation of the lower jaw, but about
equal in length to the base of the anal fin ( somewhat longer or shorter ), which is at most about 2/3“ of the length
of the head, or 7 10 of the postabdominal length, or 85 % of the length of the peduncle of the tail, which is
greater than the length of the lower jaw. Dentition feeble and without canines, but complete. Greatest thickness
of the body about 2/3 (63 — 70 %) of its greatest depth, which measures about 14 — 17 % of its length. Least
depth of the tail about 3/10 (29 — 31 %) of the length of the head. No spinif erous scales, or only vestiges thereof,
at the ventral margin. No sharply defined, silvery band embedded in the skin along the sides of the body.

Fig. 250. 5 'tolephorus enerasicholus , cp- Natural size. Taken at Striimstad in 1878, C. A. Hansson.

R. br. 12—13; D. 3 - ; .4. - ;

13—15 15*— 16

C. #+1 + 17 4- 1+<j?; L. lat. 44 — 50 ; L. tv.

P. ; V.

15—16 6

7; Vert. 46—47.

Syn. Halecula, Belon, Nat., Divers. Poiss., p. 167. Enerasicholus,
Bond., De Pise., p. 211. Lycostomus balticus, Schonev.,
Ichthyol. Slesv. Hols., p. 46, tab. V (fig. inf.). Clupea
maxilla superiore longiore, Art., Ichthyol., Gen. Pise., p. 7 ;
Syn. Pise., p. 17. Anchois et Melet, Duham., Pitches, pt.
II, sect. Ill, p. 457, tab. XVII, fig. 5; sect. VI, p. 157,
tab. Ill, fig. 5.

Clupea Encrasicolus, Lin., Syst. Nat., ed. X, tom. I, p. 318;
Bl., Naturg. Fische Deutschl., pt. 1, p. 212, tab. XXX,
fig. 2; Retz., Fn. Suec. Liu., p. 354; Pall., Zoogr. Ross.
Asiat., tom. Ill, p. 212; Cuv., (subg. Engraulis ), Reyn.
Anim., ed. I, tom. II, p. 175; Risso ( Engraulis ), Eur. Mer.,
tom. Ill, p. 454; Cuv., Val., Hist. Nat. Poiss., vol. XXI,

P- 7,

tab. 607 ; Kr., Damn.

Fisk.,

vol. Ill

, p.

221 ;

Nilss.,

Slcand. Fn., Fisk., p.

531: v.

Bemm.

in Herkl.,

Boniest

. Fn. Nederl., pt. Ill,

p. 383;

G-thr,

Cat. Brit.

M/is.,

Fish., vol. VII, p. 385

; Coll.,

Forh.

Vid.

Seb k.

Cbrnia

1874, Tilleegsh., p. 194; 1879

, No.

1, p.

. 99;

N. Mag. Naturv. Cbrnia, Bd. 29 (1884), p. 112; Malm,

Gbgs, Boh. Fn., p. 569; Wxnth., Naturh. Tidskr. Kbhvn,
ser. 3, vol. XII. p. 47 ; Mor., Hist. Nat. Poiss. Fr., tom.
Ill, p. 460; Day, Fish. Gt., Brit., Irel., vol. II, p. 206,
tab. 138, fig. 1: Mob., IIcke, Fisch. Osts., p. 142; Peter-
sen, Vid. Meddel. Naturh. For. Kbhvn 1884, p. 159; Lillj.,
Sv., Norg. Fish., vol. Ill, p. 33; Ehrenb., Sonderbeil.
Mittheil. Sekt. Ktist., Ilochseef., Jahrg. 1892 (D. Fisch. Ver.).

Engraulis meletta, Cuv., R'egn. Anim., edit. 2, tom. II, p. 323
(ex Duham.).

Engraulis vulgaris, Nilss., Proclr. Ichthyol. Scancl., p. 25.

The Anchovy sometimes attains a length of 2 dm.c,
but is commonly no more than 12 — 15 cm. long. As
we have already mentioned, it is characterized by an
elongated body, fairly terete both at the dorsal margin
and the ventral, and of a handsome and regular, fusi-
form shape, pointed in front. Only the head is com-
pressed below, its sides converging towards each other.
The pointed snout is one of the most distinctive char-
acters belonging to the Anchovy and its nearest con-

a 65 — 6 0 1 /., % in our specimens.
b Sometimes 14, according to Day.
c 20 1 2 cm., according to Ehrenbaum.

ANCHOVY.

993

geners. The ethmoidal region is prolongated in front
of the mouth and above the intermaxillaries, which
are even more reduced than in the preceding Clu-
peoid forms.

The head in its remaining form and structure
lends itself most readily to a comparison with that of
the Greater Scopelus ( Myctophum elongatum). This
resemblance may be seen in the large, horizontal cleft
of the mouth, with the gill-slits extending far forward
in the lower jaw, the very strongly constricted cheek-
region below the eyes, and the backward and down-
ward prolongation of the opercular apparatus, the pre-
operculum having lost the lower, horizontal arm that else
projects in a forward direction. The cheek behind the
eye is also of the same triangular form as in the said
Scopeloid.

In Anchovies 13 — 15 cm. long the length of the
head occupies about 21 V2 — 2278 % °f that, of the body.
So great, however, is the extension of the pharyngeal
and opercular apparatus that the head itself, from the
tip of the snout to the occiput, measures only about
V7 of the above length. The longitudinal diameter of
the eyes, which are round, but incline to an oval shape,
is between 24 and 2 11/2 %, their vertical diameter about
20 %, of the length of the head. The eyes and the
sides of the snout in front of them are covered by an
adipose membrane, which is quite continuous, without
aperture or eyelid, but transparent on the pupil. The
interorbital width at the middle of the eyes is slightly
greater than their vertical diameter, but less than the
longitudinal. The length of the snout is slightly less
than the said vertical diameter, the difference being
least in old specimens. The nostrils on each side are
set close together, being divided only by a narrow
strip of skin, which may, however, be elevated so as
to form an obliquely cut groove, open in front. They
are set at the edge between the flat or faintly convex
top of the snout and its converging sides, about half-
way between the tip of the snout and the anterior
margin of the eyes or a little nearer to the latter.
The small, stiletto-shaped, and curved intermaxillaries
are attached to the lower anterior margin of the long
maxillaries. The latter bones increase uniformly, but
slightly, in breadth behind, where each of them is fur-
nished with two narrow jugal bones, pointed in front,

a On the sides of the head of the vomer, on the palatines,
ectopterygoids.

their greatest breadth, including the jugal bones, being-
only about 7 10 of their length. The lower jaw is also
shallow and long, in contradistinction to that of the
preceding Clupeoids; it is slightly turned up at the
tip. The length of the maxillaries is about 61 — -65 %,
that of the lower jaw about 68 — 71 %, of the length of
the head. The intermaxillaries, maxillaries, and lower
jaw are each furnished with a row of small pointed
teeth, somewhat curved and of uniform size. Similar
teeth may be found on almost all the bones of the pa-
late", but are more or less deciduous. In old specimens
the teeth of the maxillaries, and sometimes, at least in
part, those of the lower jaw, also disappear. The tongue
is small, cartilaginous, and toothless, but free. Close
to its base begins the fine denticulation of the long and
narrow row of copular bones. Both the upper and
lower pharyngeals are armed with small teeth. The
gill-rakers are long and fine, numbering about 62 — 72
on the first branchial arch.

The top of the head, which is almost flat, is coursed
at the middle by a longitudinal ridge from the very tip
of the snout (the upper margin of the ethmoid bone)
to the occiput; and from the inner side of each pair of
nostrils, obliquely outwards to each supraorbital mar-
gin, and thence obliquely inwards to the occiput, there
run the same ridges as in the Clupeoid forms im-
mediately preceding the Anchovy. The cephalic sy-
stem of the lateral line is generally less developed than
in the said forms, but here too varies considerably in
its development. In some instances only faint traces
thereof appear; in others the whole top of the head,
the temples, the upper parts of the scapular region, and
the triangular cheeks are covered with numerous, ver-
rucose elevations and pores, sometimes with a network
of confluent ducts belonging to this system, and the
hind margins of the operculum, the preoperculum, and
the scales on the scapular region are pierced by the
straight, backward ramifications of these ducts. The
preoperculum is crescent-shaped, though the lower cor-
ner is somewhat prolongated, forming a thin, rounded
lobe. The operculum is quadrangular and of uniform
breadth, but curved like a sickle, its upper anterior
margin, above the articular head, being perpendicular,
and its whole posterior margin forming a fairly regular
curve of about a quarter of a circle. Its lower posterior

on the entopterygoids. and, in a small card, on the hind part of the

994

SCANDINAVIAN FISHES.

margin shows an arcuate indentation. Obliquely across
its surface, from the articular head to the lower pos-
terior corner, runs a groove, in front of which the
lower anterior, triangular part of its surface is some-
what tumid, but smooth. The suboperculum has a
rounded triangular form, with the hind margin sharply
curved. The length of its upper margin, which coincides
with the breadth of the operculum, is about 10 — 12 %
of the length of the head. The interoperculum is of
about the same shape as the suboperculum, but much
smaller. All the opercular bones are very thin. The
branchiostegal membranes are narrow, hardly perceptible
at the hind margins of the operculum and suboperculum.
In front (below) they follow the branches of the lower
jaw, their breadth being but slightly greater than that
of these bones or, at the extreme front, less. The
branchiostegal rays are also comparatively short, gradu-
ally increasing behind (with the exception of the last
one) in length and breadth. The anterior are subulate
and curved, the posterior of more and more uniform
breadth and sabre-shaped, with obliquely set, terete shaft;

' 1 1 i ii AuL.’A*'

Fig. 251. Composite scale from the left side of the inferior caudal
lobe in an Anchovy 16 cm. long. X 3.

the longest ones (in the inferior lobe). On each side
of the fin are set, as in the Clupeoid forms immediately
preceding the Anchovy, a pair of so-called composite
scales — much smaller, however, than the largest body-
scales — one on the superior lobe, the other on the in-
ferior, and each consisting of a scalpel-shaped dermal
flap, with diamond-like reticulation, and covered with
a considerable number (20 — 25) of small scales. These
are set in imbricated rows, and each of them is marked
with a row of brownish black pigment-spots (fig. 251).

The pectoral fins are comparatively short, their
length being about equal to the height of the dorsal fin
or a little greater, and are obliquely truncate, their shape
reminding us strongly of the pectoral fins in those An-
chovies where the first ray of these fins sIioavs a fila-
mentous elongation. The ventral fins are also obliquely
truncate, but much shorter than the pectoral, their length
being only about 7 72 — 7 % of that of the body. The
position of the ventral fins is such that the distance be-
tween them and the tip of the snout measures about

but the hindmost is of a rounded triangular shape, re-
sembling in form and position a lower suboperculum.
The nine middle rays are set on the ceratohyoid bone,
the last two on the epihyoid.

The distance between the dorsal fin and the tip of
the snout is about 44 (4472) — 48 %, its height as well
as its base about 10 — 11 %, of the length of the body.
The first ray is extremely small, the second about 73 as
long as the third (the last simple ray), which is slightly
shorter than the first branched ray. The last ray in
the fin is scarcely elongated. The distance between the
anal fin and the tip of the snout is about 60— 63 72 0/oi its
base about 14 %, and its height (the length of the first
branched ray) about 6x/2 — 7 % , of the length of the
body. It is thus both longer and lower than the dorsal
fin, and has a more concave outer margin, the la,st ray
being somewhat elongated. The first of the two simple
rays at the beginning of the fin is hardly 73 as long
as the second, which is almost equal in length to the
first branched ray. The caudal fin is deeply forked,
the length of the middle rays (about 5 — 57 2 % of the
length of the body) being even less than 73 of that of

Fig. 252. A scale from the left side of an Anchovy 16 cm. long.

Three times the natural size.

40 — 44 % of the length of the body, or 90 — 92 % of
that between the dorsal fin and the same point. The
preabdominal length is about 18 — 22 %>, and the post-
abdominal about 20 %, of the length of the body.

The scales are extremely thin and transparent, ra-
ther large, but short, and of a quadrangular shape,
rounded at the corners (fig. 252). Their posterior (free)
part is as large as the anterior (covered) portion, and
is also finely striated, the concentric striae being even
twice as numerous as on the latter. The radiating
grooves are sharply defined, originating both at the an-
terior margin and the posterior. The scales of the pec-
toral fins, and the pointed axillar scales of the ventral
fins are long, extending respectively 4/5 and 3/4 of the
distance along the fins. The basal scales of the dorsal
and anal fins are about the same as in the Shads. The
median line of the belly is covered, it is true, with
ordinary scales; but vestiges of the spiniferous scales
that appear in the preceding Clupeoid forms may some-
times be found. These take the shape of linear thicken-

ANCHOVY.

995

ings of the skin in the preabdominal region, answering
to the lateral processes of the spiniferous scales; and
outside (below) each pelvic bone, just in front of the
outermost (first) ray of the ventral tins, lies an extreme-
ly thin, squamoid covering-bone, which sends out a
wand-shaped process in the skin, extending about V3 of
the distance up the sides of the body, and corresponding
to the ordinary double process that rises at this point
in the Herrings (cf. p. 958, tig. 240, c).

The coloration is essentially the same as in the
Herrings, but is characterized by a steely longitudinal
band on the upper part of the sides, forming a sharp
line of demarcation between the back, which gleams
chiefly with a greenish lustre, and the lower parts of
the sides, which are silvery white. On the articular head
of the operculum lies a dark, elliptical spot, the length
of which is scarcely half the diameter of the eyes.

The Anchovy’s range in the Atlantic coincides Avith
that of the Pilchard". Its true habitat lies in the Mediter-
ranean and the neighbouring part of the Atlantic, and
hardly extends north of England and Holland, though
numerous specimens have been taken in Christiania
Fjord, and solitary individuals occur along the Nonve-
gian coast up to Bergen. It also roves into the Cat-
tegat and the Baltic, probably in the train of the Her-
ring and Stromming, up to the island-belt of Stockholm,
where Professor Hjalmar Holmgren took a female An-
chovy about 18 cm. long* * 6 on the 15th of August, 1869.
In the south-west of the Baltic it is caught somewhat
more frequently — “though rare”, Avrote Schonevelde
in 1624 — off Kiel (Mobius and Heincke) and Trave-
mttnde (Lenz). On the north coast of Funen Feddersen
obtained 7 specimens in the middle of June, and Pe-
tersen caught 4 at the end of the year 1885. On the
north coast of Zealand Kroyer saAv two Anchovies. In
the Sound north of Saltholm Winther secured a speci-
men 16 cm. long in June, 1870. The Royal Museum
has received from Baron Gyllenstjerna a specimen
17 cm. long, taken off Kullen in August, 1829. Burman
sent two specimens that had been caught in Herring-
nets off Stromstad in July, 1862, and the Museum has
more recently received through Mr. C. A. Hansson tAvo
more from the same locality. Malm mentions 4 speci-

mens from Bohusl&n. The Anchovy, though rare, is
thus not to be reckoned among the rarest fishes of the
Cattegat and Skager Rack. In Christiania Fjord it is
so common, according to Collett, that it appears every
summer, at least in small numbers, at the fish-market
of Christiania; and in August, 1873, Collett met Avith
a considerable number, both large and small, in the
fjord. In October, 1875, he found several specimens
among common Herrings off Christiania, one of them
nearly 19 cm. long, another only 68 mm., Avhence he
infers that the Anchovy spawns in those waters. The
adequacy of this evidence may, however, be questioned,
if Ehrenbaum be correct in his opinion that Anchovies
of the latter length may be in their second year and
have passed one Avinter in the sea.

That the Anchovy is outside the bounds of its true
habitat, or at the northern limit thereof, on the coast
of Holland — though it spawns there, and is sometimes
extremely plentiful — appears from the great irregularity
to be observed in its occurrence. Van Bemmelen re-
lates, on Martinet’s authority0, that in 1765 or 1766
fifty fishermen could not secure more than three An-
chovies during the Avhole summer in the Zuyder Zee,
though some years before the Zee had been “so full of
Anchovies as to be almost unnavigable for smacks.”
Plven so far south as in the English Channel it is far
less common, according to Moreau, than in the Medi-
terranean.

From the large gape of the Anchovy and its other
resemblances to the Greater Scopelus avc may conjecture
that its manner of life is also much the same, and that
it is strictly a pelagic fish-of-prey. But in quest of
food and in order to spaAvn it approaches the coast,
and for the former purpose it enters brackish or even
fresh Avater. It is found both in the Zuyder Zee and
the mouths of the Dutch rivers. The Seine it ascends,
according to Valenciennes, up to Quillebeuf. It spaAvns
in the Mediterranean, according to Risso, and on the
coasts of Holland and Hanover, according to Hoffman'*
and Ehrenbaum, in spring and early summer, from
April till the beginning of July. This seems also to
be the case on the English coast, to judge by an ob-
servation of Mr. Jackson’s (Day), Avho took Anchovies

a According to Gunther (1. c.) the Anchovy is found off Tasmania and New Zealand, in a variety distinguished by a somewhat

greater number of rays in the anal fin.

6 The specimen is so .greatly damaged that the length of the body cannot be stated exactly.

0 Verh. Haarlem 1754 — 93, XI, pars 2, p. 236.

d Verslag van den Staat der Nederlandsche Zeevisschereien over 1884 en 1886.

996

SCANDINAVIAN FISHES.

full of roe off Southport (Lancashire) on the 9th June,
1 87 8 The actual course of the spawning has not yet
been observed; but the eggs, which are easily distin-
guished from the more or less spherical ova of other
fishes by their ellipsoidal form, and which measure
0‘7* — 1'5C mm., have been found on several occasions.
Ehrenbaum met with them in large numbers at the
beginning of June at the surface and in open water
9 — 10 fathoms deep off the island of Norderney (East
Friesland). In the innermost parts of the Zuyder Zee
Anchovy-eggs have been found; but the actual spawn-
ing is probably performed in salt water. According
to Wenkebach (Ehrenbaum), the ova are hatched in
three days. According to Hoffmann, the fry mea-
sure 32 — 55 mm. at the beginning of August, and 50
— 100 mm. at the beginning of September. A Dutch
Anchovy-salter stated too that in August and Septem-
ber such enormous quantities of young Anchovies 50
— ■ 7 0 mm. long are often taken that they are salted
and consumed in the same way as the larger ones, in
spite of the fact that a firkin contains 8,000—10,000
of these small fish, whereas only 3,200 — 4,000 full-
grown Anchovies go to the barrel. Ehrenbaum is, how-
ever, of opinion that these small Anchovies belong to
the spawning of the preceding year'', and assumes that
larvae less than 3 cm. long grow very slowly during
their first, winter and spring in the sea. If this opi-
nion is correct, the Anchovy does not attain sexual
maturity until its third year, which we should else
date a year earlier.

The food of the Anchovy consists principally of
Copepodce, Mysidce, and young shrimps.

The Anchovy is taken partly in fine-meshed drift-
nets, sailing-seines, and shore-seines, partly with hoop-

nets and stationary engines, such as ryssjor (traps) etc.
In the Mediterranean the fishery is carried on chiefly by
torchlight, the glare enticing the Anchovies to the boat,
where the fisherman splashes in the water so as to
frighten them into the meshes of the nets spread round
about. The value of the fishery is generally much less
than that of the Pilchard -fishery ; but the Anchovy
commands a higher price than the Pilchard, and in
certain places the Anchovy-fishery is consequently the
more valuable. The annual take of Anchovies on the
Palermo coast is stated" at 400,000 kilo., valued at the
same number of francs; while the Pilchard-fishery brings
in 600,000 kilo, of fish, but only 200,000 francs. In
France the take of Anchovies for 1881 was nearly
6,000,000 kilo., valued at rather more than 500,000
francs; whereas in 1882 the quantity fell to 1,500,000
kilo., but the price rose to somewhat over 700,000
francs '. Holland had an average annual export for
the years 1883 — 1890 of nearly 1,750,000 kilo, or
35,000 barrels of salt Anchovies, which fetched a price
per barrel of 8 — 78 Dutch guldens* * 6' (on an average,
according to Ehrenbaum, 20 — 30 guldens).

The Anchovy is recommended not only by its
fleshiness and fatness, but also by its flavour, a pecu-
liar bitter taste, which combined with its saltness tick-
les the gourmand’s palate and whets his appetite. This
bitterness, which becomes disagreeable unless the head
as well as the liver and intestine be removed before
salting, was prized by the Romans, to whom Anchovy
sauce ( garurn ) was known. From this too the ancient
Greeks derived their name for the fish, a name which
the species still retains, encrasicholus (with the gall in
the head).

a Both Day (1. c.) and Olsen ( Piscatorial Atlas), however, state that in English waters the Anchovy spawns in autumn, September

— December.

6 According to Wenkebach, quoted by Ehrenbaum.
c According to Hoffman, quoted by Ehrenbaum.

d On the assumption that a growth from 4 — 32 mm. is impossible during the interval between the end of the spawning-season and
the beginning of August. But this growth is not so inconceivable from the beginning of the spawning-season.
e Esposiz. intern, di Pesca, Berlino 1880, Sez. Ital., Gatal., p. LXVII.
f Statistique des peclies maritimes , 1882, pp. 235 and 237.
g A Dutch gulden = Is. 8d.

PIKE-FISHES.

997

ESOCIFOKMES.

Physostoms ivith

bones. The first four abdominal
without connexion ivith the cranial

opercular apparatus.

The Esociform series corresponds to the order
established, under the name of HaplomP, by Cope6.
It includes the Pikes ( Esocidat ), the Mud-Minnows
( Umbridce , named after the Austrian and Hungarian
dogs-fislT ), the Toothed Carps ( Cyprinodontidce — see
above, p. 702 — three species of which family occur
in Southern Europe), and the Blind-fishes ( Heter -
opygii , inhabitants of caves and brooks in North Ame-
rica, a family whose maxillaries, like those of the
Toothed Carps, form no part of the margin of the
mouth, and which is especially distinguished by the
forward situation of the vent, in front of the pectoral

Teleosts, from the head , and without precoracoid
development. Air-bladder , where present , simple ,
and pterygopalatine arches complete , as well as the
No adipose fin.

fins). In the Scandinavian fauna the series is repre-
sented by the first-mentioned family alone.

The series contains very dissimilar forms, predatory
fishes like the Pikes, and more harmless fishes of small
size, principally insectivorous or even mud-eaters. This
dissimilarity manifests itself, as usual, most clearly
in the form of the head and especially of the mouth.
In addition to the above-given characters, hotvever,
the series has a peculiarity characteristic of the great
majority among its members in the backward posi-
tion of the dorsal fin, which lies above or even behind
the anal.

the shoulder-girdle suspended , as usual in the
vertebrae normal in form and
cavity. Hyomandibular

Maxillaries fully developed.

Fam. ESOCIDiE.

Dorsal fin situated , at least for the greater part , behind the perpendicular from the vent,
depressed, and, long (at, least a.s long as the postorbital part of the head or only a little shorter),
upper jaw formed by the tip of the ethmoid bone , the intermaxillaries, and the maxillaries.
canine teeth. The pneumatic duct of the simple air-bladder runs

The family of the Pikes differs considerably from
the preceding family: a Pike and a Herring have not
much in common. But one of the most characteristic
external peculiarities of the Esocoid family, the caudal
position of the dorsal and anal fins, reappears in a
Mediterranean fish Alepocephalus, the type of a family
very nearly allied to the Herrings, and also in the
above-mentioned Paralepidince. The more predatory
instincts of the Pikes, however, accompanied as they
are by a simplification of the intestinal canal, the py-
loric appendages having entirely disappeared, range
these fishes nearer to the Salmonoids, in spite of the
numerous pyloric appendages of the latter. The lateral

Gape large. Snout
Margin of the
Mouth f urnished with
from the anterior part thereof to the oesophagus.

margins of the upper jaw are also formed almost as
in the Salmonoids — short, toothed intermaxillaries (fig.
253, C and F , pmx) and long, though here toothless,
maxillaries ( mx ), each with a well-developed supple-
mentary bone (j) — but show a pecularity which we
have not seen in the preceding fishes, and which is all
the more important, from a systematic point of view,
to the explanation of the forms immediately following
this family. The intermaxillaries, which we have
hitherto found to be more or less closely applied to
each other at the tip of the snout, are here (fig. 253,
C , pmx) separated by the cartilaginous end of the
ethmoid bone (tig. 253, A, C, and E, etcr), which

“ arckoog, simple and loyog , shoulder; i. e. without precoracoid in the shoulder-girdle.
b Trans. Amer. Philos. Soc., Philad., n. ser. vol. XIV (1871), p. 455.
c Heckel and Kner: Susswasserf. Ostreich. Mon., p. 291.

998

SCANDINAVIAN FISHES.

forms the very tip of the snout, and its inferior
covering-bone, the head of the vomer. This peculiarity
shows that the Pikes occupy a comparatively low rank
in the system of the Teleosts, during whose evolution
the rostro- ethmoidal part of the snout has suffered re-
duction, while the parts of the upper jaw have been
more strongly developed. In the head of the Pikes
too the primordial chondrocranium is persistent to about
the same extent as in the Salmonoids. The hyoman-
dibular bones of the Pikes (tig. 253, A and F, hm )

show a great resemblance to those of the Gadoids in
the long, backward process {hmp) with the top of which
the operculum {op) articulates. The anterior of the
two articular heads {lima) which serve to articulate the
hyomandibular bone with the cranium, is directed so
obliquely inwards that it meets the petrosal bone {ptr,
prooticum , fig. 253, E), instead of having its articular
cavity, as is usually the case, on the under surface of
the postfrontal bone ( spho , sphenoticam).

This family contains only one genus.

Genus ESOX.

Body elongated , with the abdominal part longest and of uniform depth °, more or less terete or compressed. The
dorsal and anal fins , which are opposed to each other, short and of the same size. Caudal fin forked. Middle-
sized or small cycloid scales cover the body entirely, the head partially. Pseudobranchice hidden.

Branchiostegal rays

The genus of the Pikes, with its five or six spe-
cies, is distributed over the rivers and lakes of the
north temperate zone both in the Old and New Worlds,
a relic of the period when a land communication, with
collections of fresh water within its bounds, existed
between Europe and North America b But only one
species is found in the Old World.

numerous {11 — 20).

The generic name of Esox was adopted by Artedi1
from Pliny d, who applied it to a large fish of the Rhine.
GesneiP had already conjectured that the Pike was pro-
bably the fish meant; but not even he was confident of
the correctness of his opinion. In his works, as in most
of the ancient authors, the genus was called Lucius, the
Italian Luzzo, from the Greek Xvxof.

THE PIKE (sw. gaddan).
ESOX LUCIUS.

Plate XLIV, fig. 4.

Cheeks and temples, together with the upper part of tfie opercula and the posterior part of the occiput {to a line
with the upper end of the preoperculum) covered with scales; the rest of the head naked. Usually 14 {13 — 15)
branchiostegal rays. Sides of the body spotted with yellow or yellowish white, the unpaired fins with brown.

V.

6—8

R. br. 13 — 15?; D. ; A.

15 — 17

— — — ; C. a? + 1 + 17 + 1 + a:; Lin.

(5)6-8 1

1 2 — 1 3 ( 1 4) ' ‘ 14 — 15’

lat. ca 125 — 130: L. tv.

ca 26 — 28 supra pinn. ventr.; Vert. (58)59 — 62.

Syn. Esox(?), Pun. 1. c. Lucius, Auson., Mos., vers. 122; Belon,
Fat., Divers. Poiss., p. 292; Rondel., Pise. Lacustr., p.
188; Gesn., 1. c., p. 500; Schonev., Ichtliyol. Slesv. Hols.,
p. 44; Willughb., Hist. Pise., p. 236. Esox rostro plagio-
plateo, Art., Syn. Pise., p. 26; De.scr. Spec. Pise., p. 53;
Lin., Fn. Suec., ed. I, p. 114.

a With the exception of gravid females with pendent belly.

b Smitt, Ryggradsdjurens geologiska utve elding och slagtskapsfdrliallanden, p. 59.
c Gen. Pise., p. 14.
d Lib. IX, cap. 15.
e Hist. Anim., lib. IV, p. 368.

f Belon derived the name from the Latin lucere, “because the fish when dried shines in the dark” (see Gesner, 1. c., p. 502).
? Sometimes 16, according to Fries and EkstrOm.

PIKE.

999

Esox Lucius , Lin., Syst. Nat., ed. X, torn. I, p. 314; Bl.,
Naturg. Fiscli. Deutschl., part. I, p. 229, tab. XXXII;
Retz., Fna Suec. Lin., p. 350; Pall., Zoogr. Ross. Asiat.,
vol. Ill, p. 336; Ekstr., Vet. Akad. Hand]. 1831. p. 75;
Nilss., Prodr. Ichthyol. Scand., p. 36; Richardson, Fna
Bor. Amer., part. Ill, p. 124; Fr., Ekstr., Skand. Fislc.,
ed. 1, p. 49, v. We., tab. X; Cuv., Val., Hist. Nat. Poiss.,
vol. XVIII, p. 279; Sundev., Ofvers. Vet. Akad. Fork. 1851,
p. 164; Kr., Danm. Fisk., vol. Ill, p. 236; Nilss., Skand.
Fna, Fisk., p. 348; Sundev., Fiskyng. Utveckl., Vet. -Akad.
Handl. 1855, p. 11; Stockh. L. Hush.-Sallsk. Handl., H. 6
(1855), pp. 82, 91, 167; Hckl, Kn.. Siisswasserf. Ostr.
Mon., p. 287; Sieb., Siisswasserf. Mitteleur., p. 325; Mgrn,
Finl. Fiskfna, (disp. Helsingfors 1863), p. 66; Gthr, Cat.
Brit. Mus., Fish., vol. VI, p. 226; Canestr., Fnaltal., pt. Ill,
p. 21; Lun., Poiss. lac Lem., p. 161, tab. XIX; Coll., Forh.
Vid. Selsk. Chrnia 1874, Tillsegsk., p. 175; 1879, No. 1,
p. 94; Olss., 6fvers. Vet. -Akad. Forh. 1876, No. 3, p. 131;
1882, No. 10, p. 48; Malm, Gigs, Boh. Fna, p. 550; Fed-
ders., Naturh. Tidskr. Kbhvn, ser. 3, vol. XII, p. 81; Bncke,
Fisch. Fischer., Fiscliz. 0., W. Preuss., p. 165; Mor., Hist.
Nat. Poiss. Fr., tom. Ill, p. 466; Mela, Vertebr. Fenn .,
p. 355; tab. X; Jord., Gilb., Bull. U. S. Nat. Mus., No. 16,
p. 353; Day, Fish. Gt. Brit., Irel., vol. II, p. 139, tab.
CXXVI; M6b., Hcke, Fisch. Osts., p. 134; Reut., Sundm.,
Finl. Fisk., tab. XII; Lillj., Sv., Norg. Fisk., vol. II, p.
476; Fat., Fne Vert. Suisse, vol. V, p. 419.

Esox estor, Le Sueur, Val. p. p., nec Dekay; vide Jord.,
Gilb., 1. c.’

The maximum size attained by the Pike in Sweden
may be estimated, on the authority of several trust-
worthy statements from different places, at a length of
19 dm. and a weight of 26 kilo". Pike of this size
are, however, at least in our times, of rare occurrence;
most of the specimens taken measure between 3 and
12 dm. The relation between the length and Aveigbt
varies in different individuals. In old and nearly full-
grown Pike the Aveight increases in a greater propor-
tion than the length.

The elongated body becomes shallower at the head
and tail; but between these points the depth is almost
uniform, so long as the belly is not distended with
food or tumid organs of generation. The greatest depth
measures as a rule 1572 — 17 % of the length, and the
greatest breadth is rather more than half, as a rule
about 52 — 57 %, of the greatest depth. In a transverse
section the body is more or less distinctly quadrilateral,
Avith rounded angles, and broadest in the dorsal region,
groAving more or less compressed towards the belly.

The back is ahvays broad and convex, the belly mostly
flat. The least depth of the body measures in the fry
about 35- — 36 %, in adult Pike about 38 — 43 %, of the
greatest depth, or in the former about 17 — 20 %, in the
latter about 24 — 28 %, of the length of the head.

The length of the head measures in the fry (about
V2 dm. long) about 32 % of that of the body, in full-
groAvn Pike about 27 — 30 % thereof. Its posterior out-
lines form an uninterrupted continuation of those of the
body, but in front of the eyes it is depressed into a
hollow, with the sides somewhat widened in front, and
the tip rounded in an horizontal direction, thus acquir-
ing a resemblance to a duck’s bill. The cleft of the
mouth is large, extending beloAv the anterior margins
of the eyes6. The mouth shows a structure peculiar to
the Pike and intimately connected Avith its well-knoAvn
voracity, in which respect it is surpassed by feAv fishes.
The bones entering into the apparatus of the jaAvs are
endoAved Avith great mobility, the mouth being capable
of considerable distension; and both jaAvs, the palate,
and the pharynx, including the branchial arches, are
Avell armed Avith sharp, retrorse teeth. The mobility
and expansiveness of the jaws are supplemented by
the free articulation of the opercular apparatus.

As Ave have mentioned above, the intermaxillaries
(pmx, fig. 253) are widely separated by the broad, flat,
prominent, and cartilaginous ethmoid bone ( etcr ), Avhich
groAvs broader in front, and, supported by the vomer
(vom), forms the anterior margin of the upper jaAv.
The ethmoidal cartilage is furnished not only Avith this
inferior covering-bone, the vomer, but also Avith two
pairs of upper covering-bones, the supraethmoids, called
by others the nasal bones ( spet , 1 and 2), the anterior
pair situated above and beside the long nasal processes
(frn) of the frontal bones, Avhich processes extend al-
most to the tip of the snout. By means of cartilaginous
joints the small intermaxillaries articulate Avith the an-
terior outer angle of the ethmoidal cartilage and the
last-mentioned nasal bones. An ascending process, flat
and incurved, of the intermaxillaries overlies the an-
terior extremities both of the maxillaries and of the
concave articular head on the anterior outer end of the

° The Fisheries Commission of 1881 — 83 received from several provinces information of Pike 15 — 19 dm. long and 17 — 26 kilo,
in weight.

h The length of the upper jaw from the . middle of the tip of the snout to the hind extremities of the maxillary bones occupies in
the fry (about ]/2 dm. long) about 38 %, in Pike 4 — 5 dm. long about 46 — 49 %, of the length of the head; and the length of the maxil-
laries measures in the latter about 38 — 40 %, and their greatest breadth (across the supplementary bone) about 7 — 8 %, of the same. The
length of the lower jaw is about 65 — 69 % (in the young about 60 %) of that of the head.

Scandinavian Fishes.

126

1000

SCANDINAVIAN FISHES.

palatines (pi), where the anterior extremity of the
maxillaries has its articulation; while the posterior part
of the intermaxillaries is inserted under the anterior
extremity of the maxillaries. In addition to the above-
mentioned covering-bones the ethmoidal cartilage has a
further ossification at each anterior outer corner, an-
swering to the true ethmoid bone (et), which forms
the prominence where the anterior extremity of each
palatine bone has its mobile articulation. The hind
extremity of the palatine is joined as usual by a firm
suture to and covered by the geniculate pterygoid bone
(pt), the lower arm of which is applied and united by

a firm suture to the anterior margin of the quadrate
bone (qu), while the upper arm sends out above the
bend an ascending process which is linked by a mobile,
cartilaginous articulation to the lateral ethmoid (etl,
prefontal bone). The articulations of the hyomandi-
bular with the under surface of the squamosal bone
(squ) and with the upper posterior angle of the petrosal
(ptr), which there meets the latter bone, have already
been mentioned. The whole apparatus of the upper
jaw, the palate, and the suspensory apparatus of the lower
jaw are thus capable both of lateral and upward disten-
sion, the lower jaw simultaneously expanding, for

Fig. 253. Bones of the head in a Pike. l/2 the natural size. A, from the right; B , the skull, seen from "behind; C, the head, seen from
above; D , the skull, seen from the right; Fj, the skull, seen from below; F, the upper jaw, palate, and suspensory apparatus of the lower

jaw, right side, seen from without; G , the lower jaw.

alsp, alisphenoid; art, articular part of the lower jaw; de, dental part of the lower jaw; ept, ento pterygoid; et, ethmoid; etcr, tip of the
cartilaginous part of the ethmoid; etl, lateral ethmoid; fr, frontal; frn, part of the frontal bones that projects above the ethmoidal cartilage;
ftr, exit of the nervus trigeminus; hm, hyomandibular; lima, anlerior articular process of the liyomandibular ; imp, hyomandibular articular
process, with the end of which the operculum articulates; iop, interoperculum; j, supplementary (jugal) bone of the maxillary; mb, tip of the
lower jaw; mpt, metapterygoid; nit, mastoid; mx , maxillary; ocb, basilar part of the occipital; ocl , lateral part of the occipital; ocs, squa-
mosal part of the occipital; op, operculum; par, parietal; pi, palatine; , pmx, intermaxillary; pob, preorbital, the foremost bone in the sub-
orbital ring; pop , preopereulum; psp, parasphenoid ; pt, pterygoid; pimp, post-temporal; ptp, ascending process of the pterygoid; ptr, petro-
sal; qu, quadrate; scl, supraclavicula; sob, hindmost (fifth) bone in the suborbital ring; sop, suboperculum; spet 1 and 2, supraethmoids ;
spho, postfrontal bone; spl, os symplecticurn ; spo, supraorbital; squ, squamosal part of the temporal bone; st, styloid (opisthoticinn) ; vom, head

of the vomer; vomp, shaft of the vomer.

vom

etcr

PIKE.

1001

its two halves are joined in front (mb), at the sym-
physis, merely by cartilage instead of the ordinary
firm suture.

The long maxillaries ( mx ), which are somewhat
curved and twisted, are toothless, and the single or
sometimes (in front) double row of intermaxillary teeth
are not very large; but all the greater is the size of
the teeth on the head ( vom ) of the vomer, in the inner
rows on the palatines (pi), and, above all, in the lower
jaw (de). In the long, posteriorly pointed patch of
teeth that occupies the shaft ( vonip ) of the vomer, ex-
tending back to a line with the nasal cavities, the
dense teeth decrease in size behind, a diminution which
also appears, both outwards and backwards, in the some-
what broader card of palatine teeth, which extends a
little further back than the vomerine card. The man-
dibular teeth of young Pike are set throughout in one
row; in large Pike there are two rows at the extreme
front, both being incurved (recurved) nearest to the
symphysis, but only one row throughout the rest of
the jaw margin. The front teeth are comparatively
small, about twice as large as the intermaxillary teeth;
but the size increases backwards, while the teeth be-
come more scattered, the posterior two-thirds of the
jaw margin being armed with only about 6 large ca-
nines. Among these canines the middle ones are the
largest, being of so considerable a size that in Pike 9
dm. long’ the largest mandibular tooth measures together
with its base 19 mm. and above the gum 14 mm.
The tongue is cartilaginous, somewhat broader in front,
but incised at the truncate tip. The hind part of its
lingual bone and the three copular bones (the first and
largest of which is formed by the coalescence of two),
are set with dense cards of small teeth. The roof of
the pharynx is armed with two pairs of cards, on the
upper pharyngeals, containing teeth about equal in size
to the foremost in the lower jaw, the anterior cards
pointed in front, and both cards together of about the
same size and form as either of the two cards on the
lower pharyngeals. The last-mentioned bones are slen-
der (resembling branchial arches), but furnished on the
posterior inward side with an elevated, flat process,

over which the card of teeth extends, being thus raised
free above the hind part of the pharyngeal itself. All
the hones already enumerated in the mouth and pha-
rynx may be dentigerous in most of the Teleosts; but
in the Pike we further find that the gill-rakers, on the
front of the branchial arches, are transformed into dense
cards of teeth, divided into quadrilateral or rounded
(verrucose) groups, and containing teeth about equal
in size to the intermaxillary ones. The number of the
teeth is thus augmented so considerably that Keoyer
succeeded in counting “ certainly more than three thou-
sand teeth” in the mouth and pharynx of a Pike. All
the teeth are ophidian in type, pointed, for the most
part curved, rigid in an outward direction, flexible to-
wards the interior of the mouth, with the tip directed
inwards and backwards, thus affording a ready ingress
to the prey, but preventing its egress. Exceptional in
form are the straight small teeth on the branchial ar-
ches, and the largest canines in the lower jaw, which
are also straight, but dagger-shaped and two-edged.

The number of the branchiostegal rays varies be-
tween 13 and 16, being not unfrequently different on
either side. All of them are sabre-curved, the posterior
sharp, the anterior rather more terete. The anterior
(lower) 6 or 7 are set on the ceratohyoid bone, and
are somewhat separated from the posterior (usually 8),
which belong to the epihyoid. The branchiostegal mem-
branes are free and separate underneath to the root of
the tongue, where they are attached, the very extre-
mities crossing each other. This arrangement further
contributes to the distensibility of the pharynx.

The top of the head is flat, with a concavity be-
tween the eyes. These are large", oval5, rather far
apart0, and set high, about half-way along the head A
Above their anterior part projects a semi-elliptical or
rounded quadrilateral bone (fig. 253, spo) that calls to
mind the superciliary shield in birds of prey, and gives
the Pike a similar fierce and savage look. The nasal
cavities are large, and lie just in front of the eyes,
each containing two apertures, the anterior round, the
posterior furnished with a rounded dermal lid, origi-
nating from the fairly broad ridge of skin between them.

° The longitudinal diameter of the eyes measures in the young about */5, in Pike 4 — 5 dm. long about J/8 — 1/9, of the length
of the head.

b The vertical diameter of the eyes is about 80 — 85 % of the longitudinal.

c The interorbital width at the middle of the eyes is about 1/s (19 — 21 %) of the length of the head.

d The length of the snout is about 42 (sometimes 40) — 45 %, and the postorbital length about 43 (in the fry 37)— 46 %, of the
length of the head.

1002

SCANDINAVIAN FISHES.

The cephalic system of the lateral line has very
distinct and deep pores in the thick skin. In each
fronto- rostral branch there usually appear 4 pores along
the forehead and snout; in the suborbital branch (tig.
253, sob — pob) 9 or 10 in a row surrounding the lower
part of the eye and advancing along the snout to a
point just in front of the nostrils; in the mandibular
branch 5 pores on the under surface of the lower jaw
(de) and -6 at the preopercular margin. The occiput
is crossed on each side by a serpentine groove (the
supraorbital branch, cf. above, fig. 104, p. 368), ending
at the outer extremity in a similar pore, from which
a row containing 3 or 4 pores runs back, along the
temple, to the upper angle of the gill-opening. The
body is covered with soft, middle-sized, and imbricated
scales (fig. 254), each enclosed in a thin follicle and
naked only at the tip, which appears in a small, an-

of the length of the abdomen, measured from the an-
terior (lower) end of the shoulder-girdle.

Among the seven fins the vertical, which are si-
tuated on the tail, are most developed, forming the
propeller by the aid of which the Pike darts like an
arrow on its prey. The dorsal fin is high and rounded.
It begins vertically above the vent, at a distance from
the tip of the snout measuring 67 — 69 % of the length
of the body, and terminates above the end of the anal
fin. Its base measures about 1 1 lj2 — 1372 %b, and its
longest ray about 40 7 2 — 1 1 72 0/o° •> °f P‘e length of the
body. The anal fin is similar to the dorsal, but shorter,
usually almost as high as it is long. Its distance from
the tip of the snout is about 71 — 74 %d, its base about
107* — 972 %i and its height about 10 — 11 %, of the
length of the body. The caudal fin is slightly forked.
The middle rays, which occupy about 6 — 7 % of the

A.

B.

c.

Fig. 254. Scales of the Pike.

X 2: A, from the anterior part of the body, above the lateral line; B, from the lateral line;

C, from the ventral side.

gular opening, usually of a distinctive colour. In shape
they are rounded and oblong, or (on the belly) more or
less pointed behind; and they are most distinguished
by the cleavage of the anterior (inserted) part into three
or four lobes. The lateral line is straight, and runs
somewhat nearer to the back than to the belly. It is
marked by scales incised at the tip, but is interrupted
here and there by scales with entire tip. On each side
of the lateral line, especially on the tail, there are
scale-rows similar in the structure of the scales to the
lateral line, but varying in number, extent, and position.
At the outer angles of the ventral fins the usual scale-
flap" is wanting.

The anal aperture, which is large, lies so near to
the caudal fin that its distance therefrom is only 7 3

length of the body, are about half as long as the long-
est rays in the fin.

The pectoral fins are obliquely rounded and set low.
Their length in adult Pike is about 10 — 13 % of that
of the body. The ventral fins are of about the same
size. They are situated half-way along the body, and
the preabdominal length is about 7 4 (in gravid females
as much as 7io)> 4iie postabdominal about 1/5 (in gravid
females somewhat more), of the length of the body.
Without any examination of the internal organs, the sex
may be determined by the fins, which are generally larger
and more rounded, with thicker rays, in the male than
in the female. The length of the pectoral fins, in parti-
cular, is somewhat more in the male, less in the fe-
male, than half of the preabdominal length.

“ Cf. above, on the Loaches (p. 709 and note d) and on the Minnow (p. 756).

b In the fry about 36 — 38 %, in Pike 4 — 5 dm. long about 44 — 47 %, of the length of the head.

c In the fry about 34 — 38 %, in Pike 4 — 5 dm. long about 38 — 43 %, of the length of the head.

'7 In the fry about 2/3; in the males commonly less than in the females.

PIKE.

1003

The coloration of the Pike is black above, white
below. The sides are grayish green, with transverse
bands of greenish yellow, more continuous in the
young, broken up in old Pike into almost round or
oval spots, sometimes elongated, like a ribbon, in the
longitudinal direction of the body. The ground-colour
of the fins is grayish yellow, waved with a darker tint.
The anterior margins of the ventral and anal fins and
the inferior lobe of the caudal are of a purer yellow.
The iris is yellow, with a dark, grayish brown spot
both in front of and behind the pupil. From the eye
two sinuous, greenish yellow stripes run to the hind
margin of the operculum, and some irregular spots of
the same colour appear on the lower part of the said
bone. The snout has an elongated, triangular patch,
coursed by a longitudinal black stripe, on the sides in
front of the eyes. On the sides of the body the scales,
as we have mentioned above, have a naked, crescent-
shaped or angular spot, with the angle pointing for-
wards, and of a metallic lustre, witli a dash of yellow
or white. The ordinary dress of the fish, as described
above, shows variations dependent on age, the fluctua-
tions in the supply of food, and the nature of the sur-
rounding water. Old specimens have a purer, more
defined coloration. The Pike that live in turbid water
are always darker than those that inhabit clear lakes
or streams; and in small lakes with dark water almost
black specimens have been found. In the inner island-
belt. of the Baltic small Pike are taken at the beginning
of winter which have a dark lemon-yellow coloration
instead of the white tone. The islanders - believe that
these Pike have just arrived from the outer islands or
the sea outside, and have therefore conferred upon them
the name of Nylcindare (new-comers).

On an examination of the internal organs the liver
is found to consist of a single lobe, long, obtuse, and
arched, which lies principally on the left side of the
body, and extends back to a point, just short of the in-
sertions of the ventral fins. On the concave upper sur-
face of the liver, at the extreme front, but to the right,
lies the gall-bladder. The intestinal canal is simple.
When the stomach is empty, it apparently composes
together with the oesophagus a tube of almost uniform
width, extending to about a line with the beginning of

the last third of the ventral tins when folded. At. this
point it. bends abruptly, and with a slight constriction
at the pylorus is continued in a forward direction by
the intestine, which advances below its right side to
the gall-bladder, and bends there with equal abruptness,
to return in a straight line to the vent. The spleen is
triangular, and lies close t.o the termination of the sto-
mach. The gall-duct is long, and enters the intestine
at some distance below the pylorus. The air-bladder is
long, almost cylindrical, and is connected with the oeso-
phagus by a short, and narrow duct from its anterior
extremity. The testes and ovaries are long, and follow
the direction of the abdominal cavity. The kidneys,
which lie above the air-bladder along each side of the
spinal column, are dark red. The urinary bladder is
thin, cylindrical, and rather long, and shares with the
sexual organs a special aperture behind the orifice of
the intestine.

The Pike has a very extensive geographical range
— about the same as the Minnow’s (see above, p. 757) —
from North-eastern Siberia west to the east, of North
America. According to Pallas it inhabits the River
Amoor“, which falls into the Sea of Okhotsk, and the
Rivers Indigirka and Chatanga, which disembogue into
the Arctic Ocean. Brehm found it in the lower course
of the River Obi'', and in the great lakes of the Baraba
Steppe (the upper basin of the Obi and Irtish) it is ex-
tremely common, according to Pallas, and attains a
considerable size. It also occurs in the Caspian Sea, but
not in Transcaucasia, nor in the Black Sea, though it is
found in the Sea of Azov and the basin of the Danube.
It is met with in rivers and lakes throughout Russia,
North and Central Europe, including Great. Britain and
Ireland, and Italy, including Sicily (Canestrini) ; but is
said to be wanting in Greece and on the Pyrenean Pen-
insula. It- inhabits almost all the waters of Scandinavia;
but. in Norway Collett has observed that its range,
like that of the Perch and several other fresh-water
fishes, is interrupted, the gap extending over the Pro-
vince of Trondhjem, Nordland, and the whole west coast..
Though really belonging to lakes and rivers with fresh
water, it is also found in the island-belt of the Baltic.
That salt water is not its true home, appears from the
fact that among the said islands the Pike shows a de-

“ Assuming that Pallas here refers to our common Pike. From the Onon, a Daurian tributary of tire Amoor, he describes a “va.
riety” of a coloration more suggestive of the American Esox nobilior.

On the Kamchatkan Peninsula, according to Pallas, the Pike is wanting.

4 Brehm’s Thierleben , 2:te Aufl., Or. Ausg. (1879), Bd. 8, p. 249.

1004

SCANDINAVIAN FISHES.

crease both in size and numbers the nearer it ap-
proaches to the open sea, at last disappearing entirely. It
ascends to a great altitude among the mountains; but
in Switzerland, according to Fatio, stops short at a
height of 700 — 800 m. above the level of the sea,
though it has been planted with success 1,100 m. or
even 1,400 m. above the sea-level. In Tornea Lapp-
mark it inhabits the alpine lakes above the birch-region;
in Jemtland, according to Olsson, it ascends to lakes
at least 650 m., perhaps 740 m. above the level of the
sea. In its American range too, the Pike is common
from the Arctic regions (Richardson) south to Ohio;
but it is not included in Bean’s list of the fishes of
Alaska. From Greenland and Iceland it is unknown.

The Pike chooses its spring and summer haunts
by preference in shallow inlets with a weedy bottom
and shores overgrown with reeds and rushes. Towards
autumn it betakes itself to precipitous, stony shores,
which it again forsakes when the winter is at hand,
and the inlets freeze. Most of the Pike then return to
their summer stations; but the larger ones seemingly
follow the shoals of other fishes to the depths, being
seldom caught during winter in shallow water. To
these a more plentiful supply of food is, no doubt,
necessary than the shallows afford in winter-time.

When the Pike has chosen its station for the season,
it restricts its wanderings to the immediate neighbour-
hood, leading a solitary life, and never seen in com-
pany except during the spawning. Its attachment to
one spot may be gathered from a circumstance related
to us on trustworthy authority. A Pike that had ta-
ken a trolling-hook and escaped was recaptured after
several years in a seine at the same place, and the
hook was found in its stomach.

The Pike is undoubtedly the most voracious among
the fresh-water fishes of Scandinavia. It devours in-
discriminately other fishes, young waterfowl, small
mammals, and carrion. From the dense bed of grass
or rushes where it usually passes the day in stationary
watch, it pounces with the speed of an arrow on its
unwary victim. It almost always seizes its prey cross-
vise, and retains its hold until the latter is dead or
so exhausted as to desist from all struggles. Then the
Pike turns the prize in its jaws till the head points
towards the interior of its mouth, and commences its

meal. This operation is a protracted one when the
victim is large, for the end first swallowed and received
in the stomach must digest to make room for the
remainder. Mr. C. Fk. \Wern has kindly communi-
cated the following observation in point: “I have kept
Pike together with a Salmon in a pond fed with run-
ning water. The former were usually stationary, the
latter kept constantly on the move. On one occasion
I saw a Pike of 7 or 8 lbs. weight dart forward and
seize the Salmon, which was quite as large, in its for-
midable jaws right across the body. The combat was
fierce. The Salmon leapt out of the water and made
desperate, but fruitless struggles to shake off its re-
lentless captor. In a couple of hours’ time the Salmon
was utterly exhausted, and the Pike began to swallow
it head first. The meal lasted three days before the
whole body had disappeared. The process of digestion
must have taken much longer, for all the following
week the Pike had a very swollen appearance, and
could hardly be induced to move by touching it with
a long stick.” This voracity grows to ravenous and
insatiate hunger, and blood-thirsty tendencies are in-
herent in the Pike from its earliest youth. Baron
C. J. Cederstrom relates® from his experiments in
pisciculture that on the 17th of June he had four
small Pike, two of them measuring about 26 mm., the
other two about 20 mm., living together in a bowl of
water. They had been fed with small Ide-fry, the
larva? of gnats, and water-fleas ( Daphnia ). In dark-
ness they ate nothing, their rapacity was excited only
by the moving objects appreciable to their sight. Dur-
ing the night their bowl had been covered with a lid.
When he removed the lid at 5 a. m., they were to be
seen all four; but when he returned a quarter of an
hour later, one of the larger pair had begun to devour
one of its smaller comrades. Half of the victim’s body
still stuck out of the other’s mouth, and moved for
some moments, while the stomach of the latter was
monstrously distorted and distended. Soon afterwards
he observed how the other pair kept a jealous watch
on one another. The larger of them soon made an
assault, but was foiled, the smaller escaping by a dex-
terous movement, yet without fleeing to any distance.
But the second attack was rewarded with complete
success, and the two victors now swam about, glutted

In Sundevall, bfvers., ]. c.

PIKE.

1005

with prey. After two hours had elapsed, the tails of the
victims had disappeared into the maws of their devourers.

The strength of the Pike is fully proportionate to
its size and voracity. We have just seen that a Pike
can come off victoriously from a battle with a Salmon
of its own size; but Ekstrom adduces® a more singular
proof of its powers. In the back of a Pike that did
not weigh more than 8V2 kilo., he found the skeleton
of an osprey ( Pandion haliaetus), which had rashly
attacked the fish, but been drawn into the depths and
drowned. Similar tales are told of sea eagles that
have perished in the attempt to capture and carry off
the Pike.

Though not very tenacious of life, the Pike may
be kept alive for a long time in a well, especially
if it has some smaller fish to prey on, for it re-
tains its rapaciousness even in captivity. In contra-
distinction to other fishes, it continues to feed during
the spawning-season. On the 9th of June, 1893, we
purchased in Stockholm a gravid female 472 dm. long
strongly distended'' by the ovaries, which were quite
ripe, and voided the eggs at the least pressure. In
the stomach, however, lay a Roach 12 cm. long, only
the head of which had digested.

The spawning is of long duration, its season de-
pending on the age of the fish. The young spawn
first. When they have finished, the middle-aged Pike
begin, and last of all the oldest and largest. In spring,
before there is open water in the lakes, the Pike com-
mences to ascend towards the shores. A tradition, handed
down among the fishermen of Central Sweden from
time immemorial and still surviving, states that on
St. Gregory’s Day (March 1 2th) the Pike turns its
head towards the shore, and on St. Gertrude’s Day
(March 17th) begins its ascent. It repairs to those
parts of the shore where streams and brooks fall into
the inlets, rendering them in the Swedish fisherman’s
parlance landlosa {landless). The Pike that arrive first
are known as Gertrude’s or Ice Pike. A great por-
tion of the roe they deposit is probably destroyed,
for it is often committed to the open lake, where it
is exposed to so many dangers. When the spring is
so far advanced that the lakes are ice-free, the brooks
clear, and the low-lying meadows round the shore
under water, the larger Pike make their way to these

inundated places, and begin to spawn. As this gene-
rally coincides with the pairing-time of the frog, the
Pike that breed at this season are called Fro-gaddor
or Gloss-gciddor, from Kdllfro and Glossa, names ap-
plied by the peasantry to the frog {B,ana temporaria).
They are also known as Gras-gaddor (Grass Pike)
and Angs-gaddor (Meadow Pike), from the nature of
their spawning-place. Last of all come the largest

Pike. These usually begin to breed at the end of
May, when the trees are in leaf, and several flowers
in bloom; and are hence called Lof-gaddor (Leaf Pike)
or Blomster-gdddor (Flower Pike). They are few in
number, like the Ice Pike, and spawn in water of
some depth within weedy inlets. The earliest Pike,
which are often of a brassy yellow colour, also bear
the name of Messing s-gciddor (Brass Pike). The some-
what older fish are known, on account of their slender
form, as Langstjert-gaddor (Long-tailed Pike), and the
oldest, with their thicker body, as Kortstjert-gaddor
(Short-tailed Pike).

The further north the Pike has its home, the
later is the spawning-season. In the Tornea Elf the
Pike does not breed until the middle of June. The
Fro-gadda or Gras-gddda always spawns in shallow
water; and as the breeding fish are neither timid nor
massed in numbers at the same place, they may be
closely observed. The females, which are always larg-
est, come first, each attended by two or three, seldom
four males. The female swims so high in the shallow
water that, when the weather is calm, the surface is
faintly rippled by her movements. Now and then the
dorsal and caudal fins may be seen above the surface.
As soon as the female halts, the males approach and
surround her, one on each side, and, if there are
more than two, one under the tail and another above
the back. They rub themselves against her body,
during which operation she keeps still, only moving
the fins. After a while she disperses the males with
a sudden lash of her body, and darts to another
point, where the same operation is renewed. Mean-
while she deposits on the grassy bottom the yellowish
and coarse-grained roe, which is impregnated by the
milt. In a Pike weighing 6 German pounds (24/5
kilo.) Bloch counted 136,500 ova; in a female weigh-
ing 28 lbs. avoirdupois (12*7 kilo.), and with ovaries

0 Vet.-Akad. Handl. 1831, p. 79.

6 The greatest deptli 21 % of the leugth of the body.

1006

SCANDINAVIAN FISHES.

weighing 21 oz. (607 grammes), Buckland® found
292,320 eggs, and in another weighing 32 lbs. (141/ 2
kilo.), and with ovaries weighing 5 lbs. (21/* kilo.),
595,200. The Pike is thus fairly prolific, but its fe-
cundity is not above the average, as compared with

B

Fig. 255. A, young Pike 2 days old and hardly 10 mm. long.
The vent has not yet divided the vertical embryonic fin. Pectoral
fins rudimentary. No fin-rays. Immediately after hatching the larva
is hardly 9 mm. long. B , young Pike 11 days old and 15 mm.
loDg; a, from the side; b, from above. Vent open. Ventral fins
have appeared about half-way along the embryonic fin in front of
the vent. No fin-rays. Distinct teeth in the lower jaw. Opercula
developed. C, young Pike 5 or 6 weeks old and 20 mm. long.
Dorsal and anal fins begin to separate, before the rays are formed.
In the caudal fin the base seems to develop first, the rays being
rather short. D , young Pike 7 or 8 weeks old and 26 mm. long.
Of the vertical embryonic fin only a small remnant to be seen under
the caudal fin and another, very narrow, from the anal aperture to
the ventral fins. All the fins, though thick and rigid, have the com-
plete number of rays. In the ventral fins the rays are separated at
the base. The upturned tip of the tail still lies, rather large, in the
form of a compressed, membranous lobe, above the caudal fin, but
is confluent therewith, and no longer shows increase in size. —
SUNDEVALL.

Even in a larva 15 cm. long and between 6 and 7 months old
(killed on November lOtli and born of an artificial impregnation per-
formed in April) the development of all the fins and the cleavage of
the branched rays are not much further advanced than in Hemibranchs
and Lophobranchs (see above), these rays being compressed, with re-
mote articulations, and with short and imperfect, almost exclusively
simple division.

that of other fishes. The eggs, which are at first
rather adhesive, lie free on the bottom, and in spring
(April) require about three weeks to hatch.

“The new-hatched fry”, writes Sundevall, “are
short and thick in shape, with rather large belly (fig.
255, A). The coloration is yellowish, but quite trans-
parent, and densely punctated on the surface with
black dots, a dark band running from the eye along
the sides of the belly. In coloration, external form,
and movement they have some resemblance to tadpoles;
but some are much more sparsely punctated with black,
and consequently of a lighter tone. At first the larva
remains for the most part still, preferring to lie close
to the surface beside plants or floating straws and the
like, to which it seems, as it were, to hang, or else
at the bottom in less than an inch of water. On being
touched, it swims rapidly about, with hasty movements
of the tail, but soon resumes its former position. In
about 10 or 11 days (fig. 255, B) the yolk is ab-
sorbed, and the belly much reduced in size, but the
head prolongated, with large mouth. It now begins
at once to swim more steadily, in the same manner
as its elders, and goes in quest of prey. It soon
abandons the habit of lying at the bottom or resting
alongside floating objects, repairs to somewhat deeper
water, remaining for the most part stationary, as if
on the watch for prey, and seizes small fishes and
other aquatic animals of a size considerable enough
in comparison with its own, but only those which it
sees moving, just as older Pike .... The external
form in which the specific characters of the Pike may
be traced (fig. 255, I)) seems to be fully developed at
an age of nearly 2 months and a length of about a
Swedish inch (25 mm.).” Subsequently the growth
proceeds rapidly, as usual, at first, but with very con-
siderable variations, depending on the different supply
of food under circumstances favourable in other re-
spects. According to some observations a one year
old Pike is only 15 cm. long, according to others 30
cm. Blanchere states the growth as follows* 6:

Maximum length of the Pike at the age of 1 year .

n ?? '>') n v 5? >> ^ years.

?? 11 'll 'll >i i) ii i) ^ n

?! ) ? ii ii a a a a a ^ a -

1 2

>, >> >) ii a ii ii ii ii x “ ii -

M. M.
0-25—0-30
0-36—0-42

0- 55—0-60

1- 00
1-35.

a Nat. Hist. Brit. Fish ., p. 387.

6 Dictionn. Gen. Peches , p. 116.

PIKE.

1007

How widely such computations may differ, appears
from Ekstrom’s observations. He found that Pike-fry
37 — 49 mm. long, kept in a spring with muddy bottom,
only attained in 5 years the size of a common Herring;
but that a specimen 15 cm. long, kept in another spring
with smaller fish to feed on, attained in 5 years a
length of 4 dm. The statements we possess of the Pike’s
duration of life — according to old accounts over 200
years — whatever may be their authenticity, only show
that this fish may probably live to a very great age.

As a food-fish the Pike is of no small value. The
flesh is white, firm, wholesome, and comparatively free
from bones. Fresh Pike is by no means bad eating;
and the flesh has an advantage over that of many other
fishes: it may be kept for a long time, without de-
teriorating, in a salted or dried condition. The great
havoc wrought by the Pike among other species of fish
has given rise to proposals for its extermination, at
least in small pieces of water. But the only result of
such a procedure would be the increase in number of
the smaller species to so great an extent that the
supply of food would at length fall short, the develop-
ment of the fish be arrested, and a. great proportion
of them die out.

Some stated peculiarities in the history of this
species may lastly be adduced. The fishermen in ge-
neral believe that at certain seasons of the year the
Pike entirely abstains from food ( tager icke svalg as
the Swedish fishermen say), and at others is excessively
voracious. These seasons are said to be periodical and
regular in occurrence, the observant fisherman being
able to predict the time when the Pike is on its feed
(i taget), as it is called. But these periods are said
not to occur at the same time year after year, and
according to some observations they are determined by
the spawning-season, for the period of voracity begins
in the same change of the moon (waxing or wane) as
the Pike finishes spawning. There is one exception,
however, the Pike being always on its feed throughout
the dog-days. This periodical voracity and moderation
is said to depend on the circumstance that, when the
Pike is off its feed , the gums are so swollen that the
points of the teeth hardly project above the flesh, some
tenderness of the gums being thus the curb of the
Pike’s usual rapacity. Perhaps we have here some
observation of the manner in which the Pike casts its
teeth, or we may find a more probable explanation in
the fact that the fish requires some time to digest the

great quantity of food which it devours during the pe-
riod of voracity. Another singularity is that the Pike
can disgorge its prey, a consequence of the structure
and simplicity of the intestinal canal.

Most of the fishing methods employed for the Pike
depend for their success on its rapacity. It is this
quality too that entices the Pike to the spawning-places
of most other fishes, where it finds prey in abundance,
and is always on the alert. The Pike is consequently
taken on many occasions in the tackle set for other
spawning or schooling fishes. We may also assume
that the Pike is caught with almost all kinds of fishing
engines. We shall confine our attention, however, to
those which are used in Sweden especially for Pike.
Among them we find:

The ryssja (fig. 7, p. 33, above), in which Pike
are taken principally during the spawning-season, when
they ascend into inlets and channels, to low-lying, in-
undated meadows or marshland, and to grassy shores
where the water is shallow.

Nets (ordinary gill-nets, Pike-nets, with a mesh
of more than 25 mm.) may be employed all the year
round, so long as there is open water, but are al-
ways unproductive tackle. When used for Pike, they
are set in shallow, grassy inlets, and sometimes to
close broader pieces of water and thus guide the Pike
into ryssjor or similar traps.

The trammel-net (fig. 183, p. 741, above) is plied
only in summer, and always with success off shores
fringed with reeds and rushes, where the Pike usually
takes its station on bright and warm summer days.

The long-line ( backa , g add-ref) is used from the
end of May, when the Pike has finished spawning, till
late in autumn. The warmest part of the summer is,
however, an unfavourable season, partly because the
high temperature of the water soon kills the bait, which
should always be alive in this method of fishing (fig.
256), partly because the Pike then has an abundance
of small fish to feed on. In early summer the lines
should be set on grassy shores and round shallows
and islets, some yards outside the edge of the reeds,
but from September and during the rest of autumn
in deep inlets, with stony or sandy shores thinly fringed
with reeds.

The angelkrok (really triangle-hook , originally in
the form of a triangle with one side open, fig. 257).
The winter is the only season. The fisherman chooses
his pitch in bays or creeks of no great depth, with

Scandinavian Fishes.

127

1008

SCANDINAVIAN FISHES.

Fig. 256. Different methods of fastening live bait on the snood hooks of a Pike-line.

reedy shores, but else clear of weeds, so that the hooked
fish cannot run the line foul. Though this fishing may
be practised all the winter, it succeeds best and is
attended with least difficulty early in the season, be-
fore the ice has attained any considerable thickness,

Fig. 257. Different forms of triangle-hook for Pike-fishing under
the ice with live bait. The bait is attached by inserting the point
of the hook in the fleshiest part of the back beside the spinal co-
lumn and drawing it out in the direction of the head, so that, when
the bait is left in the water, the hook keeps it in a natural position,
and the point of the hook is directed backwards.

and after some snow has fallen, to render the ice less
transparent and to give the fisherman a foothold.

The stdngkrok (fig. 258), with the slang (a thick
rod) obliquely thrust into the bottom, and with the
line wound up, so that the Pike can run to its entire
length until the fisherman comes to examine his tackle,
may be used not only in summer, but also, with some
modification of its setting, in winter. In all seasons

Fig. 258. a and b, the tops of two rods as used in stdngkrok
fishing (as much as should be visible above the water); «, with the

line wound on a boiv (hazel best); b, with the line wound on a forked
stick (juniper best), c, bait fastened on the hook (see above, fig.
256), but also suspended from the line by a fine thread (attached to
a pin stuck through its back), so as to keep it in an horizontal posi-
tion. d, dead bait suspended in a triangle, where the line (the thick
line in the figure) forms one side of the triangle, the hook and the
bait form the base, and a fine thread, attached as in c, forms the

third side.

the stdngkroh is invariably set near the edge of the
reeds, off rocks, promontories, and holms, in such in-
lets as the Pike is supposed to haunt.

The klumpkrok (fig. 259), with the rod of the pre-
ceding method replaced by a floating klump (a piece
of timber 3 or 4 feet long), one end of which is an-
chored with a stone, the line, wound up on a bow,

Fig. 259. Klumpkrok.

being suspended from the other end. Used at the same
season as the stdngkrok and at similar spots.

The palkrok (fig. 260), with one end of the line
anchored by means of a stone but fastened at a dis-
tance from this point somewhat less than the depth of
the water, by means of a running noose, in a notch
at the end of a. floating pale (a splinter of spruce-fir
about 3 feet long), so that about a third of the splinter
projects above the surface. Employed at the same
season and in the same places as the klumpkrok , to
which it has no slight resemblance.

PIKE.

1009

Fig 260. Palkrolc.

Trolling (Sw. slant), an old and well-known me-
thod of angling, the description of which would carry
us too far, if we attempted to give every detail. Iiire
derives the Swedish name from the Belgian slonden (to
gorge). The rod and line are plied from a rowing-

grass or weeds that grow at the bottom or in mid-
water without reaching the surface, in which case they
may be avoided by the angler. Lastly the snood — a
suitable violin-string (silver-string) is best — is fastened
with a peculiar kind of knot (fig. 261, g ), tied in the
loop at the end of the line. Now the trolling begins:
the bait is dropped where the fisherman hopes to find
a Pike, but soon drawn up again, so as to keep it bob-
bing up and down. This motion entices the Pike to
seize the bait, which the angler easily perceives by the
check. As soon as the Pike has taken the bait, the
fisherman instantly jerks back the rod sufficiently to
allow the loop to slip off the pin at the end of the rod.
The line is now free, and is let out until the Pike stops,
when the boat is kept stationary, either by heading it
towards the reeds and holding it fast, or by thrusting
the butt of the rod into the bottom of the lake. The

Fig. 261. a, reel ( lekare ); b and c, baiting-needles of different form (7 or 8 in. long, of wood or brass), in which the snood is wedged
when it is to be passed through the bait; d — /, gorge-hooks of different form; g, lower part of the loop at the end of the line and the

upper end of the snood, showing how the knot between them is tied.

boat by one man, while another pushes the boat cau-
tiously along the edge of a bed of reeds or in some
other place supposed to be haunted by Pike. The line
runs through a loop at the end of the rod, or is at-
tached by a loop ( oska ) to a straight pin of wire fixed
there, and can be let out or taken in at pleasure by
means of a reel ( lekare ) fastened above the butt of
the rod. The snood with the hook at its end is passed
through the bait (a Roach, Bleak, Crucian Carp, or
other Cyprinoid with bright scales) from the mouth
to the end of the tail. The hook is shanked with lead,
has one or two prongs (fig. 261, d — ■/), and should
be barbless, the prongs being left to project backwards
from the mouth of the bait. The snood is lashed with
thread to the tail of the bait, and all the fins are then
cut off, to prevent the bait from running foul of the

fisherman now waits a while, at most 15 — 20 minutes,
for the Pike to pouch, when it again begins to move;
and now is the time to haul in both line and fish.
Trolling may be practised with success from spring,
after the close of the spawning-season, till late in autumn,
though it should be observed that cloudy weather and
a light breeze are more favourable than calm, fine days,
on which the fisherman is seldom rewarded. The fish-
ing-place is chosen during summer in grassy ancl
shallow inlets, during autumn off stony shores and in
deep water.

Springkrok (fig. 262) is a modification of the
preceding kind of fishing, and is conducted most com-
monly at times when the fisherman suspects that the
Pike is off its feed. The place should be the same
as in trolling.

1010

SCANDINAVIAN FISHES.

b

Fig. 262. Springkrolc, hollow, with the elastic arms held back by
their pressure against the sides of the tube, as in a, or
drawn out, as in b.

Drag (spinning), also called Drag span, Oppror ,
Bottenror, etc., is a well-known kind of angling. A de-
ceptive hook (tig. 264), or a range of hooks on a bright
metal bait imitating a fish, is towed with a long line,
which can be wound in on a reel (fig. 261, a ), after a boat
rowed at an even and rather rapid pace. The fisher-
man’s object is to attract the attention of the Pike in
the same parts of the water as when trolling; but in
this case he can manage without assistance if he holds
the line between his teeth. Spinning is practised

Fig. 263. Spinning-bait for Pike.

throughout the summer and autumn, but the best
season is from the middle of August till the end of
September.

Spearing by torchlight (eldstodjning) is perhaps
one of the oldest kinds of fishing, and has advantages
as being one of the least expensive. The method of
using the light we have seen in the Stickleback-fishery
(see above, p. 657, fig. 164); but when spearing Pike,
the fisherman must find out the spot where the fish is

standing, and when he strikes with the leister ( ljuster ,
fig. 264), he must aim the blow at the head of the fish,
in case it should take to flight. Spearing is practised
in spring and autumn', when the nights are dark, and
off shores with shallow and clear water.

Fish may also be speared in broad daylight. This
kind of fishing is called ljustring (leistering), and is
most often employed for Salmon on their upward jour-
ney, but also for Pike when they are spawning on in-
undated meadow-land. The fisherman always goes
alone, and either stands on the shore or wades, with
his creel on his back to receive the speared fish. But

Fig. 264. Salmon and Pike spear, with the lower part of the shaft.

as the fish are more difficult of approach in the day-
time, he must be skilled in the special art of hurling
the spear at his quarry.

Spearing by torchlight is now less in vogue than
formerly: more effective methods of fishing have been
invented, and men of the craft have learnt by expe-
rience that it frightens the fish from shore, and thus
harms the fishery.

We should mention one more way of catching
Pike, though it can never be rewarded with any great
success. When the Pike is spawning or standing among
grass where it thinks itself hidden from sight, it may
be captured by cautiously passing a noose of copper
wire, attached to a stick, over its head until the noose
is past the gill-openings, when the fish should be pulled
up at once.

(Ekstrom, Smitt).

EEL-FISHES.

1011

ENCHELYMOKPHI.

Shoulder-girdle detached above from the head and suspended at the sides of the spinal column behind it. No

prcecoracoideum. Anterior vertebrae normal: air-bladder
Opercular apparatus complete; but the liyomandibular

arch, incomplete.

Regarded either from a biological or a morpho-
logical point of view the Enchely morph series is one
of the most remarkable among fishes. The life of the
Eels, especially their sexual life, though studied for
thousands of years, was shrouded in a veil of mystery
to a time within the memory of man; and the mor-
phological elucidation of their structure shows the Te-
leosteous type in a singular simplicity, whether this
should be explained as a primordial condition or as a
retrogression in the direction of older types.

The suspension of the shoulder-girdle in our com-
mon Eel has a certain resemblance to the analogous
arrangement in the Rays and Sharks, though the sus-
pensory bones do not correspond. The posttemporal
bone as well as the post-clavicular is wanting, the girdle
itself consisting on each side only of the clavicle and
supraclavicle. The latter of these (tig. 265, set), with
its flattened, more or less distinctly forked top, hangs
loose, embedded in the outermost layer of the dorsal
half of the great lateral muscle, but the clavicle (fig.
265, cl), which retains its ordinary relation to this
muscle, is attached by means of an horizontal tendon
(a part of the aponeurosis between the dorsal and
ventral halves of the lateral muscle) to the sides of
the fifth and sixth abdominal vertebrae6. The shoulder-
girdle is similarly suspended, behind the head, in the
said cartilaginous fishes; but an essential difference con-
sists in the absence of both clavicle and supraclavicle
in the last-mentioned forms, the suprascapular part (the
upper parts of the primordial shoulder-girdle) being
attached to the upper side of the spinal column. In
the Eel this resemblance to the cartilaginous fishes is

destitute of osseous connexion with the auditory apparatus,
arch without symplecticum. Palatine arch and maxillary
Ventral fins none.

consequently of a secondary origin, probably connected
with the great development of the branchial cavity,
which has forced the shoulder-girdle backwards, loosen-
ing its attachment to the cranium, and causing the
disappearance of the posttemporal bone. The girdle is

Fig. 265. Half of the shoulder-girdle in Anguilla vulgaris , seen from
in front, 3/2 times the natural size, scl, left supraclavicle; cl, left
clavicle; sc, left scapula; cr, left coracoid; b, brachial bones of the
left pectoral fin.

completed below by the ligamentous connexion between
the two clavicles, which are curved at an obtuse angle;
and this lower part of the girdle is joined by a long
muscle to the short urohyoid bonec (fig. 268, A and
fig. 271, B and C, uh), which is directed backwards,

a Gr. ey%ehv g, eel, and fiogcpf form.

b In our Conger the shoulder-girdle is suspended in the same manner and at the same point, but still more superficially, with the
upper parts nearer to the skin. The posttemporal bone is thus wanting there as well; but a row of cartilaginous ducts, belonging to the
lateral line, runs under the skin from the head (the temples) to the supraclavicle.

c In our Conger the urohyoid bone is longer, but preserves the ordinary Teleosteous type, being flattened only at the base, where it
articulates with the basihyoid bones (coalescent, as in the common Eel, with the ceratohyoids), and furnished with a raised median keel on its
Tipper surface. Its posterior extremity is more or less deeply divided into three spinous points (fig. 282, p. 1036).

1012

SCANDINAVIAN FISHES.

and has the form of a flattened cone. High up on the
superior arms of the clavicles are attached the scapular
disks. These are thin and in great part cartilaginous,
each consisting of a circular scapular part (fig. 265, sc),
with an angular incision above and a small, round hole
(scapular fenestra) below, and a semicircular coracoid
part (fig. 265, cr), both of them fitted into the mem-
brano-cartilaginous disk. A scapular disk as simple as
this belongs to the earliest stages of development in
other Teleosts; but a manifest relic of the morphological
alliance with more primitive piscine types appears in
the great number of supporting bones (brachial bones
— fig. 265, b) possessed by the pectoral fins of the Eels.
In almost all" the preceding Teleosts the number has
been 4; in our common Eel it is 7 or 8. Anna, a Gan-

the primitive forms. The loss of the ventral fins, on the
other hand, as we have seen in many of the preceding Te-
leosts, is, here too, secondary in its significance; and in
foreign Eels the pectoral fins may share the same fate0.

The skeleton of an adult Eel is firmly ossified, and
characterised by its numerous vertebrae ■ — generally
more than 100, sometimes about 160 — of almost uni-
form shape and with feebly developed appendages. In
our common Eel the anterior vertebrae, to the seventh
inclusive, have their shallow but long, almost contiguous
neural spines broken up into several (as many as 9)
spicules'*. The said vertebrae are furnished both with
upper" and lower-' transverse processes. The former are
alary spines, directed backwards and of fairly uniform
strength, which are, however, exchanged on the seventh

Fig. 266. Three abdominal vertebra; (44 — 46) in Anguilla vulgaris , seen from the left and magnified.
na, scleral bone (epinenral, cf. fig. 236, p. 947); psps, upper spinous process; ptrp, lower posterior transverse process {processus transuersus
posterior ); ptra, lower anterior transverse process; ha, haemal arch; pspi, lower spinous process {processus spinosus inferior)', pi, ribs.

oid genus, has the same number; and Polypterus, an-
other Ganoid genus, has 13 — 17 brachial bones. These
Ganoids have besides retained one or more bones be-
longing to the innermost (proximal) row of the prim-
ordial cartilage (Gegenbaur’s pterygium) of the pectoral
fins, which has entirely disappeared in the Teleosts.
During the evolution of the Teleost type the basal parts
of the pectoral fins have thus suffered reduction6; and
the Eels are most nearly approximated in this respect to

and following vertebrae for scleral bones, directed back-
wards and upwards in the dorsal half of the great lateral
muscle (fig. 266, na). The lower transverse processes
(ptra) of the first vertebra are only small, pointed pro-
tuberances. On the following vertebrae they grow
broader, but from the seventh vertebra inclusive they
become pointed spines, directed outwards, backwards,
and downwards, and bearing at the tips short and weak
ribs (pi), which in their turn give place towards the

a For exceptions see the Batrachoids (p. 133, above) and the Lophioids (p. 136). Ostracion, a Plectognate, has 5 basal bones.

5 Cf. Smitt, Ur de hogre djurens utvecklingshistoria, pp. 222 seqq.
c Cf. the Lophobranchs, see above, pp. 667 seqq.

d In the Conger this division is restricted to the formation of lateral grooves on the neural spines, but is perceptible in this form
even in the 13th vertebra.
e Diapophyses.
f Parapophyses.

EEL-FISHES.

1013

tail to scleral bones in the ventral half of the lateral
muscle. Throughout the greater part of the abdominal
region, from the 8th vertebra to the 44th inclusive, we
find in the common Eel another pair of lower transverse
processes (parapophyses, ptrp), behind the last-mentioned
processes and always smaller than these. On the 45th
vertebra, or sometimes on the 44th, this posterior trans-
verse process on each side of the bone disappears, or
is at least reduced more perceptibly than before; but
the anterior transverse process divides instead into two
(three) branches,' the anterior (outer) supporting the
rib, the posterior (ha), which is simple or double, curv-
ing inwards. On the 46th vertebra this posterior
branch springing from one side of the vertebra joins
the corresponding branch of the other side, to form

(an inner) transverse process, the former continued by
a row of transverse processes at the middle of the
sides of the caudal vertebrae (which row is wanting in
the common Eel), the latter by the row which curves
downwards and inwards to form the haemal canal of
the caudal region. The skeleton of the Eel, for all its
simplicity, thus shows with the greatest distinctness how
the vertebral appendages may be homologous in the
lower grades of differentiation, although, when the
differentiation is more advanced, they are different
both in origin and functions.

The tip of the tail presents the appearance of a
rather primitive (diphycercal) structure. Only the outer-
most (last) two vertebral — the outermost of which is
composed, at least in our common Eel, of two prim-

U t

Fig. 267. Skeletal parts of the tip of the , tail in Anguilla vulgaris , seen from the left side and magnified.

A: from a young Eel (Elver, Fr. Uivelle ), after Robin; B: from a full-grown Eel.
v, vertebra, originally the antepenultimate, subsequently the penultimate; p, its h temal arch; n, its neural arch; »>,, its spinal canal; u, origi-
nally penultimate vertebra, afterwards confluent with the last vertebra; i, primordial articulation between these vertebra*; h, hindmost haemal
arches, surrounding the cavity of the lymph-heart; £, termination of the spinal canal; r , lowest, s, uppermost ray of the caudal fin;

w, posterior expansion of the urostyle.

the closed htemal arch and its hannal spine ( pspi ). In
the Conger the said posterior parapophysis appears only
on a few vertebral in the hindmost part of the ab-
dominal region, and even there is not always present,
or is sometimes developed only on one side of the ver-
tebra?. But the parapophyses of the abdominal vertebra?
are broad, as in the Codfishes, and pierced at the base";
and in the last 9 or 10 abdominal vertebra? the anterior
parapophysis divides into an upper (outer) and a. lower

ordial vertebra? — enter into the structure of the appa-
ratus supporting the caudal tin. Although the vertical
tins of the Eels are, as a rule6, so confluent that no
separate caudal fin can be distinguished externally, the
presence of such a fin is indicated internally by the
circumstance that the hindmost fin-rays — 10, both in
our common Eel (fig. 267, r — s ) and in the Conger0 —
do not articulate like the others by means of a flat ar-
ticulary surface with separate interspinal bones, but

a In the Murasna the posterior parapophysis divides on the 25th vertebra, and from, the 73rd vertebra. inclusive its lower branch enters
into the structure of the closed haemal arch: — see Owen, Anat. Physiol. Vertebr., vol. I, p. 45.

* In some, however, as in Ophichthys , the tip of the tail is free, without any trace of caudal fin.
c In the Murrena 6.

1014

SCANDINAVIAN FISHES.

include within their cloven base the margin of the
hypural bones and urostyle. The spinal canal extends
straight back towards the end of the last vertebra ( t ),
the body of which is compressed to form the urostyle
(r), also directed straight backwards. The first (posterior)
hypural bone (h) is really composed, in analogy to the
primordial confluence of the last two vertebrae, of two
haemal parts, which have been united in an arch at the
base (proximally), just below the last vertebra, and have
closely applied their outer (distal) extremities to each

B

other, thus including an elliptical cavity, in which the
pulsating lymph-heart of the Eels is situated". The
lower (anterior) hypural bone (between p and r) an-
swers to the haemal spine of the penultimate vertebra,
which also supports the hindmost haemal arch (p). In
young Eels, as well as in the Conger, Robin found this
hypural bone to be fairly well developed6 and originally
divided by a suture from the haemal arch. In older
Eels (fig. 267, B) it is extremely narrow, and lies, in
the form of an osseous rod, close to the under margin

Fig. 268. Skull of an Anguilla vulgaris. 3/2 times the natural size. A: with branchial apparatus, jaw-bones, and orbit. B: cranium, seen
from above. C: the same, from the right; D: the same, from below; E: the same, from behind.
alsp, alisphenoid; bo , basilar part of the occipital; et, etv, and v, ethmoid (et) and vomer (v), confluent in front; fr, frontal; hm , hyomandi-
bular; iop , interoperculum; /, lingual; lo , lateral parts of the occipital; mdb, mandible; mt, mastoid part of the temporal; op, operculum; par ,
parietal; pob, preorbital; pop , preoperculum; psp, parasphenoid ; ptr , petrosal; qu, quadrate; Ii. br., branchiostegal rays; so, squamosal part of
the occipital; sob, hindmost part of the suborbital ring; sop, suboperculum; spet, supraethmoidal ; splio, postfrontal (sphenotic); squ, squamosal

part of the temporal; uh, urohyoid.

a In the Conger the upper (posterior) part of this hypural bone coalesces with the lower edge of the urostyle. See Robin, Journ.
Anat., Phys. 1880, pi. XXV, figs. 1 and 2.

b Cf. Robin, 1. c., pi. XXV, fig. 3 and pi. XXIV, fig. 4.

EEL-FISHES.

1015

of the first (posterior) hypural bone. A continued re-
duction would thus seem to obtain here; and the di-
phycercal caudal fin of the Eel, a character which would
else be the token of a low degree of evolution, rather
suggests a far advanced, retrogressive metamorphosis.

Among the peculiarities that characterize the skull
of the Eel, the first to attract attention are the com-
paratively great expansion of the parietal bones (fig.
268, par), which form almost half the roof of the cra-
nial cavity, and the excessively elongated squamosal
bones ( squ ), which by means of a long process on the
upper side of the skull extend forward to a line with
the hind extremities of the upper ethmoids ( et and etv ),
and at about the middle of their length support the
postfrontals (spho). The last- mentioned bones rest below
on the petrosals ( ptr ), and have lost their importance
as a point of suspension for the hind extremity of
the suborbital ring. This (sob — pob), which in our
common Eel — as well as the prefrontals (its anterior
suspensories) and turbinals (spet) — is cartilaginous, is
attached behind on each side of the skull to a process
(fr in D), broken up into spines and originating from
the side of each frontal bone just before the anterior
extremity of the squamosal bone. The breaking-up of
this process into spines, whereby firmer muscular at-
tachments are secured, and which we have previously
seen in the upper spinous processes of the anterior
abdominal vertebrae, extends to the lateral occipitals
(lo) and the squamosals, that expand backwards like
wings, and, as well as the other bones in the lateral
walls of the skull, are characterized by a spongy tex-
ture. There is no orbito-muscular canal; and the true
sphenoid (basisphenoideum, bsp in fig. 282) is closely
superposed on the parasphenoid, with upward processes
towards the descending processes (fr below in fig. 268,
C) of the frontal bones and towards the alisphenoids
(alsp) behind them. There are no orbitosphenoid bones.

Great diversity of opinion still prevails as to the
composition of the palatine and maxillary arches — as-
suming that both these arches should really be re-
garded as distinct in the present series. In that case
each of them consists on each side of only a single
bone. The inner bone (fig. 269, pt), in the common
Eel of a thin and oblong, lanceolate form, often im-
perfectly ossified in front, joins the oblique hyoman-
dibular (Jim) and the quadrate (qu) to the anterior

“ Zeitschr. Gres.' Naturw. 1867, p. 270, taf. I, fig. 1 («).

part of the parasphenoid (fig. 268, psp). In the Con-
ger it is expanded and bifid behind, one branch meeting
the hyomandibular, and the other forming the con-
nexion with the quadrate. The outer bone (fig. 269,
mp) lies anteriorly and throughout the greater part of
its length in the same plane a"s the inner, and is there
united by ligaments with the latter, the two bones thus
forming on each side a palatine roof continuous with
the parasphenoid bone. But the hind extremity of the
outer bone is twisted and bent downwards, and is folded
outside the mandible, being united by strong, but loose
bands of sinew to the outside of the coronoid process
of the latter. In the common Eel this end is pointed,
in the Conger flattened, somewhat expanded, and in-
serted between the outer and inner bands of sinew by
which the great cheek-muscles are attached to the man-
dible. In the latter species the hind extremity of the
outer bone also glides on the outside of the coronoid
process of the lower jaw, by means of a flat and car-
tilaginous, articulary surface on each bone. In the Mu-
rama, according to Jacoby", it is joined by a special
band of sinew to the outside of the quadrate. The an-
terior extremity of the outer bone is tightly articulated
with the compound bone formed by the coalescence of
the vomer and the ethmoid (fig. 268, et, etv , and v ),
a striking resemblance to the corresponding connexion
between the intermaxillaries in the Pike, where we
found the said bones (fig. 253, pmx) similarly furnished
with a flat, ascending process. But here these processes
(fig. 268, A) are so closely applied to the constricted
and carinated anterior extremity of the ethmoid that
they almost meet above, behind the expanded head of
the vomer (etv, to the right in B). This connexion
reminds us most of the ordinary position in the Teleosts
of the palatine bones. The analogy between the outer
bone and a palatine is further strengthened by the
disposition of the teeth. In most of the fishes we have
hitherto seen furnished with teeth both on the vomer
and the palatines, the transverse row of teeth on the
anterior extremity of the first-mentioned bone (the
head of the vomer) forms an arch together with the
palatine teeth, and here too the teeth are so arranged.
The long and narrow vomer ( v ), the posterior end of
which extends on the under side of the parasphenoid
bone a little beyond the orbits in the common Eel, is
armed underneath and in front, for rather more than

Scandinavian Fishes.

128

1016

SCANDINAVIAN FISHES.

half its length, with an unbroken0 card of teeth, which
are straight or slightly recurved at the tip, in young
Eels pointed, in old more obtuse. In the extreme front,
at the tip of the snout, the bone has an expansion,
like a vomerine head (fig. 268, D), and behind this
point it is constricted. The form of the dental card
follows that of the bone. Into the said constriction on
each side is fitted the outer bone with its card of teeth,
which resembles in composition the vomerine card, and
extends, gradually narrowing behind, to the very cor-
ner of the mouth, working against an exactly similar
card on each half of the mandible.

It is easy to perceive that the structure of the
palatine roof and the upper jaw may give rise to the
most conflicting interpretations. That a comprehensive
reduction has taken place, is obvious. The inner bone

Fig. 269. Bones of the palatine roof and jaws in Anguilla vulgaris.
3/9 times the natural size.

hm, hyomandibnlar ; hma, its anterior articulary process; limp, its
posterior articulary process, with the end of which the operculum ar-
ticulates; qu, quadrate; pt, entopterygoid ; mp, palato-maxillary ; mdb.
mandible; art, its articular part; cl, its dental part.

(pt), which serves to connect the hyomandibular and
quadrate with the anterior part of the parasphenoid
and with the bone which we have hitherto called the
outer bone, cannot be explained as anything but a pte-
rygoid. Such is the position occupied on each side of
the palate in all the Teleosts — where these bones are
distinct from each other — by the inner pterygoid ( en -
topterygoideum 1. mesopterygoideum ) and the posterior
pterygoid ( metapterygoideum ), the latter of which has
coalesced in the. Eels, according to Owen* 6, with the
hyomandibular. The outer bone (mp) has received three

interpretations: as a palatine (1), a maxillary (2), and
an intermaxillary (3).

The first-mentioned interpretation (1), which was
suggested by Owen0, presupposes that the maxillaries
as ivell as the intermaxillaries have disappeared in the
process of reduction, and that the tip of the snout is
formed, as in the Pike, by the ethmoid and vomer.
Tendencies to such a reduction we have seen above,
e. g. in the maxillaries of the Glanomorphs and in the
intermaxillaries of Argentina. The hind extremity of
the assumed palatine occupies a peculiar position, it is
true, in relation to the lower jaw; still it is not so
superficial and free as the extremity of the maxillaries
in other, more typical Teleosts, and it is also joined,
according to Jacoby, by a band of sinew to the quad-
rate bone.

The second explanation (2) — according to which
the margin of the upper jaw is formed on each side by

Fig. 270. Opercular apparatus of Anguilla vulgaris. X s/2.
op, operculum; pop, preoperculum; sop, suboperculum; iop, inter-
operculum.

the maxillaries — was first given by Rosenthal'4 and
Meckel0, and has subsequently been adopted by Peters
and his pupil Jacoby in Berlin, by Brattstrom7 and
his teacher Lilljebobg, and others. It is based on the
presumption that palatines are wanting, and that the
intermaxillaries, in their confluence with the ethmoid
and vomer, form the tip of the snout. The relation
of the assumed maxillary to the pterygoid and the ho-
rizontal expansion inwards of its anterior part, how-
ever, do not agree with this interpretation, nor can the
assumed coalescence of the intermaxillaries and the
other bones at the tip of the snout be regarded as by
any means fully demonstrated.

a The gap mentioned by Jacoby (1. c., p. 291) I have not been able to detect, even in young Eels 66 mm. long.

6 Compar. Anat.. Physiol. Vertebr., vol. I, p. 122.
c 1. c., pp. 113 and 118.
d Tchthyotom. Tafeln , taf. XXIII, i* .
e Syst. Vergleich. Anat., Bd II, p. 356.

■' Om krauiet och skuldergordeln hos Murcona anguilla , Lin., disp. Ups. 1875, p. 17.

EEL-FISHES.

1017

The third interpretation (3) — according to which
the margin of the upper jaw consists laterally of the inter-
inaxillaries and anteriorly of the vomer (there coalescent
with the ethmoid) — was first- proposed by Cuvier",
and has long been the most generally accepted. It pre-
supposes that true maxillaries and palatines are wanting.

The history of the Eels’ development has not yet-
pronounced judgment on these three opinions; and a
fourth explanation might reasonably be suggested, name-
Iv that the palatines and maxillaries were originally
independent, but have coalesced, as Meckel and, after
him, Peters assumed of the relation between the inter-
maxillaries and the ethmoid and vomer. From the
evolutional history of the higher vertebrates we know6
that the maxillaries as well as the palatine roof are
primordial excrescences of the mandibular arch, and
that the incipient palatine roof may appear as a se-

Fig. 271. Hyoid arch of Anguilla vulgaris. X 3/2.

A: hyoid arch with branchiostegal rays; B: lingual hone; C: the same,
seen from below.

/, lingual; uh, urohyoid; bbr, first copula (basi-branchial); cell, cera-
tohyoid; eph, epihyoid; 11. hr ., branchiostegal rays.

condary growth in an inward direction, originating from
the maxillary arch. Whether an equally intimate rela-
tion between these parts has possibly arisen during their
reduction in the Eels, is a question which only the his-
tory of evolution can solve. We accordingly consider
it still doubtful whether intermaxillaries and maxillaries
are wanting in the Eels or have coalesced with more
internal bones, and in the present state of the question
the describer is fully entitled to speak of maxillary
and vomerine teeth.

In conjunction with the great development of the
branchial cavity the Eels have very broad branchiostegal

membranes and long branchiostegal rays (B. hr.). But
the former are united throughout by far the greater
part of their margins to the skin and joined to each
other, thus leaving only small, slit-like gill-openings;
and all the latter, except the uppermost (hindmost) ones,
are slender, filamentous, and coiled backwards, upwards,
forwards, and finally downwards, so as to include within
their curve both the operculum and the suboperculum.
In the Conger (fig. 282, p. 1036) the last two bran-
chiostegal rays of each membrane are expanded at the
top; in the common Eel (fig. 271) the expansion is con-
fined to the last ray, but is so great that the ray is
sometimes equal in breadth to the comparatively nar-
row, obliquely semi-elliptical operculum (fig. 270, op).
The sabre-shaped — in the Conger (fig. 282) elongated
triangular, posteriorly pointed, but curved — suboper-
culum (sop) follows the curved inferior margin of the
operculum, and is weaker both in the common Eel and
the Conger than the triangular, anteriorly pointed inter-
operculum ( iop ), which is even larger than the thin,
crescent-shaped preoperculum (pop).

The Eels are exceedingly voracious; and their di-
gestive canal is accordingly rather short and simple.
Our common Eel may serve as an example. The main
abdominal cavity of this species varies considerably in
length, between about- 26 and 30 % of the length of
the body; but- in the caudal part it has on each side
a prolongation, of which we shall give a- fuller descrip-
tion when treating of the sexual organs. The thin-
walled and rather wide oesophagus passes into the sto-
mach without a break, except that the longitudinal
folds (ridges) in which the mucous membrane lies on
the inside become thicker and higher in the stomach,
while the wall of the stomach is also thicker than that
of the oesophagus. Straight backwards the stomach is
prolongated in the form of a cone; but from the right
and lower side of its anterior part it sends out a pyloric
portion, with equally thick walls, which runs forward
to the hind margin of the liver, where it bends abruptly
in a narrow crook, so that the beginning of the intestine,
which runs backwards, lies close to its under surface.
There are no ceecal diverticula. At the middle of the
crook is a funnel-like valve (an annular fold on the in-
side), marking off the pylorus from the intestine; and
on the inside of the latter the mucous membrane is
disposed in a honeycombed network of deep, confluent

a Mem. Mus. cl' Hist. Nat., tome I, p. 118.
h Cf. Smitt, Ur de hogre cljurens utvedclingslnstoria, p. 167.

1018

SCANDINAVIAN FISHES.

folds. Beside the termination of the stomach the in-
testine forms a curve of varying size, with one or more
small convolutions, before passing into the rectum, from
which it is divided internally by a more or less distinct,
annular fold". The liver is generally almost entire,
having only a more or less deep incision in the hind
margin. Its length is about 1/i of that of the abdomi-
nal cavity proper, somewhat more than V2 of that of
the stomach, or 3/4 of the distance between the termi-
nation of the stomach and the vent. Sometimes, how-
ever, its length is equal to the last-mentioned distance.
Its upper, concave side surrounds the under side of the
oesophagus and the beginning of the stomach, as well
as the gall-bladder, which is rather large and lies to
the right. The spleen is oblong, rather more than half
as long as the liver, and is situated on the right side,
at the fold between the pylorus and the intestine. On

and Maggic as testes. The air-bladder varies consider-
ably in length, between V2 and 1/3 of that of the ab-
dominal cavity proper. • It is of a pointed fusiform
shape, but in front, for almost half its length, is divided
into an upper and a lower cone, the latter joined by
the pneumatic duct to the oesophagus, though in the
specimens examined by us the communication has in-
variably been closed. The kidneys lie, as usual, close
below the spinal column, above the air-bladder and
without the peritoneal cavity. They extend not only
throughout the length of the abdominal cavity proper,
but also into the backward continuation of the haemal
canal, in the caudal region, above the prolongations of
the abdominal cavity, to which structures we shall re-
turn in our description of the generative organs. The
urinary bladder lies just behind the downward curve
of the rectum towards the vent, and has its orifice just

Fig. 272. Abdominal viscera of Anguilla vulgaris. After Syrski. A, female; D, male.

A: a , right, b, left ovary; c, right, d, left prolongation' of the ovary in the cando-abdominal cavity; e, septum (prolongation of the mesen-
tery) between the caudo-abdominal cavities; f, vent; g , urinary bladder; //, fold of adipose membrane on the right side of the intestine; 7/,
fold of adipose membrane on the stomach; i, fold of adipose membrane on the left side; k, stomach; 7, pylorus; m, liver;

n, gall-bladder; o, pectoral fins.

B: a, right, b , left testis; c, right, d, left prolongation of the testis in the caudo-abdominal cavity; e, septum between the caudo-abdominal
cavities; /, efferent duct of the testes ( vas deferens)', g, seminal vesicle; h, vent; i, urinary bladder, mainly concealed by the seminal vesicle;
/<:, k', and l, corresponding to h, h' , and i in A (see above); m, stomach; n, pylorus; o, liver, turned up to show how it is applied to the

oesophagus and stomach; p, gall-bladder; q, pectoral fins.

the left side of this fold, as well as on both sides of
the intestine and rectum, hangs a lobulated peritoneal
fold, which has been developed into an omentum-like
fold of adipose membrane, consisting of connective-tissue
meshes filled with fat-globules. In appearance these
folds are so like generative organs that they have been
described both by Ercolani* * 6 and by Balsamo-Cravelli

a The honeycombed mucous membrane of the intestine and

part, is apparently subject to considerable variations.

6 Mem. vAccad. Sc. Bologna, 1872, p. 529.

c Mem. 1st. Lomb. Sc., Lett. Milano, XII, 1872, p. 229.

behind the anal aperture. In our common Eel it has
the form of an equilateral triangle, with one angle (the

s

so-called neck) directed downwards and passing into
the urogenital aperture.

The generative organs are suspended on each side
of the abdominal cavity and throughout the greater
part of its length, side by side with and outside the

rectum, as well as the thickness of their walls and that of the pyloric

EEL-FISHES.

1019

above-mentioned folds of adipose membrane, the right
beginning farther forward than the left, but not ex-
tending so far back. The structure and significance
of these organs, especially in our common Eel, has
long been an obscure question, which was not eluci-
dated until recent times.

From the preceding pages (p. 829) we know that
the Salmonoid family and some other fishes, among
them the family of the Eels, are destitute of oviducts,
at least closed ones, such as appear in the remaining
Teleosts, and that the eggs of these fishes fall, when
ripe, loose into the abdominal cavity, whence they are
expressed through a more or less developed peritoneal
canal on each side, opening into the genital pore just
behind the vent or into the urogenital aperture com-
mon to the genitals and the urinary bladder. The
testes, on the other hand, are furnished, even in the
Eels, with efferent ducts ( vasa deferentia), which have

tsc rc tdc ura tel vd d

hand, they occupy as caudo-abdominal cavities the same
relation to the skeleton as the abdominal cavity proper.
The peritonea] fold in which the intestine is suspended
( mesenterium ) crosses from the intestine to the urinary
bladder, which lies behind the rectum — leaving in the
females a triangular opening (peritoneal canal) between
the lowest part of the rectum and the neck of the
bladder — and is continued backwards in the middle
of the haemal canal (or a little obliquely). It thus forms
a partition wall between the two caudo-abdominal ca-
vities, suspended from a continuation of the peritoneum
proper, which lines the walls of these cavities and di-
vides them from the superposed prolongations of the
kidneys (caudal kidneys). The most striking peculiarity
in the generative organs of the common Eel is, how-
ever, the duplication of the parts contained in the
caudo-abdominal cavities, each organ consisting there
of two blades running side by side.

ra t vd

t

B

Fig. 273. Portions of the testes in Anguilla vulgaris. A , abdominal cavity (posterior part) and caudo-abdominal cavity, opened and with
the walls folded back to show the enclosed organs; natural size; after Brock. B , a portion of the young testes, at an early stage of deve-
lopment, in an Eel 23 cm. long, taken at Trollhattan in 1848; magn. about 8 diam., to show how the testicular lobes ( t ) originate in a thin

and transparent mesorchial fold (at first of uniform breadth) of the peritoneum.

ts, left, tel, right testicle in the abdominal cavity; tsc, left, tdc , right testicle in the caudo-abdominal cavity; vds , left, veld, right testicular
duct ( vas deferens)', vs, seminal vesicle; vu, urinary bladder; ra, abdominal, rc, caudo-abdominal divisions of the kidneys; ura , abdominal,
lire, caudo-abdominal urethra; r, rectum, detached and laid to the left; ved, vena cava dextra; cl, cloaca (emits).

their common aperture, as usual, in the anterior wall
of the neck of the urinary bladder. Characteristic of
the majority of the Enchelymorph genera, but most
developed in our common Eel, are two backward pro-
longations of the abdominal cavity and the continua-
tions in these cavities of the generative organs, one
below each side of the caudal (post-anal) prolongations
of the kidneys. We have indeed seen a similar pecu-
liarity in the Flounders, though the secondary abdo-
minal cavities are there situated outside the hasmal
canal (see above, pp. 370 etc.). Here, on the other

The ovaries have the appearance of frilled bands;
but their inner (median) side is smooth, and the ap-
pearance of frills is caused by elevated, transverse (ver-
tical), simple or double, foliate lobules (ovarial lamella?,
in which the eggs are developed) on the outer (lateral)
side, though so arranged that in the caudo-abdominal
part of the ovary the inner blade has these lobules on
its inner side, and turns its smooth side outwards, to
face the outer blade of the same ovary. The testes, on
the other hand, so long as their real nature was un-
known, bore the name, conferred on them for their

1020

SCANDINAVIAN FISHES.

appearance, of l-obulate organs (Sw. flikorgan , Germ.
Lappenorgan). They consist of a row of rounded la-
mella;, coherent to each other just at the base, and
suspended from the efferent duct ( vas deferens).

In their internal structure the ovaries show a mesh-
work of connective tissue, so closely packed, when they
are unripe, with strongly refractive fat-globules, that it
is difficult enough to distinguish among these the ex-
ceedingly small eggs, which in the common Eel usually
measure during the summer months at most about 1/10
mm. in diameter, and late in autumn or even in De-
cember attain a size of about 1/i mm. The testes are
essentially similar to the ovaries in the texture of their
stroma (connective-tissue fold), but are without adiposis,
being consequently more transparent and difficult of
detection. Most easily perceptible is their white efferent
duct ( vas deferens). Their seminal cells (spermatogonia)
are also exactly similar at first to newly-formed ova,
and remain extremely small, measuring 0.015 — 0.03
mm. in diameter; but they multiply by fission as they
pass from the germinal epithelium into the stroma,
where they gather in their special cavities (spermatic
ducts). In the Conger and Mu ram a the development of
the spermatozoa has been traced; in the common Eel
spermatozoa have not yet been observed.

Some Enchelymorphs, among them probably our
common Eel", develop without any great metamorphosis.
Others, such as the Conger, have a larval existence dur-
ing which they are scarcely recognisable. These larva;
Avere first described under a generic name, Leptoceplia-
lusb or Hehnichthysc , and were long regarded as types
of a distinct family Leptocephalidce or Helmiclitkyidce)
or even of an order ( Lemniscatf ). The structure of
these larvm was described by Kolliker in 1852b They
are thin (ribbon-shaped) or more terete (cylindrical),
and so transparent that only the black eyes with sil-
very iris betray their presence, as they move Avith sIoav
and languid undulations of their Avhole body, sometimes

near the coast, sometimes in the middle of the ocean.
The head is extremely small — hence their name7 —
and the skeleton most feebly developed, Avith hardly
any signs of vertebras, these being confined, as a rule,
to the hindmost part of the body. The notochord
( chorda dorsalis), entire or only marked Avith annular
constrictions, advances Avithout a break into the carti-
laginous mass of the skull, and lies in the body, en-
closed, together Avith the spinal cord and the great
blood-vessels, in a mucous mass. Avhich separates these
parts from the surrounding musculature. The muscular
mass is fastened to the skeleton only in front (at the
head) and behind (at the tail), being attached elseAvhere
to the skin; but it is divided, as in other fishes, by
the zigzag transverse bands ( myocommata ) into flakes
{myomeres), each Avith an angle (directed forwards)
at the middle of the sides of the body, another (di-
rected backwards) at the back, and a third at the
ventral margin. The mouth is armed, both in the
upper jaAv and the loAver, Avith comparatively large and
strong, rather scattered, straight arid pointed teeth, set
in a single toav and directed obliquely fonvards. Each
nasal cavity is a depression in front of the eye, Avith-
out proal nostril. The tubular abdominal cavity ex-
tends along the greater part of the ventral margin.
The straight digestive canal includes, according to Kol-
liker, a caecum-like stomach, Avith two upward, caecal
diverticula and two similar appendages belonging to
the intestine. The liver is faintly developed; spleen,
air-bladder, and sexual organs are wanting. From this
larval structure Facciola5' has succeeded in tracing a se-
ries of transition-forms that bring it to perfect identity
Avith the young of the Congers, a metamorphosis at
which Gill7' had already hinted. The most singular
point is, hoAvever, that these larvae, retaining the struc-
ture proper to their first stage of development, can
attain dimensions considerably greater than those of
the stages in Avhich the terete type and internal struc-

a In the Ball. Accad. Gioen. Sc. Nat. Catania (Fasc. 34 — 35; Nov. 1893 e Genn, 1894, p. 4) Gbassi and CalandRticcio have ex-
pressed the supposition that the ova of Anguilla vulgaris are to he found floating in the sea, and that Leptocephalus brevirostris is the
larval stage of this species.

b Ghonovii Zoophylac., Fasc. I (1763), p. 135. Morris, an acquaintance of Pennant’s, had found these fishes in St. George’s Chan-
nel, off Holyhead, North Wales.

c Rakinesque, hid. Ittiol. Sic., where the name is, however, written Helmictis.
d Richardson.

e Zeitsclir. f. Wiss. Zool., Bd IV, p. 360.
f From Gr. Xercvog, fine, small, and VXqtaXf head.

■' II Naturalista Siciliano, Anno XII (1893), p. 194.
h Proc.. Acad. Nat. Sc. Philad. 1864, p. 207.

EEL-FISHES.

1021

ture of the species may be recognised without difficulty.
Gunther mentions Leptocephaloids as much as 10 Paris
inches (270 mm.) long"; and Facciola found larva) of
the common Conger in the first stage of development
(Leptocephalus incequalis) that measured 76 — 140 mm.,
while almost typical fry of the same species were only
75 mm. long. Gunther also considers it probable that
not all these larvae follow the normal course of deve-
lopment and become Congers, but that some of them
— perhaps when they are borne by the current or some
other agency out into the open sea or too far from the
coast where the species develops under normal condi-
tions— may lead a pelagic, but morbid (hydropic) life,
increasing in size, though never attaining the typical
structure of their species or propagating themselves.
Among the Leptocephali of the Royal Museum we find
a specimen, otherwise uninjured, and taken in the
middle of the Atlantic, south-west of the Azores (Lat.
31° 13' N.; Long. 35° 46' W.), that has five vesicular

dubitable and 63 doubtful species; in 1883 Jordan and
Gilbert d estimated the number of the species then
known at about 280. The great variability of form
indeed deprives any such estimate of conclusive author-
ity; but the number of species is at all events con-
siderable enough to necessitate the arrangement of the
series in several divisions. Bleeker established six
families within the series. Gunther was of opinion
that, considering the transition-forms and the varia-
bility of the Enchelymorphs, the whole series might be
included within one family {Murcenidm) ; but at the
same time he established ten subfamilies, the majority
of which might well lay claim to the rank of distinct
families. The Scandinavian fauna contains only two
members of one among these families; but for the
European fauna several of them possess interest. On
the English coast there occurs, though seldom, the Me-
diterranean Mu ram a (Mur am a helena), which we have
mentioned above, the type of the Gymnothoracida (Blee-

Fig 274. Leptocephalus Morrisii (?), from the Atlantic;

tumours — two of large size, occupying the whole
depth of the body, and three smaller ones, in the dor-
sal region — full of fluid and with the myomeres more
or less atrophied. Sickliness and an abnormal manner
of life might well explain, as Gunther has pointed out,
the inconstancy both of form and structure that marks
these larvae. From the Mediterranean 35 species of the
so-called genus Leptocephalus have been described* 6, but
without evidence to show that the specific characters
are more than individual variations.

The Enchelymorph series is fairly rich in forms.
Bleekerc had examined in 1864 more than 250 species.
In 1870 Gunther adopted in his Catalogue 227 in-

a Cat. Brit. Mus ., Fish., vol. VIII, p. 143.

6 Carus, Prodr. Fnce Medit., vol. II, f>. 546. But Carus re
species distinct* vix describi possint.”

c Ail. Ichth. Lid. Orient. Ne'erl., tome IV, p. 5.
d Bull. U. S. Nat. Mus., No. 16, pp. 355, seqq.
e GOnther ( Deep Sea Fauna , Chall. Exped.) knew 14 such s]
36, exclusive of the Leptocephaloids.

taken in Lat. 31° 13' N. ; Long. 35° 46' W. Natural size.

ker’s Gymnothoracoidei, Gunther’s Murcenidce engy-
schistce, with narrow slits between the branchial arches),
a family very rich in forms, especially in India, and with
naked body, laterally compressed, and no pectoral fins.

The Enchelymorphs live both in fresh and salt
water; but most of them are marine fishes. A great
number are deep-sea forms6 — some of them taken in
more than 4,000 fathoms of water ■ — and show reduc-
tions still more comprehensive than those we have
described above, the piscine type consequently appear-
ing in such simplicity that it is sometimes difficult
to decide in which order the forms have their right-
place.

iarks with regard to these species, “Not* specific* tantum variant, ut
icies, and Vaillant {Exped. Scient. Travailleur, Talisman ; Poissons )

1022

SCANDINAVIAN FISHES.

Fam. ajvguillidje.

Gill-slits ( between the branchial arches) wide; gill- openings (■ in the skin ) widely separated. Heart situated just
behind the branchial cavities. Vertical fins well-developed and confluent round the tail. Pectoral fins present.
Anal fin separated from the tip of the snout by a distance of more than twice the length of the head. Posterior

nostrils distant from the margin of

Among Gunther’s Platyschistce some (the Synaplio-
branchidce ) are characterized by the downward removal
of the gill-openings towards the ventral side and their
union. Others (the Ptyobranchidce ) are marked by so
great a prolongation of the forepart of the body that
the heart is situated far behind the branchial cavity.
Others again (the Nemich thy idee) have the vent far ad-
vanced, close to the gill-openings or separated from
them by a distance less than the length of the head,
and the beginning of the anal tin just behind this
point. In others (the Saccopharyngidce “) the mouth

the upper lip. Tongue with free tip.

and pharynx are developed into a monstrous, funnel-
shaped gorge, so large that it exceeds the abdominal
cavity in length. Another division (the Ophichthyidce
and Myridce, the former with Unless tip of the tail)
are distinguished by a singular downward removal of
the nostrils, sometimes quite to the margins of the
upper lip. Some (the Murcenesocidce) are without free
tongue; others (the Heterocong rides ) are similar in all
these respects to our common Eel, but destitute ot
pectoral fins. The rest compose the family that has
its most marked type in the

Genus ANGUILLA.

The slimy body furnished with scales immersed in the skin.

By this solitary character it is always possible to
recognise one of the so-called Fresh-water Eels b, all
other distinctions being far too inconstant to be im-
plicitly relied on. Another character, which is not
peculiar, however, to Anguilla , though within the Scan-
dinavian fauna it is sufficient to denote the genus, and
besides has the advantage of being more conspicuous,
consists in the fact that the tip of the snout falls short
of the point of the lower jaw, or at least does not
project beyond it.

“An infinite number of species,” says Gunther,
“have been described; but most are so badly charac-

terized, or founded on individual or so trivial cha-
racters, that the majority of ichthyologists will reject
them.” Kaufc, who was the greatest authority on
this head until Gunther’s time, adopted no less than
49 species; Guntiikr'z reduced the number to 23. Da-
reste* acknowledged only 4 or possibly 5. As Gun-
ther has remarked-'’, the genus is known throughout
the Tropical and Temperate Zones, with the excep-
tions of South America, the west of North America,
and Western Africa. In Scandinavia and the rest
of Europe only one species is recognised at the pre-
sent day.

° For the Saccopharyngidce see G-ill and Rydeb (Proc. U. S. Nat. Mus., vol. VI, p. 262; vol. VII, p. 48), who founded a separate
order ( Lyomeri ) to receive this family, and Vaillant ( Exped . Scient. Travailleur, Talisman , Poissons, p. 193), who considered them to be as
nearly allied to the Anacanthines as to the Enchelymorphs. Among their peculiarities it should first be remarked that the number of the
branchial arches is 5 instead of 4, the usual number in the Teleosts.

6 Fresh-water or at least brackish-water fishes indeed occur within other Enchelymorph genera, as in Gymnothorax , Pisoodonophis,
Sphagebranchns , Moringua, and Murcenesox ; but in these genera salt-water Eels predominate, and they are besides entirely foreign to the
Scandinavian fauna.

c (Jatal. Apodal Fish., Brit. Mus. (1856), p. 32.
d Cat. Brit. Mus., Fish., vol. VIII, p. 23.
e Arch. Zool. Experim., tome IV, p. 224.
f Introd. Study of Fishes, p. 671.

EEL-FISHES.

1028

The
by a
what
29—

P. 17
Syn .

THE COMMON EEL (sw. alen).

ANGUILLA VULGARIS.

Plate XLV, fig. 1.

dorsal fin commences far behind the tips of the folded pectorals , being separated from the tip of the snout
distance of more than twice the length of the head , and precedes the anal by a distance about equal to ( some-
greater or less than ) the length of the head , the distance between it and the tip of the snout being about
33 %, that between the anal and the same point about 40 — 45 %, of the length of the body. The jaw-teeth
traight ( of an obtuse, conical form), or somewhat recurved at the tip, and set in quite continuous cards.

Li. br. 10 — 12«; D. ca 245— 275; A. ca 205— 225; C. 10;
— 206; Vert. 112 — 115s.

Murcena nnicolor; maxilla inferiore longiore, Art., Ichthyol.,
Gen., p. 24; Syn., p. 38 (ubi synon. vet. videantur): Spec.,
p. 66; Lin., Fna Suec., ed. I, p. 109.

Murcena anguilla, Lin., Syst. Nat., ed. X, tom. I, p. 245;
Fabr., Fna Groenl., p. 137; Bl., Naturg. Fisch. Deutschl.,
pt. Ill, p. 4, tab. LXXIII; Ahi. ( Ophichtus ), Spec. Ichtli.
Mur. Opliicht. (disp. Ups. 1789), p. 11; Retz., Fna Suec.
Lin., p. 311; Fab. {Murcena,), Fisch. 1st., p. 59; Pali..,
Zoogr. Ross. A*'., tom. Ill, p. 71; Ekstr., Vet. Akad.
Handl. 1831, p. 285 (= Mur. oxyrhina + Mur. platyrhina ,
p. 287); Nilss., Proclr. Ichthyol. Scand., p. 65 ( + Mur.
latirostris); Id., Skand. Fna, Fisk., p. 661; Sundev., Stockh.
L. Hush. Sallsk. Hand]., VI (1885), pp. 83, 92, 181;
Wdgrn, Landtbr. Akad. Hand]. 1858, pp. 181 et 210; Ny-
strom, laktt. Fnan Jemtl. Vattendr. (disp. Ups. 1863), p.
20; v. Bemm. in Herkl., Bouivst. Fna Nederl., part. Ill, p.
388; Olss., Ofvers. Vet. Akad. Forh. 1876, no. 3, p. 139;
1882, no. 10, p. 51.

Anguilla vidgaris, Turt., Brit. Fna, p. 87 ; Mitch., Trans.
Lit., Phil. Soc. N. York, vol. I, p. 360; Flmng, Brit.
Anim., p. 199; Costa, Fna Regn. Nap., Peso., pt. I, Mala-
cott. Apod., Anguilla, p. 52 (+ Angu. platyrhyncJius, p.
50, ex Cov., + Angu. acutirostris, p. 51, ex Yabr.), tabb.
LIII — LX; Blanch, Poiss. cl. eaux douces Fr., p. 491
(cujus formre: Angu. latirostris, p. 495, ex Risso, Ear.
Mer., tom. Ill, p. 199, + Angu. rnediorostris, p. 496, ex
Risso + Angu. oblongirostris, ibid., + Angu. acutirostris, p.
467, ex Risso, 1. c., p. 198); Sieb., Siisswasserf. Mittel-
eur., p. 342; Mgrn, Finl. Fiskfna (disp. Helsingf. 1863),
p. 33; Gthr, Cat. Brit. Alas., Fish., vol. VIII, p. 28
( + Angu. bostoniensis, p. 31, ex Lesueur, Journ. Acad. Nat.
Sc. Philad., vol. I, p. 81, + Angu. texana, p. 32, ex Kaup,
+ Angu. latirostris, p. 32, ex Risso, + Angu. Kieneri, p.
35, ex Kaup); Dareste, Arch. Zoo]. Exper., tom. IV (1875),
p. 217; Coll., Forh. Vid. Selsk. Chrnia 1874, Tillasgsh.,
p. 196; 1879, no. 1, p. 99; N. Mag. Naturv. Chrnia, Bd
29 (1884), p. 113; Winth., Naturh. Tidskr. Kbhvn, ser.
Ill, vol. XII, p. 50; Fedders., ibid., p. 93; Mop,., Hist.

Nat. Poiss. Fr., tom. Ill, p. 560; Bncke, Fisch., Fischer.,
Fisch z. O., W. Preuss., p. 173; Mela, Vertebr. Fenn., p.
357, tab. X; Day, Fish. Gt. Brit., Irel., vol. II, p. 241,
tab. CXLII, fig. 1; Mob., Hcke, Fisch. Osts., p. 143; Br.-
Goode, Fisher., Fisher. Industr. U. S., sect. 1, p. 630, tab.
239; Seeley, Freshw. Fish. Ear., p. 373 (+ Angu. eury-
stoma, p. 380, ex Hckl, Kn.); Lillj., So., Norg. Fna , Fisk.,
vol. Ill, p. 375; Reut., Sundm., Finl. Fisk, tab. XXXIII;
Car., Prodr. Face Medit., vol. II, p. 540.

Murcena rostrata (+ bostoniensis ) Lesueur, Journ. Acad. Nat.
Sc. Philad., vol. I, p. 81 (+ Mur. cirgentea + Mur. ma-
crocephala , p. 82); Dek., N. York Fna, part. IV, p. 312
(+ Angu. tenuirostris, p. 310); Jord., Gilb., Bull. U. S.
Nat. Mus., no. 16, p. 361.

Anguilla fluviatilis, Agass. (ex Cuv. ?), Rech. Poiss. foss., tom.
V, p. 130, tab. F, fig. 2; Hckl, Kn., Siisswasserf. Ostr.
Mon., p. 319 (+ Angu. eurystoma, p. 325).

Anguilla canariensis , Val. in Webb, Berth., lies Canar., Poiss.,
p. 88, tab. 20, fig. 1.

Anguilla japonica , Schleg. in Sieb., Fna Japon., Poiss., p.
258, tab. CXIII, fig. 2.

Anguilla callensis, Guich., Explor. Alger., Zool., V, p. Ill,
tab. 7, fig. 1.

Anguilla migratoria, Kr., Damn. Fisk., vol. Ill, p. 616
(+ Angu. acutirostris, p. 642, ex Yarr. + Angu. latirostris,
p. 656, ex Yarr.).

Anguilla, Kieneri, Kaup, Cat. Brit. Mus., Apod. Fish., p. 32
(+ Angu. Cuvieri + Angu. Bibroni, p. 33 + Angu. Sa-
vignyi + Angu. capitone, p. 34 + Angu. rnorena + Angu.
melanochir, p. 35 + Angu. marginata + Angu. microptera,
p. 36 + Angu. ancidda + Angu. rnediorostris + Angu.
altirostris, p. 37 + Angu. platycepjhala + Angu. latirostris,
p. 38 + Angu. acutirostris, p. 39 + Angu. nilotica + Angu.
cegyptiaca, p. 40 + Angu. callensis + Angu. canariensis,
p. 41 + Angu. novceorleanensis , p. 43 + Angu. tenuirostris
+ Angu. punctatissima + Angu. cabana, p. 44 + Angu. no-
vceterrce + Angu. texana, p. 45 + Angu. wabashensis, p. 46);
Carus, 1. c.

Anguilla Linnei, Malm, Gbgs, Boh. Fna, p. 590 (+ Angu.
latirostris, p. 591).

“ Sometimes 9, according to Lilljeborg.

,, 16, ,, ,, Kroyer.

c „ 116, „ „ Day.

129

Scandinavian Fishes.

1024

SCANDINAVIAN FISHES.

The Common Eel, with which most of our readers
are, no doubt, familiar, attains a length of about 1 x/2 m.
and a weight of about 5, perhaps 6 kilo." Its ordinary
length and weight, however, are respectively under 1 m.
and 2 kilo. The snake-like body is so terete in front
that the greatest breadth (thickness) is sometimes equal
to the greatest depth, or even (when the branchial ca-
vity is distended) greater than it; but the former is
commonly no more than i/5 or 3 * */4 of the latter, and
at the termination of the abdominal cavity the body
shows greater and greater lateral compression. The
greatest depth, which occurs just in front of the dorsal
tin, increases with age in the young, though even at a
length of 4 1/2 dm. it sometimes measures, according to
Kroyer, only about 57a % thereof; but the rule is that
it subsequently decreases during growth, falling in Eels
measuring 4 — 7 dm. from 7 to 6 % of the length of

length of the body increases from about 272 to 67a dm.,
that of the head decreases in our specimens from 13 to
11 % of the former6, or from about 41 to 36 % of the
distance between the dorsal fin and the tip of the snout.
But in this respect we find that older Eels — we can
speak here only of females — and certain individuals
which it has been proposed to treat as a separate spe-
cies or variety ( latirostris ), revert to the juvenile cha-
racters, or even exceed the limits of these, the length of
the head sometimes measuring 15 % of that of the body
or 46 % of the distance between the dorsal fin and the
tip of the snout. The form of the head is also highly
variable, in front more or less conical (acutir ostris) or
broader and depressed, wedge-shaped ( latirostris ), and
behind more or less tumid, depending partly on the
greater or less distension of the branchial cavity, partly
(behind the eyes) on the varying development of the

Fig. 275. Three varieties of Anguilla vulgaris. A, Anguilla latirostris; B, Angu. merliorostris ; C, Angu. acutirostris.

After Blanchere.

the body. Still large Eels grow to the thickness of a
man’s arm. The males seem always to remain smaller
than the females, for up to the present large specimens
have been found in every instance to be of the latter sex.

The head passes so gradually into the rest of the
body that the limit between them is externally almost
indistinguishable, being indicated only by the position
of the small gill-openings, just in front of and partly
below the pectoral fins. The size of the head is ex-
tremely variable. After the larval stage its relative
length (to the middle of the anterior margin of the
gill-opening) shows the usual diminution. While the

masticatory muscles. The usual proportions of the se-
veral parts of the head are such that the length of the
snout is 20 % or somewhat less (17 %), the longitudinal
diameter of the eyes 10 % or somewhat less (in young
Eels at least 13 in old sometimes 7 %), and the
postorbital length about 70 % (varying in individual
cases between 68 and 75 %c), of the length of the head.
The eyes, which are covered with a thin skin that still
further enhances the snake-like appearance of the Eel,
are sometimes set vertically, sometimes rather upturned.
Partly in consequence of this variation in the position
of the eyes, partly following the alterations of growth,

a According to the reports made to the Swedish Fisheries Commission of 1881 — 83, the Eel- — which occurs in every province of

Sweden — attains in Wermland, Westmanland, and Westernorrland a length of 5 Sw. ft. (148 cm.) and in the last-mentioned province a weight

of 15 Sw. lbs. (S'/g kilo.). According to Desmarest the Eel attains a length of at least 17 dm. and a circumference of at least 3x/2 dm.

6 According to Kroyer this percentage sometimes falls to 10.
c According to Kroyer sometimes nearly 77 %.

COMMON EEL.

1025

the young having, as usual, comparatively larger eyes
partly in connexion with the different environments and
habits of the fish, and partly owing to the above-men-
tioned retrogressions of development, considerable varia-
tions appear in the relative dimensions of the eyes. In
young Eels (sometimes even in specimens 372 dm. long)
and in the variety known as latirostris, the longitudinal
diameter of the eyes is about equal to the interorbital
width or at least 3/i thereof. The eyes are compara-
tively smallest, as a general rule, in the migrating Eels,
usually measuring less than 2/3 (sometimes only about
43 %) of the interorbital width. In the last-mentioned
specimens the interorbital width is also hardly less"
than the length of the snout, in the former sometimes
scarcely 3/4 thereof, though the percentage for this re-
lation increases with age. Each nostril in the pos-
terior pair lies, in the form of a dermal fissure, set
obliquely or in the longitudinal direction of the snout,
just before the upper anterior corner of each eye; the
anterior nostils are situated, in the form of dermal
tubes, one on each side of the tip of the snout. The
mouth is horizontal, with the tip of the lower jaw more
or less prominent — most distinctly in old Eels and in
latirostris — and with the corner situated below the eyes
or, especially in latirostris , somewhat behind the per-
pendicular from the hind margin thereof. The lips are
double, fleshy, and tumid, the outer folds being more
or less expanded at the sides, in the upper jaw as though
the free maxillaries of other Teleosts were here replaced
by this dermal growth; but the underlips are usually
broadest. The length of the upper jaw (of the cleft of
the mouth), from the tip of the snout to the hind mar-
gin of the buccal corners, measures about 26 — 33 %,
that of the lower jaw about 35 — 47 %b , of the length
of the head, the latter proportion in each case being
characteristic of latirostris. The dentition of the mouth
is partly described above. The most characteristic point
in the dentition of our Eel, as opposed to East Indian
and Australian forms otherwise very closely resembling
it, is the absence in the jaw-cards of the longitudinal,
toothless gap shown by the last-mentioned forms, though
the innermost row in both jaws, even in the European
Eel, is made up of smaller teeth, and may, as a rule,
be distinguished more easily than the others. The
tongue is fleshy, free, toothless, and elliptical, with a

small, short tip. Gill-rakers and p s eu d o b ranch i aa are
wanting. The fine and pointed pharyngeal teeth, weaker
than the jaw-teeth, form dense, velvety cards, two above,
of an oval shape, on the upper pharyngeals, and two
below, of a more elongated shape, on a thin inward
expansion of the lower pharyngeals, which else resemble
branchial arches. The upper half of the gill-openings,
which are slit-like or curved forwards in a crescent,
lies just in front of, the lower half below the base of
each pectoral fin, their height being equal to this base
and about the same as the interorbital width.

The dorsal tin is of about uniform height, though
low in front and very gradually increasing in height
behind, the increase being proportioned to the percep-
tible decrease in depth shown by the posterior third of
the body. The longest rays, which measure about V 2
— 2/ 5 of the greatest depth of the body, thus lie in the
hindmost part of the fin, just before it slopes to form,
together with the caudal and anal fin, a lanceolate tip
at the end of the tail. The anal fin is similar to the
dorsal, but much shorter, commencing, as we have men-
tioned, considerably further back, so that the distance
between the dorsal fin and the tip of the snout is about
70 — 75 % of that between the anal fin and the same
point. In Eels 21/2 — 7 dm. long this difference increases,
as a rule, during growth, until the body attains a length
of about 472 dm., but afterwards diminishes, the length
of the head thus being generally less in middle-sized
specimens, especially in the migrating Eels, than this
difference, in large Eels somewhat greater than the
same. But the individual variations appear to deprive
the difference of all systematic significance; nor does
it seem to possess any importance as an external sexual
character.

The pectoral fins, when expanded, are oval, often
somewhat pointed, and the middle rays (about the 10th)
are the longest. The relative length of these fins ex-
presses both a distinct alteration of growth, the young
having even relatively shorter pectoral fins than the
old, and a difference of sex, the males in general having
shorter pectoral fins than the females. In 6 Eels (4 cd
and 2 ?) 23 — 41 cm. long the average length of the
pectoral fins proved to be 3'6 % of that of the body
and 8"6 % of the distance between the anal fin and
the tip of the snout. In 7 Eels (all ?) 45 — 68 cm.

a According to Jacoby even greater.
6 According to Kroyer up to 54 /.

1026

SCANDINAVIAN FISHES.

long the corresponding percentages were respectively
4'3 and 10. In the males the former percentage varied
between 3'0 and 3*7, in the females between 3'8 and
4‘5. The latter percentage varied in the males between
7'5 and 8*6, in the females between 9 and 10‘9. That
this sexual difference — as it appears in the above
measurements — is fully constant, enabling us in every
instance to distinguish between males and females, is
hardly probable, for it coincides with a marked altera-
tion of growth; but it would seem at least to show
that the males retain the characters of youth longer
than the females®.

The thick and tough skin completely envelops even
the tins, but owing to the underlayer of lax connective

Fig. 276. Scale of Anguilla vulgaris. About 23 times the natural size.

tissue and fat is so loosely attached — as every kit-
chenmaid should know — that it can easily be stripped
off entire. The copious slime that covers it has ren-
dered the Eel’s slipperiness proverbial; and the system
of the lateral line is well developed. The lateral line
itself, which is straight, lying about half-way up the
sides, in front somewhat higher, so as to meet the
temples, is made up of comparatively scattered pores,
on the tail often indistinct, and not supported by spe-

cial scales, but usually marked off by their light hue
from the ground-coloration. The cephalic system of
the lateral line shows a transverse roAv of pores across
the top of the head behind the eyes, from which they
are separated by a distance about equal to the length
of the snout; and on each side runs a i*oav, usually
containing three distinct pores, straight fonvard from
the transverse toav, toAvards the superior margin of the
eyes, but not extending half-Avay thither. More distinct
are the pores on the snout, one roAv on the turbinal
bones and the suborbital ring, another on each half
of the mandible. The scales, which are entirely im-
mersed in the skin, are thin, flexible, and of a pro-
longated elliptical shape, someAvhat constricted at the
middle of their length, Avhich is about half the dia-
meter of the eyes. On the sides of the body their
arrangement is such that a certain number, usually
3—5, lie beside and contiguous to each other, obliquely
across the longitudinal direction of the body, and on
each side of this group lie others, the scales of Avhich
are set at about right angles to the former. Or each
group may be longer, containing up to a score of
scales, and forming a straight or curved row. Or the
scales may be juxtaposited one by one, in the said
zigzag arrangement. On the belly the scales are more
scattered, and more nearly approach to the longitudinal
direction of the body. The texture of the scales is
highly characteristic. When slightly magnified, the
entire surface seems to be composed of dense strings
of lustrous beads, all parallel to the margin. When
the scale is more powerfully magnified (fig. 276), this
conformation dissolves into a honeycombed network,
due to the union of the concentric striae (ridges) round
the elongated, central nucleus by radiating ridges of
equal elevation.

The coloration of the Eel varies Avith age, the
season of the year, the haunt of the fish, and partly
too Avith its sex. The dorsal side is of a lighter or
darker, greenish or grayish colour, the ventral yelloAv
or Avhite; the middle of the sides has a bronze lustre
of varying intensity. The colour of the fins usual ly
resembles that of the back, except in the case of the

a How futile it is to rely on the external sexual characters hitherto assigned in literature to the Eel, is most clearly shown by the
conflicting results at which different writers have arrived. According to Jacoby ( Fischf . Comacchio, Berlin 1880, p. 40) the males have
pointed snouts. From his measurements (p. 41) it appears that the breadth of the snout at the nasal ducts — (I assume that “Breite der
Schnauzenspitse zwischen den Nasaltuben” must be taken in this sense, for “between” the nasal tubes I have never found the distance so
great) — measured in 8 males (319 — 480 mm. long) less, in 8 females (313 — 480 mm. long) more, than 12 % of the length of the head.
According to Lilljebobg (Sv., Norg. Fisk., vol. Ill, p. 382) the males most commonly have blunt, “sometimes pointed” snouts.

COMMON EEL.

1027

anal fin, which in front shows the colour of the belly.
The yellow tint of the belly is characteristic of the
young and of the Eels that lead a somewhat stationary
life among the seaweed on the coast, or in the mud
and among the grass of the lakes. The white belly
belongs to the migration uniform, the spawning-dress
of the Eels®. The back is darkest, sometimes even
black, in the migrating males. Among colour-varieties
there have been observed light specimens, almost yel-
lowish green on the back, and irregularly spotted fish,
with light, clouded spots on the dorsal side or with
“dorsal streaks of a golden yellow” (Benecke). Albi-
nos have also been found6.

In the basin of the Atlantic the Eel is dispersed
between the latitudes of the West Indies and of Nor-
Avegian Finmark, and from the United States eastAvards
over North Africa and throughout Europe, including
the regions drained by the Baltic and the Mediterra-
nean, but, strange to say", originally with the exception
of the Black and Caspian Seas and their feeders. Fa-
bricius mentions it among the fishes of Southern Green-
land, and even Olafsen d kneAV it from Iceland, Avhere
according to Faber, hoAvever, its length seldom exceeds
U/2 ft-! but from the Arctic seas and rivers it is else
unknown. Within the basin of the Pacific it is found
in Japan, China, Formosa, Borneo, and, according to
Gunther, Ncav Zealand. The geographical range of the
Eel is thus one of the most extensive; but the gaps
— its absence, for instance, from the Avest coast of
North America — are difficult to explain.

In Scandinavia the Eel becomes rarer and rarer
inland and toAvards the north. Solitary specimens have
indeed been found in the Fjords of Varanger and Tana;
but into the innermost parts of Finland the Eel does
not penetrate, and in Stveden, as avcII as in Norway,
it seldom, if ever, ascends into the mountain-regions.
On the Norwegian coast it is common, according to
Collett, up to Lofoden, rarer on the coast of Finmark

further north, and in Exaavand, a lake in Bjellerud
(P rovince of Christiansand), it has been met Avith at a
height of 1,600 feet (500 m.) above the sea-level". The
Eel occurs in every province of Sweden, but to the far
north only on the seaboard. Widegren described in
1860 the Eel-fishery of the lower Lulea Elf and the
islands at its mouth. From the basin of the Angerman
Elf Trybom9 Avas told that Eels Avere found in Lake
Malgomaj, situated 356 m. above the level of the sea;
and according to Olsson the Eel ascends to the neigh-
bourhood of Gaddede, on Qvarnberg Water, near the
Nonvegian frontier. In the basin of the Ljunga, accord-
ing to Olsson, it has been found on rare occasions in
Herjeadalens Storsjo, 363 m. above the sea-level. In
Southern SAveden, as in Denmark, the Eel is common
in rivers, lakes, and meres, Avherever it can find a con-
genial haunt. Throughout the basin of Lake Wener,
however, it is said to have been wanting9, previous to
the construction of the Trollhatte Canal, Avhen the Eel-
fry made their Avay through the locks up the fall.

The Eel is tenacious of life and supple of body,
adapting itself to the most confined abode and the most
meagre circumstances; but in order to attain any con-
siderable size it requires abundant food and ample space.
It reminds us strongly of the snakes in its nocturnal
habits and its fondness for hiding in holes or burroAving
in the soft bottom. Like the viper it gathers in large
bunches — “ Icigger sig i vret,,h says the SAAredish fisher-
man. To these serpent-like traits is allied its poAver of
sustaining life for a long time out of the Avater: just as
the common snake is partly an aquatic animal, the Eel
can traverse considerable distances by land. Its serpen-
tine, wriggling movements enable it to make rapid pro-
gress though the water, so as speedily to find a hiding-
place; and its great poAver of muscle endoAvs it Avith
great endurance during its rovings in the sea ; but it
seems to rank among the sluggish fishes, and by nature
belongs to the ground-SAvimmers. “Slippery as an Eel,”

“ Cf. F. H. T. Leth: lakttagelser over Aalene, Dansk Fiskeritidende 1882, p. 393.

Feddersen, Dansk Fiskeritidende, 1891, p. 395.

0 The Eel’s original absence from the basin of the Danube — where it has been planted in recent times, and seems to thrive, having
voluntarily spread to the Black Sea and the lower courses of the Russian rivers — may perhaps be explained by a comparison witli the equally
singular presence in the Danube of the Huch (see Smitt, Riksrn. Salmonider , p. 148). The Eel is wanting in Siberia, being compelled to
enter salt water in order to breed, and incapable of enduring the cold of the Arctic Ocean, into which the Siberian rivers fall; and the same
ancient configuration of the ocean as enabled the Siberian Huch to spread to the basin of the Danube may have cut off the advance of the
Eel in this direction.

a Reise igiennem Island (1772), vol. I, p. 594.

e In the Swiss Alps, according to Fatio ( Fne Vert. Suisse, vol. V, p. 458), the Eel ascends to an elevation about twice as great.

. f Nord. Aarsskr. Fisk., l:ste Aarg. (1884), p. 306.

9 See Lloyd, Scand. Ada., vol. I, p. 143, and Nilss., Skand. Fna , Fisk., p. 675, note.

h Vret is probably connected with vrida , to writhe, twist. Tr.

1028

SCANDINAVIAN FISHES.

is a proverbial expression; and this quality is of great-
service to the fish in every strait. Nor does the Eel forget
to take advantage thereof, but slips through the tiniest
openings. In order to widen the aperture, when this
is too small to afford a passage for its head, it has re-
course to the same method as that employed by a
Fierasfer in gaining entrance to the body of an Holo-
t-huria (see above, p. 598, fig. 143). It first inserts the
end of the tail. From a live-well or other receptacle,
left without a lid, the Eel has often escaped tail first,
lifting itself over the edge with this part of the body.
The toughness of its skin is equalled by the obstinacy
with which it keeps stationary in the same spot, unless
impelled by hunger, the sexual instinct, or fear. It lies
motionless among the dense water-plants of the lakes
or river-banks, or among tufts of weed in the sea; hides
under stones and in the crevices between them, or lies
in the tunnel which it has burrowed in the mud or
loose sand, its head and tail projecting at either end.
Its caution is extremely great, and it avoids every sus-
picious object. An old tradition is current among the
fishermen that if, on its migration in a river, the Eel
comes to the unbarked trunk of a birch-tree placed
across the channel, it halts; and by laying down such
obstacles they force it to take the path leading into
Eel-weirs or similar contrivances. Linnaeus makes re-
ference to this in his “coercetur t-runco albo Betuhe.”
To changes in the weather it is very sensitive, and
becomes greatly distressed and very restless in a thun-
derstorm, quitting its retreat, and falling an easy victim
to the seine. In winter it burrows to a depth of se-
veral feet in the mud, and lies torpid, often in large
companies, to escape the cold. But it may be frozen,
though not too hard, and again thawed to life. Its tena-
city of life is known to most by experience. Though
skinned and chopped to pieces, it still moves. Its suf-
ferings, before death finally releases it, must awake the
pity of all. Ghastly is the description given by Jacoby"
of the Eel-roasting carried on by the Italians at Co-
macchio. “A large establishment for the roasting of
Eels is a sight during working-hours that none will
forget. You see before you a living picture of hell,
where the damned suffer all the torments that the pious
imagination of mediaeval painters could conceive; and
you are surprised at every moment by the perfect re-
semblance to the work of their pencils. In the back-

ground a huge door opens now and then, to let in the
full flood of daylight. Through the door and over the
murky water of the canal a rower guides his broad boat,
mouthing execrations. He brings with him the souls of
the damned, the Eels, which lie in writhing heaps at
the bottom of the boat. The victims are now scooped
up with nets and thrown into tubs. In front of each
tub sits a fiend, armed with a sharp hatchet. He chops
into three or four pieces the wriggling bodies of the
large Eels, which vainly strive to escape. The small
Eels are cast as they are, together with these pieces,
into other tubs. The work is now taken up by other
hands, with another diabolical duty to perform, that of
spitting on huge skewers, up to two yards long, the
pieces and the live Eels in coils, one after another. The
spits, loaded with the still writhing pieces and the
wriggling small Eels, are now taken to the fire. Eight
or nine large furnaces, heated with great blocks of wood,
spread a violent heat over a great portion of the dusky
room. In each furnace, before and in the fire, hang
seven or eight of these loaded spits. They are kept
turning by women, who in face, age, and figure har-
monize well with their infernal surroundings. Each
gang of workers, men and women, chants its song, the
flames roar, the smell of burning fat rises from the
victims, and the picture of hell is almost complete.”
Man is here no more merciful than the beasts, which
in the struggle for existence reck little of each other’s
sufferings. A different proof of the Eel’s endurance was
afforded us by the dissection of a porpoise that had
been found dead in the Catt-egat off the coast of Bo-
huslan on the 29th of April, 1878. In the abdominal
cavity lay a dead Eel 465 mm. long. It had evidently
been swallowed alive by the porpoise, but had retained
strength and sense enough to gnaw its way through
the wall of one of its devourer’s stomachs into the ab-
dominal cavity, thus inflicting death on the porpoise,
though itself unable to escape from its prison6. In
spite of these powers of endurance it is easy enough
to make a wriggling Eel lie still. Only cut a slit
across the hind part of the tail; and so great is the
sensitiveness of this part that the Eel becomes motion-
less, probably with pain.

The Eel feeds principally by night, the time when
it is most active in every way. Even in the daytime
it may be enticed with a tempting bait, but, as a

a Der Fisclifang in der Lagune von Comacchio nebat einer Darstellung der Aalfrage, Berlin 1880, p. 83.
b Cf. above, p. 621, Darwin’s anecdote of a Diodon.

COMMON EEL.

1029

rule, only when the water is turbid after a storm or
in wet weather. It devours almost all animal food,
live or dead, fresh or putrefied, that it can swallow.
The small Eels content themselves with lower marine
animals of every kind, small crustaceans (Gammarids),
worms, and mollusks. Their elders begin with small
fishes, such as Sticklebacks, Sand-eels, and Lampreys,
and often gorge themselves with fish-roe. In fresh
water they wage a ruthless war of extermination against
the crayfish, which they most appreciate just after the
old shell has been cast, and while the new one is still
soft. They also attack one another; and instances are
recorded of the victor in these combats being suffo-
cated by its prey. The latter, the head of which was
already swallowed, had bent its tail right back and
forced it out through the gill -opening of its captor.
Soon they do not shrink from assailing higher ani-
mals, such as frogs, young waterfowl, and water-rats.
They even pursue the water-rats into their subaqueous
holes, and Eels are frequently found that have turned
these into their own hiding-places. But vegetable
substances, such as grains of corn, demonstrably enter
into their diet"; and Eeddersen relates* 6 that at Copen-
hagen it was a popular diversion^ to feed the Eels in
a pond in Orsted’s Park with bits of bread. The vo-
racity of the Eel thus renders it omnivorous.

In this connexion we may consider an observation
repeated on many occasions, but as often explained
away or even ridiculed: — the Eel, like the above-
mentioned Climbing Perches ( Labyrintliici ), voluntarily
travels in quest of food by land. “The Eel is said
sometimes by night to crawl out of the water on the
fields where it finds lentils, peas, or beans sown”, wrote
Albertos Magnus' in the 13th century. “This mig-
ration”, wrote BociG in the 18th century, “explains
the mysterious fact that in Prussia and Pomerania
fish are caught on dry land and with the plough.
On warm nights, when the Eels betake themselves to
the peas, the peasants plough a few furrows along the
water towards dawn, before the day lias broken; and
these are the nets in which the Eels are taken. For,
though the Eel can drag itself along on the grass, its

retreat is cut off by the upturned sods. The rustics
consider it a sign of approaching storm when the Eel
quits the water for dry land.” The Royal Museum
possesses in its collection of manuscripts a communi-
cation made to Prof. B. Fries in 1836, and relating
how the Dowager Countess B. M. Hamilton in the
early part of the century instituted the most cautious
observations of some Eels and caused their capture
during their land excursions on her estate. They
wandered of a night from Lake Hedenlunda into a
field, and ate pea-pods “with a smacking sound, like

that made by sucking-pigs when they are eating

On investigation it was found that the pods were not
gnawed in pieces or eaten up, but that the Eels only
consumed the outer soft and juicy skin covering the
young pods; and after this discovery the Eels, which
were kept at Hedenlunda in a live-well, were fed with
pea-pods.” The communication further contains a
description of the acuteness with which the Eel ap-
prehends the slightest noise, even on land, and the
celerity with which it then retreats to its proper ele-

Fig. 277. Intestinal worm (Ascaris labiata) of the Eel compared
with a young Eel of the same size. After Benecke.

ment. Ekstrom rejected all similar anecdotes as fa-
bles; Nilsson' assumed that the Eel’s object in these
journeys tvas the quest of a food more suitable than
peas, namely slugs ( Limax ). This opinion may be

more reasonable, but credence can no longer be re-
fused to the above-cited observations, for we have
evidence from other sources of the Eel’s taste for vege-
table food.

The propagation of the Eel was a riddle for thou-
sands of years, and has given rise to the most mar-
vellous conjectures. According to Aristotle7 no one
had found eggs in the Eel up to his time; but even
then its intestinal canal had been observed to contain
“hair-like or worm-like growths” (fig. 277), that were

a See Trybom, Dausk Fiskeritidende, 1885, p. 411.

6 Same periodica], 1885, p. 341; 1891, p. 397.
c Quoted by Siebold, 1. c-., p. 314.

d Wirtschaftl. Naturges chi elite von dem Konigreich Ost- und Westpreussen , 1784, quoted by Benecke, 1. c.. p. 175.
e Skand. Fna , Fisk., p. 669.
f Anim. Hist., lib. IV, cap. XI.

1030

SCANDINAVIAN FISHES.

supposed to lie young Eels, a mistake against which
he cautions his readers, but which often crops up, even
in modern times. He believed that Eels were born of
worms generated in the sea by decaying weed and on
the shores of fresh water by mud under the influence
of heat". According to Pliny6 the Eels rub themselves
against the rocks, and what they thus scrape off their
bodies comes to life. According to Rondelet0 even
Athenteus and Oppian (2nd and 3rd cent., A. D.) had
seen Eels knot themselves together in bunches; and
they believed that the slime the Eels thus pressed out
of one another’s bodies received animation. The old
tale of the Eelpout ( Enchelyopus , see above, p. 607)
as “Eel mother” can be traced in literature back to
Albektus Magnus. With these conjectures the Eel
question was dismissed throughout the Middle Ages;
and the founder of modern ichthyology, Artedi, re-
frained from touching the point in his writings'7. Ti-
selius, a Swedish rector and a contemporary of his,
had, however, published at Upsala in 1723 an “ Ut-
forlig beskrifning ofver den stora Svea och Gotlia Sjon
Wdtter ”, where he relates (p. 113) that “in several
Eels, close to the small of the back and the spine,
has been seen a fine and handsome roe of a reddish
appearance”. This observation probably refers to the
kidneys of the Eel, but perhaps to the true ovaries.
The first discoverer of these may- thus have been a
Swede; but in scientific literature the Italian MondinP
and the Danish naturalist 0. F. Muller7 are generally
mentioned as rivals for this honour. The ovaries were
more thoroughly investigated by RathkiT and Hohn-
baum-Hornschuch \ The male organs of the Eels
were not discovered until more recent times. Rathke
indeed speaks in several passages’ of such organs, but

whether he referred to the true testes, or had ac-
knowledge of their structure, seems more than doubt-
ful, for he so expressly states that they are without
efferent duct. SyrskP was the first to publish more
accurate information of these organs, and since then
B roc ii 7 and Ryder/ have been the principal contri-
butors to the elucidation of questions connected here-
with.

The first observation from which we can derive
a positive opinion as to the breeding of the Eel, is
old enough, even in a literary sense. Aristotle was
quite aware that the Eel goes out to sea, but he ex-
pressly denied that these wanderings were due to sexual
instinct, the Eel being destitute, in his opinion, both
of semen and ova. From the middle of the 17th cen-
tury, however, it has been known that the Eel breeds
in the sea. We are told by Franciscus Redi “Now
there are other fishes that pass the greater part of their
life in fresh water, but resort to the sea for the pur-
pose of disburdening themselves of their seed. Thus
I have arrived, by means of numerous and long con-
tinued observations, at the conclusion that year by year,
as soon as the rainy season sets in about the month
of August, especially on dark and cloudy nights, the
Eels repair in great numbers from rivers and lakes to
the sea, where they deposit their germs. The small
Eels born of these swim up the mouths of the rivers
into fresh water earlier or later, according as the
weather is more or less severe, towards the end of
January or just in the beginning of February, so that
the migration is commonly over by the end of April.
They do not arrive in one body, but in several de-
tachments and at varying intervals. They come in
such numbers that some fishermen whom I cominis-

“ There are several mediaeval receipts for the breeding of Eels by laying two moist sods with the grassy sides together.
b Hist. Mundi, Lib. IX, cap. 50.
c De Pise, fluv., p. 199.

d Linnaeus ( Syst . Nat., 1. c.) based his opinion on Fahlberg’s communication to the Swedish Academy of Science in 1750 of the
discovery of young in the intestine of Eels, a repetition of the old confusion with intestinal worms.

e De Angaillae ovariis, communicated to the Academy of Bologna in 1777, but not printed until 1783.
f Underbrochne Bemiiliungen bei den Intestinalwurmern, Schr. Berl. Ges. Naturf. Freunde, vol. I, 1780, p. 204.
g Beitr. Gesch. Thierw., 2:te Abth. (Schr. Naturf. Ges. Danzig, Heft. Ill), pp. 121, 161, 175; Weibl. Geschlechtw. Aal., Arch.
Naturg. 1838, p. 299: Bemerk. liochtr. Aal , Arch. Anat., Phys. 1850, p. 203.
h De Anguill. sem et gener., disp. G-reifsw. 1842.
i Beitr., 1. c., pp. 183, 186, 196.

I Uber die Reproductions- Or gane der Aale, Sitsber. Akad. Wiss. Wien, LXIX, i (1874), p. 315.
k Mith. Zool. Stat. Neap., Bd. 2, p. 415.

1 Bull. U. S. Fish. Comm., 1885, p. 1.

m Francisci Redi, Opusculorinn pars tertia, sive de animalculis vivis quee in corporibus animalium vivorum repermntur, observationcs.
Ex Etrusci Latinas fecit Petrus Coste, Lugd. Batav. 1729, p. 99.

COMMON EEL.

1031

sioned in 1607 to fish the River Arno at Pisa between
the bridge nearest to the sea and the middle bridge,
took within the space of live hours more than 3,000
pounds of Eels in seines alone. Another fisherman of
the same river, only half a mile (5,000 paces) from the
sea, caught just at daybreak more than 200 pounds of
Eels which were so young and small that about 1,000
of them went to the pound — I mean a Tuscan pound,
which contains 12 ounces. But not all the Eels are of
the same size when they enter fresh water ’; by far the
greater number measure, according to his appended li-
gures, about 35 — 55 mm., the remainder about 70 —
140 mm. On the same experience the great Eel-fishery
of the lagoons off Comacchio, at the mouth of the Po,
has been based since time immemorial. Here the ex-
tensive system of lagoons has been divided by means
of dams and dikes into broads and canals, where the
Eels can lead their fresh- water life. Every year, on
the 2nd of February, all the sluices are opened, free
ingress from the Adriatic being thus afforded to the
ascending Elvers, which struggle up against the current,
swollen as it is by the winter rains. By far the greater
number of them are under 6 — 8 mm. in length, accord-
ing to Jacoby. On fine days they keep to the bottom;
at night and in cloudy weather they swim nearer to
the surface. This Eel-fare , called by the Italians mon-
tata (the French montee ), lasts three months. Towards
the end of April the sluices are closed. They are again
opened at the end of September, when the summer
drought has so reduced the volume of the lagoons that
the water of the Adriatic streams in. At this season
the sexual instinct awakes in the old Eels — from 4 to
6, or, according to some, as much as 10 years of age —
and they set out on their seaward migration. But their
path is barred by the fisherman’s devices; intricate
systems of weirs guide them into large baskets, where
the Eels first congregate, to be afterwards transferred
to the broad boats which we have mentioned above.
This fishery with all its contrivances depends on a
knowledge of the migrations undertaken by the Eel,
and was already old when Torquato Tasso, in his La
Gerusalemme Liberata, compared Tancred in the am-
bush to the Eels in this labyrinth.

In Sweden too, time out of mind, the fisherman
has set Eel-nets ( lanor ) and constructed Eel-traps with

a Ann., Mag. Nat. Hist., ser. 6, vol. XII (1893), p. 35.
b Lillj., Sv., Norg. Fisk., vol. Ill, p. 392.

the entrance opposed to the stream for the admittance
of the descending Eels. In our rivers and lakes this
downward migration begins in May, but does not be-
come general until the dark nights set in, towards the
close of August. “In the neighbourhood of Upsala”,
writes Sundevall, “Baron Cederstrom was informed
that before midsummer the Eels haunt the large beds
of reeds and horsetail on the lower course of the Fyris.
During July (“till St. Olaf’s Day”, the 29th), when
these reed-beds are cut, they descend into the arm of
Lake Millar known as Ekoln, into which the Fyris falls.
At the beginning of August great quantities of Eel are
taken there on long-lines with 400 hooks. The Eels
proceed down this bay to Sko Land, the south coast
thereof, where the depth is greatest, measuring 20 — 30
fathoms. Here they are taken for a. short time in the
middle of August, after which they disappear”. During
its sojourn in fresh water the Eel retains its keen appe-
tite; but less and less food is consumed as the sexual
instinct asserts itself, and as the true migratory dress,
the pale belly, is adopted. Frequently the migrating
Eels knot themselves together in bunches, and large
bundles, often a fathom in circumference, are seen lying
still in the lakes or trundling down the streams. On
reaching the sea the Eel proceeds on its way to the
spawning, grounds. Their situation has not yet been
ascertained; but Calderwood" mentions a female 2972
in. (749 mm.) long and “almost ready to spawn” that
was taken on the 27th of December, 1892, about 12
miles south of the Eddystone, or 20 miles from the
nearest point of land, Raine Head. That the Eel of
Swedish waters spawns partly in the Baltic, seems pro-
bable, young Eels 7 or 8 cm. long having been found
both in the Roslag (Upland) and in the Ljusne Elf'.
But the main body of the Swedish migratory Eels roves
along the Baltic coast and round Scania out through
the Sound, perhaps too, in company with Russian and
German Eels, through the Belts. That the Eel spawns
in the Cattegat, is proved by the multitude of Eel-fry
1/2 — 1 dm. long so often to be observed in spring and
summer all along the west coast of Sweden making
their way into fresh water. In the north of the Sound,
nearer to the Swedish than to the Danish coast, is a
deep channel with more than 10, in places 20 fathoms
of water, where Eel-fry occur in myriads during win-

Scandinavian Fishes.

130

1032

SCANDINAVIAN FISHES.

ter. “And here”, says Villumsen®, “it may often he
observed how the Cod in winter disgorge from their
crammed stomachs extremely small and semi-transpa-
rent young Eels, only 5 — 8 cm. long. Food digests
rapidly in the stomach of a Cod, and as the disgorged
Eels are fresh and entire, we may safely conclude that
they have been swallowed quite recently and thus on
the spot. I have also seen them jump alive from the
jaws of the Cod, which had probably seized them at
the same moment as it took the fisherman’s bait. The
bottom in this part of the channel consists of mud
mixed with clay, and must be tenanted by multitudes
of Eel-fry, for sometimes every single Cod has secured
a number of them. It cannot well be assumed that
these frail creatures swim freely about at a time of
year when the larger Eels eagerly take refuge in the
security of the bottom. It is more probable that they
lie concealed in the mud, but that the Cod knows how
to rout them out from their hiding-places. This opi-
nion is supported by the fact that the stomach of the
Cod also contains other inhabitants of the mud, such as
Ascidians, Holothurians, worms, etc.”

“The Eel lays its eggs in the mud”, writes Des-
marest6, “after a kind of copulation. The eggs are bound
together by a slimy mass like that which envelops the
ova of the Perch, and form small clues or round balls.
Each female, as we have ascertained by personal ob-
servation, annually lays several of these agglomerations.
The fry are soon hatched, but remain for some days
after exclusion within the said balls. When the fry
have attained a length of 4 — 5 cm.', they liberate them-
selves from the bonds that confine them, and ascend
the neighbouring rivers in dense and extremely nume-
rous bodies”.

Not all the Eel-fry, however, repair to the rivers
at this age. A great number pass one or two years
in the sea'7, and are 2 — 4 dm. long when they under-
take the ascent. At Elfkarleby fry of this size make
their way up the Dal Elf, where they are called Al-

vimrnor or Alvinner e, without intermission from July
till October, but mostly in September. These Alvim-
mor or Elvers are the Civelles of French rivers. When
they have grown somewhat larger, and are found among
the seaweed of the littoral regions or the grass of the
lakes, they are known in Sweden as Gras-til or, on ac-
count of their watery flesh, Blot-al, in Denmark as
Visse-til.

We have still to consider the question whether the
Eel also breeds in fresh water. As we have seen, this
is hardly probable. Eels occur, it is true, in tarns of
great elevation and in isolated pieces of water which
seem to be cut off from the sea by barriers impassable
to a fish. But Eel-fry have an almost incredible capa-
city of penetrating or circumventing obstacles in their
path, and their elders can also travel, by land. Obser-
vations have been made, however, which are at least
easiest of explanation on the assumption that the Eel
propagates in fresh water as well as in salt/. Trybom
states (1. c.) that “in 1864 1,000 young Eels were trans-
planted from Elfkarleby to Lake Hagel in Dalecarlia.
The lake has no outlet, and no Eels had previously
been found in its waters. In 1871 Eels weighing lx/4
— I1/ 2 kilo, were taken there, in 1872 specimens that
turned the scale at 1*7, 2, and even 3*4 kilo. In the
summer of 1879 small Eels weighing only V5 kilo, as
well as larger ones were caught. Eel-fry have been
planted in the lake on only one occasion”. Only a few
years ago it was supposed that male Eels occur in the
sea alone, and do not ascend, at least not in any great
number, into fresh water. It was consequently incon-
ceivable that the Eel should multiply in fresh water.
But the collections of the Royal Museum had contained
ever since 1844 a male Eel 23 cm. long, taken by
Lieutenant Robsahm at Trollh&ttan ; and in 1880 Her-
mes9 found 13 male Eels in the Elbe off Cumlosen, near
Wittenberg, about 25 German miles from the mouth of
the river. Recently (the middle of June, 1893) at
Silkesborg Papermills and Holm Mill, in the Eel-traps

a Dansk. Fiskeritidende, 1892, p. 15.
b Chenu, Encycl. d'Hist. Nat., Rept., Poiss., p. 328.

0 But of. Jacoby’s statement, mentioned above, that most of the young Eels on their upward journey at Comacchio are only 6 — 8
mm. long.

d In the old outlet of the River Nissa, the so-called Svinback, open towards the sea, but now almost choked up, Eel-fry Y2 — 1 dm.
long used to take up their summer quarters without evincing any migratory tendencies; and the boys of Halmstad amused themselves by
wading in the shallow water to frighten these young Eels out of the mud and clay and catch them.

e Sundevall, Stockh. L. Hush.-Sallsk. Handl. 1855, p. 92; Trybom, Landtbr. Akad. Handh, Tidskr. 1881, Om s. k. Alvinner etc.
f See for example Lewin’s remarks in Wittmack, Deutsch. Fisch. Ver. Circular, No. 1 (1875), p. 127.

3 Bull. U. S. Fish. Comm. 1881, p. 98, and Mitth. Sect. Kiist., Hochseef., Deutsch. Fisch. Ver., 1893, p. 113, not. 2.

COMMON EEL.

1033

of the Guden, which falls into Randers Fjord (the Cat-
tegat), Feddersen® found among descending Fels 2 1/2
— 4 dm. long numbers of males with yellow belly,
amounting at the first-mentioned place to 80 per cent
of the catch. In Lakes Hjul, Bras, and Avn, within
the same river-basin, similar males were taken with
Eel-seines. A month later (15th — 17th July) the males
in and near the Lakes of Silkesborg had begun to de-
velop the white belly of the migratory dress, but judging
by the take were fewer. He made similar observations
at Maarup Mill, between Lakes Orum and Ove, on the
other side of Liim Fjord, and in Westen-see (Province
of Rendsburg, Holstein). It thus seemed probable at
all these places that a descent of young Eels, especially
males, takes place in summer, before the general descent
of older females, commences. The males, like the fe-
males, consequently spend a portion of their life in fresh
water ; but the assumption that the Eel spawns in fresh
water, side by side with its normal breeding-season in
the sea, first requires authoritative confirmation.

Not all the Eel-fry, as we have seen, ascend into
fresh water during the first year of their existence; nor
do all females of a size that indicates maturity under-
take migrations. In the lagoons of Comacchio the fisher-
men have long been aware of this, and call the large
Eels that do not migrate Pasciutib. Near the same
locality these forms are also found in the sea, off the
coast; and Jacoby described them as broad-snouted,
small-eyed, yellow-bellied, high-finned, and with glassy
ovaries, destitute of fat and containing transparent eggs
with little, if any, granulation. He explains them too
as sterile females; but whether this sterility is perma-
nent or temporary, still remains an open question, for
many broad-nosed Eels have ovaries of normal develop-
ment. In Sweden they are known on account of their
voracity as sluk-al (ravenous Eel), the Danish klcepal.
They do not rove in shoals, but lead a more solitary
life; yet are often caught, for they freely take a hook,
the best bait being Roach. Jacoby extols their de-
licious flesh, “which melts in the mouth”. Kroyer

/

describes their flesh as hard, dry, and lean, and
their skin as hard and tough. The difference may
easily be due to the recency or remoteness of the
spawning-season .

In recent times ichthyologists have been most in-
clined to the belief that after spawning the Eel diesb
This opinion may be correct, for it is partly true of
the Salmonoids, and applies to the Lampreys, from
which the Eels do not differ widely in the structure
of the generative organs. On one or two occasions
dead Eels, spent, as it is stated, have been found on
the seashore, and a skipper is said to have sailed some
distance in the North Sea among dead Eels, floating at
the surface. Large Eels have never been found mi-
grating in any number to fresh water. But this is no
adequate solution of the point, for that a re-ascent may
take place, though it is seldom observed, appears from
Robin’s d discovery of female Eels, with the stomach
full of marine animals — Annelids ( Eunice sanguined)
and shellfish (Doris) — which they had brought far up
the Adour, 40 kilom. from the sea. The so-called blot-
alar and slukdlar in the sea must also be taken into
consideration. They are demonstrably not always ste-
rile, nor has it yet been proved that the larger among
them have never spawned.

The Eel is greedily preyed on by mammals, such
as otters, seals, and dolphins, by piscivorous birds, and
by larger fishes; but its most destructive enemy is
undoubtedly man. To a good digestion its flesh is one
of the most excellent foods, whether boiled or fried
— less indigestible in the latter form — fresh or salted,
smoked or marinaded. The young are made in many
places into Eel-cake (cf. above, p. 973). The tough
skin of the Eel has been most generally employed as
material for flail-thongs.

The great Eel-fisheries have contributed in a high
degree to our knowledge of the Eel’s habits, for the
method of setting his tackle which the fisherman has
learnt from the experience of ages best indicates how
the Eel conies and goes. When the Eel descends the
rivers, large or small lanor, a kind of seine with fine
meshes in the bosom, are stretched across the channel,
or Eel-traps ( allms ) are constructed in the slopes of the
streams. These are timbered like log-houses, but on
the lower side chinks are left between the logs, wide
enough to allow of the passage of the water, but too
narrow for the Eel to slip through. On the side facing
the current a hole is made, in which is inserted a

“ Dansk Fiskeritidende 1893, p. 333.

6 “Pasturers”.

c See. Jacoby, ]. c., p. 55.

d Comptes Rendus, Fevr. 21, 1881, p. 382.

1034

SCANDINAVIAN FISHES

trough or wooden pump-barrel, blackened lest it should
frighten the Eel, and with the inner end raised high
enough above the level of the water in and below the
trap to cut off the Eel’s retreat. When the trough is
closed, the water runs out of the trap, and the Eels
are fished up. On the same principle alkistor are con-

structed beside mills and other waterworks; but their
lower end is generally closed with a fine iron grating.
Into Eel-trunks (bradsumpar), which are placed in a
strong current, the Eel is guided by two converging
rows of freshly peeled, white stakes or unbarked birch-
trunks. Wherever the Eel is known to have its haunts

in fresh water, among the reeds and in calm inlets,
Eel-boxes (al-lador) are sunk, lined with woolly sheep-
skin, the wool facing inwards, and pierced with round
holes 3 or 4 cm. wide. At these holes the sheepskin
is slit up crosswise, and the strips are left hanging

Fig. 279. Eel-spear.

there. The Eel-box is baited with pieces of meat or
the like wrapped in fresh pea-haulm, and the box is
then left in a fathom or two of water for a week or
so, when it is examined, and moved to another locality,
if the catch is too poor. In the same way Eel-baskets

Fig. 280. Danish Eel-vveir. After Drechsel.

COMMON EEL.

1 035

( dlkupor , fig. 278) are set, constructed on the same
principle as mjcirdar (see above, p. 32), but triangular,
woven with osiers or thin deal splinters, and with a
round wooden plug to stop the opening at the narrow
end. With the setting of Eel-lines all our readers are,
no doubt, familiar. They may be baited with fish, if
possible alive, shrimps, or worms. Eels may also be
taken with an ordinary hand-line, baited with worms
or fish-offal — pyloric appendages are best — but the
bait must lie on the bottom. With the Eel-spear (Iju-

in the bottom ( harkning , hutt'ning, or puttning), doing
more harm than the catch can repay. In Germany and
Denmark much use is made of the so-called al-vad
(Eel-seine), either the drif-vad (drift-seine, Germ. Zee-
sen), a net for bottom-fishing, plied from a sailing-boat,
and with a double bag as in the trawl, or the handvad
( pidsvad , bott-vad, ankar^vad, or snurre-vad ), manipulated
from a rowboat, and with a simple bag. Large or small
seines ( strandvadar ) are also drawn from the shore. Or
the fishermen wade out with an alglip, a square scoop-

Fig. 281. Different methods of setting alhomtHor, A, from the Province of Calmar; B. from Blekinge and Eastern Scania. After Lundberg.

ster, fig. 279) much Eel is caught, both in winter and
summer. Where the Eel lies hidden in the mud, or
among the grass or weed, air-bubbles rise to the sur-
face. In winter these bubbles stop under the ice, and
show the fisherman where to strike; in summer he
watches for them in smooth inlets, or where the current
is not strong, and there the Eel lies of a morning, with
its head turned towards the sun. The fisherman thus
knows where to plunge his spear so as to transfix the
Eel, even without seeing it. In many places, as in the
island-belt of Blekinge", the Eel-spearer strikes blindly

net, which one of them holds on the bottom, while the
other splashes in front of him, to drive the Eels into
the net. But the most valuable Eel-fisheries depend on
the migrations of the Eel in the sea. In Denmark Eel-
weirs ( algardar ) have been constructed from prehistoric
times. These are rows of stakes, or fences woven with
brush (fig. 280), running straight out from the beach,
and with an al-ryssja (liomma) at the outer end. Or
there, as on the Swedish coast, only biommor are used,
set singly or in a row, one outside the other (fig. 281).
These constructions have conferred names on the mi-

See Forslag till ny fslceristadga, Stockh. 1883, p. 94.

1036

SCANDINAVIAN FISHES.

grating- Eels, gard-al in Denmark, hom-al in Sweden.
The hommora are one-armed or two-armed, furnished
in the latter case with a long arm and a short one, the
former (the leader or lang-arm ) to guide the Eel into
the homma itself ( halt ), the latter (the check-arm or
gin-arm ) to head off the fish and prevent them from
passing the opening. At the outer end of the liomma
an Eel-basket ( kass'e or tina) is set, out of which the
Eel cannot creep back. The whole engine is firmly
anchored to the bottom with large stones, and the bas-
ket is secured in a sled-shaped wooden frame. When
the Eel, on its migration along the east coast of Swe-
den, comes from the north and north-east, along the
south coast from the east, and in the Sound from the
south, the hommor must be placed with the opening on
that side of the leader from which the Eel approaches.
The same rule applies, of course, to the position of the
al-ryssja at the outer end of the Eel-weirs, where the
weir may be lengthened by setting more ryssjor outside.
The season lasts through the closing months of the

year, dark and stormy nights and land-winds being
most favourable to the fishery. Most of the Eels taken
on the east coast of Sweden are sold alive to Germans
who sail in their well-boats ( qvasar ) along the coast,
buying up the supply.

The value of the Eel-fisheries in Scania and Ble-
kinge, according to Lundbeeg6, was in the years

1882 £8,240,

1883 £11,604,

1884 £15,691,

1885 £14,846,

the price fetched in Scania being about Is. Id., in Ble-
kinge about 10V2d. per kilo. The statistics of the
Eel-fisheries in other parts of Sweden are extremely
defective; but we probably do not exaggerate in assu-
ming that the Eel affords our country an average an-
nual income of between £16,500 and £’22,000. In
Denmark the Eel-fishery is still more productive, and
its combined annual value in the two countries certainly
exceeds £55,000.

Genus CONGER.

No scales in the skin. One row of jaw-teeth larger than the rest and transversely compressed at the base, but more or
less sharpened at the tip in the longitudinal direction of the jaws. Vomerine teeth set in a card containing several rows.

Fig. 282. Bones of the head in a Conger ( Conger niger ).

alsp, alisphenoid; art, articular part of the lower jaw; bsp, basispheuoid ; cell , ceratohyoid; ceth, cartilaginous parts of the ethmoids; d, dental
part of the lower jaw; et, ethmoid; fr , frontal; Inn, hyomandibular ; top, interoperculum; mp, maxillary (maxillo-palatine) ; op, operculum;
pob, preorbital (first suborbital); pop, preoperculum; psp, parasphenoid ; pt, ento pterygoid; qu, quadrate; R. br. I — IX, first — ninth branchio-
stegal rays; sob, hindmost suborbital bone; sop , suboperculum; spet, supraethmoidal (nasal); spho, postfrontal (sphenotic) ;

squ, squamosal; uh, urohyoid.

a A minute description of this fishery may be found in Lundberg, Orn dlfisket med s. k. hommor vid svenska Ostersjokusten saint
Oresund, Landtbr. Akad. Handl. och Tidskr. 1881.

6 Meddelanden rorande Sveriges Fiskerier, Part. II (1888).

CONGER.

1037

The genus of the Congers, which as defined above,
after Ivaup“, contains a few (3 or 4) ascertained species,
in the essential form of the body and fins resembles
the preceding genus, only that the dorsal fin commences
further forward. The character afforded by the form
and distribution of the teeth (fig. 282) distinguishes it
not only from the preceding genus, but also from three
other genera* belonging to the family of the Eels. The
most important skeletal differences from Anguilla are
remarked above. Here we shall merely add that the
nasal (supraethmoidal, spet) bones covering each nasal
cavity are more developed in Conger, being indeed thin,
but broad, and broken up anteriorly into a grating,
with the openings set transversely or, at the extreme
front, directed forwards.

The generic name had a specific signification in

Linnasus, and ought indeed to have retained this capa-
city; but Cuvier" raised it to its present rank, which
it has uniformly preserved in the system. Now that
it has been proved (see above) that the formerly recog-
nised genus Leptocephalus, which has occupied a place
in the system since 1763, consists of larvie and dege-
nerate forms, belonging in great part to the species of
the common Conger, it may well be asked whether the
Cuvierian generic name ought not to be replaced by
the Gronovian. But at present, so long as there are
Leptocephalus forms which cannot be referred with
certainty to any definite species, the alteration might
easily cause confusion"'. Furthermore, Conger has been
known as a generic name ever since the time of
AristotliC, and in the works of Belon7, Rondelet*,
and WillughbyC

THE CONGER (sw. hafsalen).
CONGER NIGER.

Plate XLV, fig. 2.

Distance betiveen the dorsal fin and the tip of the snout about /5* 1 of the length of the body, or at most about
'if °f that betiveen the anal fin and the same point. Length of the snout 1/i or more of that of the head,
which measures 2/3 — 3/4 of the distance betiveen the dorsal fin and the tip of the snout. Length of the pectoral

fins about 1/. — V, of the last-mentioned distance.

R. br. 9*— 10; I). ca 275—300; A. ca 205—225; C. 10;
P. 17 Vert. 153— 156 m.

Syn. Murcena supremo margine pinnas dorsalis nigro, Art., Ichihyol.,
Gen., p. 24; Syn., p. 40.

Murcena Conger, Lin., Syst. Nat., ed. X, tom. I, p. 245;
Bl., Naturg. Ausl. Fisch., pt. II, p. 37, tab. CLV; Ahl
( Ophichthus ), Mur., Opli. (disp. Ups. 1789), p. 11; Mitch.
(Anguilla), Lit., Phil. Trans. N. York, vol. I, p. 360;
Pall. (Murcena), Zool. Ross. Asiat., tom. III. p. 72; Nilss.,
Prodr. Iclithyol. Scand., p. 64; Kr. (Anguilla), Danm.

Fisk., vol. Ill, p. 603; Nilss. (Murcena), Skand. Fna,
Fisk., p. 680; Bit. -Goode ( Leptocephalus ), Fisher ., Fisher.
Inclustr. U. S., sect. I, p. 656, tab. 240.

Murcena myrus, Brunn., Ichthyol. Massil., p. 12 (false de-
term.).

Eclielus Macropterus, Raf., Caratt. Ale. N. Gen., p. 64
( + E. Gr uncus , p. 65), tab. XVII, figg. 2 et 3.

Murcena nigra, Kisso, Ichthyol. Nice, p. 93 (+ Mur. conger,
p. 92); Jord., Gilb. (Conger niger), Bull. U. S. Nat. Mus..
No. 16, p. 362; Coll., N. Mag. Naturv., Bd. 29 (1884),
p. 113.

“ Cat. Apod. Fish., Brit. Mus., p. 111. The characters were first pointed out, however, by Richardson in Ichthyol. Ereb., Terror , p. 107.
6 Ophisoma, Swainson ( Congromurcena , Kaup, Gthr), Uroconger , Kp, and Poeeiloconger, Gtiir.
c R'egn. Anim., ed. 1, tome II, p. 231.

^ Jord., Gill., Bull. U. S. Nat. Mus., No. 16, p. 362, note.

e rdyyQOg, Be Anim. Hist., lib. I, cap. V; lib. II, capp. XIII, XV, XVII; lib. Ill, cap. X; lib. VI, cap. XVII; lib. VIII, capp. XIII,
XV ; lib. IX, cap. II.

J Nat., Divers. Poiss., p. 159.
g De Pise., lib. XIV, cap. I, p. 394.
h Hist. Pise., lib. 4, cap. V, p. 111.

1 19 — 20 %; in old specimens, according to Kroyer, up to 23 %.
j 46—52 %.

k Sometimes 8, according to Bleeker.

1 „ 15, ,, ,, ,, , and 19, according to Day.

m 154- — 164, according to Day.

1038

SCANDINAVIAN FISHES.

Conger vulgaris ( Congre commun ), Cuv., R'egn. Aniin ., ed. I,
tom. II, p. 231; Yaee., IJist. Brit. Fish., ed. I, vol. II,
p. 304; Schleg. in Sieb., Fna Japon., Pise., p. 259;
Blkr, Atl. Ichth. Ind. Or., tom. IV, p. 26, tab. CXLIX
{Mur. V), fig. 2; Gthr, Cat. Brit. Mus., Fish., vol. VIII,
p. 38; Coll., Forh. Vid. Selsk. Chrnia 1874, Tillsegsh.,
p. 199; 1879, No. 1, p. 99; Daeeste, Arch. Zool. Exper.,
tom. IV, p. 227; Winth., Naturh. Tidskr. Kbhvn, ser. 3,
vol. XII, p. 51; Moe., Hist. Nat. Poiss. Fr., tom. Ill, p.
565 ( + var. niger, p. 566); Mela, Vert. Fenn., p. 359, tab.
X; M5b., Hcke, Fisch. Osts., p. 148; Day, Fish. Gt. Brit.,
Irel., vol. II, p. 250, tab. CXLII, fig. 2; Storm, Vid. Sels.
Skr., Trondhj., 1883, p. 41; Lillj., Sv., Norg. Fna, Fisk.,
vol. Ill, p. 410; Cab., Prodr. Fnce Medit., vol. II, p. 541.

Congrus leucopheeus , Richards., Ichth. Voy. Ereb., Terr., p.
108.

Conger occidentalis, Dek., N. York Fna, pt. IV, Fish., p.
314, tab. LIII, fig. 172.

Conger Linnei, Malm, Gbgs, Boh. Fna , p. 591.

Form® larvales et degeneratse:

Leptocephalus, Gronov., Zoophyl., fasc. I (1763), p. 135, No.
410, tab. XIII, fig. 3; Morris, Penn., Brit. Zool., ed. 1776,
vol. Ill, p. 139, tab. XXV, No. 67.

Leptocephalus Morrisii, Gm., Syst. Nat. Lin., ed. XIII, tom. I,
p. 1150; Yarr., Brit. Fish., ed. 2, vol. II, p. 311; Kp,
Cat. Apod. Fish., Brit. Mus., p. 147; Couch, Hist. Fish.
Brit. Isl., vol. IV, p. 348, tab. CCXXXVIII, fig. 2; Gthr,
Cat. Brit. Mus., Fish., vol. VIII, p. 139; Moe., Hist. Nat.
Poiss. Fr., tom. Ill, p. 567.

Ilelmictis punctatus, Raf., Ind. Ittiol. Sic., p. 62, tab. II,
fig. 3; Kp. {Leptocephalus), 1. c., p. 148, tab. XVII, fig.
8 — sec. Moe.

Leptocephalus Spallanzani, Risso, Fur. Mer., tom. Ill, p.
205; Kp., 1. c., tab. XVII, fig. 7 — sec. Gthr.

Leptocephalus Gussonii, Cocco, Isis 1831, p. 1340 — sec.
Kp et Gthr.

Leptocephalus candidissimus, Costa, Fna Reg. Nap., Peso.,
Malacott., Apod., tab. XX — sec. Kp., (?) sec. Gthr.

Leptocephalus inaequalis, Facciola, Atti. Soc. Tosc. Sc. Nat.,
vol. VI, fasc. 1; Natural. Sicil., Anno XII (1893), p. 194.

Obs. Dareste proposed (1. c.) to unite with this species, or
at least to regard as a variety thereof, the East Indian and East
African Conger marginatus, Val., Gthr = Conger Noordzieki , Blkr,
and judging from the variability of the preceding species, there would
seem to be good reasons for this opinion. But apart from the more
elongated (shallower) form of the body, the higher dorsal .fin, and
the black-spotted pectoral fins, characters of Conger marginatus that
Bleeker in particular laid stress upon, we find a remarkable difference
in the shorter trunk of the last-mentioned species — the beginning of
the anal fin being separated from the tip of the snout by a distance
of about 36 % of the length of the body — and in the commencement
of the dorsal fin being still more advanced — the length of the head
measuring about 79 or 80 % of the distance between the dorsal fin
and the tip of the snout. In these respects the species is conse-
quently still farther removed from the preceding genus than Coyiger niger.

The Conger, in Sweden known as the Hafsdl (Sea-
Eel) or sometimes, in Bohusl&n, as the Konger-al, so
closely resembles the Common Eel in the form of the
body that no long description is necessary. The most
prominent differences from the preceding species are

the greater extension in a forward direction of the
dorsal tin, the more pointed tail, the longer snout, and
the larger, more oval eyes. As a rule too the Conger
is distinguished by its far more considerable size. In
Scandinavian waters it is, we may almost say, rare to
find Congers so small that they might be mistaken for
Common Eels, though Krgyer mentions a specimen
34 cm. long. The maximum length which can lie
assigned with certainty to the Conger is about 2l/2 m-
Day mentions a specimen of this length and weighing
58 kilo. The largest specimen Collett had heard of
from the Norwegian coast was taken at Farsund in
September, 1883, measured nearly 23 dm., and weighed
25 kilo. The greatest depth of the body in small
Congers is about 5 %, in large ones about 10 or 1 1 %,
of the length of the body, except in the case of fe-
males with belly monstrously distended by the tumid
ovaries.

The length of the head commonly varies between
about 13VS and 15 V2 % (sometimes as much as 17 %)
of that of the body. Its form is the same as in the
preceding species, with the above-mentioned exception
that the snout is longer, and the tip of the snout
usually projects beyond that of the lower jaw, though
this is by no means constant, both jaws being some-
times of equal length. The eyes are oval, some-
times so distinctly that the vertical diameter is only
s/4 or even 2/3 of the horizontal, sometimes, in old
specimens, less perceptibly. During the growth of the
Conger from a length of 372 to one of 16 dm. the
longitudinal diameter of the eyes varies between 1 9
and 11 % of the length of the head. The length of
the snout varies between 25 (2572> according to Krd-
yer) and 28 %, and the postorbital length is about 2/3
(66 — 64 %), of the length of the head. The interorbital
width increases in these Congers from about 15 to 24 %
of the length of the head, the longitudinal diameter of
the eyes, which in young Congers is greater than this
width, being in the old only about half thereof, and
their vertical diameter decreasing during these altera-
tions of growth from about 4/5 to 2/5 or even approach-
ing 1/3 of the same. The posterior nostril on each side
is a longitudinal slit in front of the anterior upper
corner of the eye, in young specimens measuring only
about 7i2 °f the longitudinal diameter of the eyes, in
old up to about 1/i of the same and separated from
the eye by a distance about equal to its own length.
The anterior nostrils are set here too one on each side

CONGER.

1039

of the tip of the snout, and are tubular, but compara-
tively shorter than in the preceding species. The
mouth is similar to that of the common Eel, but the
gape is comparatively larger, and the lips are still
broader, especially in front, on the sides behind the
tips of the snout and lower jaw, where the outer folds
may be expanded almost like wings. Into the base of
the outer fold on the upper jaw projects the lower
posterior corner of the preorbital bone, which is fur-

anterior part of the upper jaw and on the tip of the
lower, subulate teeth are set in a card containing se-
veral rows, the two first-mentioned groups being often,
not always, separated by a toothless space at the above-
mentioned constriction. On the jaws these pointed
teeth are continued backwards in a row double in front,
single behind, farthest back on the inside of the above-
mentioned great median row of compressed teeth, for
a short distance on its outside. The tongue is free,

Fig. 283. Branchial arches of a Conger ( Conger niger ) seeri from above. Natural size. Halves of the right arches in a natural position,
those of the left arches bent outwards, so that the under surface of their upper parts is visible.
glh, glossohyoid (true lingual) bone; cop I — IV, first — fourth copulas; libr I — IV, first — -fourth hypobrauchials ; cbr I — IV, first — fourth cerato-
branchials; epbr I — IV, first — fourth epibranchials ; pbbr I and II, first and second pharyngobranchials ; stbr, stylobranchials ;

phs, upper, phi, lower pharyngeals.

nished on the outer side with a large, round hollow,
including a muciferous cavity belonging to the cephalic
system of the lateral line. The length of the upper
jatv from the tip of the snout is about 37 or 38 %,
the length of the lower jaw about 48 — 52 %, of the
length of the head. In front, both on the under sur-
face of the rostro-ethmoidal tip (of the interm axillaries,
according to Peters’ theory) and on the vomer, in the

narrowing abruptly to a point. The branchial arches
(fig. 283) are here too without gill-rakers, and the cardi-
form pharyngeal teeth resemble those of the common
Eel. The structure of the opercular apparatus we have
mentioned above. To the gill-openings essentially the
same remarks apply as in the case of the common Eel.

The dorsal and anal fins differ from those of the
common Eel in having their longest rays somewhat

Scandinavian Fishes.

131

1040

SCANDINAVIAN FISHES.

further forward, the gradual backward shortening of
the rays being thus more extended, whence the more
pointed form of the tail. The length of the head is
always perceptibly less than the distance from the be-
ginning of the dorsal tin to that of the anal, usually
only 60 — 70 % thereof; but in Congers with an exces-
sively long head this percentage may rise to 85.

The pectoral tins are oval or obliquely pointed
above, the upper rays being longest, and extending to
about a line with the beginning of the dorsal tin, some-
what beyond or somewhat short of the same. Their
length is about 6 % — in Krdyer’s youngest specimen
only 4 % — of that of the body, or about 10 — 15 %
of the distance between the anal tin and the tip of
the snout.

Iu the scaleless skin the depressed, straight lateral
line with its light (white or yellowish white) pores is
distinct throughout the length of the body.

The coloration of the dorsal side and the tip of
the tail is gray, with a more or less pronounced tinge
of chocolate-brown, lighter or darker, sometimes even
black, the ventral side is white, >14111 a dash of violet
behind. The base of the dorsal tin is of the same
colour as the back, but towards the top the tin becomes
ash-gray or violet, to a greater and greater extent be-
hind, and the extreme margin is black. This black
tint also appears on the posterior part of the margin
of the anal tin, the rest of this tin being grayish blue
or violet, growing paler in front. The pectoral tins
are black at the upper margin, ash-gray or violet with
a white or yelloAvish white rim on the outside, dark,
sometimes .quite black, on the inner (posterior) surface.
The top of the head in colour resembles the back; the
lips and its under surface are of the same hue as the
belly, the former, however, with a more or less distinct
tinge of tiesh colour; the cheeks have a more or less
powerful metallic lustre. The iris is sometimes silvery,
punctated with black, sometimes of a metallic lustre.
In old Congers the inside of the mouth and the bran-
chial cavities are principally black; in the young the
mouth and tongue at least are white.

The Conger, as the Swedish name conveys, is
distinctly a salt-water tish, and has a wide geographical
range, extending almost round the globe. It is known
from the east coasts both of North and South America,
from St. Helena and the west coast of Europe, the
Mediterranean, India, Japan, New South Wales, and
Tasmania. On the west coast of America it has never

been found. Its true home hardly extends into Scan-
dinavian waters, though it can by no means.be regarded
as rare on the west coast of Sweden, and sometimes
strays even into the Baltic. Storm received specimens
from the outer parts of Trondhjem Fjord; but farther
north it has never been taken. On the shelving west
coast of Jutland it seems to be rarer than in Norway;
but several specimens are known from the Skager Rack,
the Cattegat, and the Sound. Esmark received a Lepio-
cepJialus caught, it was stated, in Christiania Fjord,
and perhaps a proof that the Conger may spaAvn not
far from Scandinavia, though these larval forms rove
considerable distances in the ocean. In 1877 Malm
estimated that at most between 20 and 30 specimens
had been secured during the preceding 30 years on
the SAvedish coast. Nilsson received a specimen nearly
15 dm. long from Halmstad, and another (in the Avin-
ter of 1853), 24 dm. long, from Landskrona. The
latter, according to Lilljeborg, had been left by the
waves on a sandbank; and a female 16 dm. long Avas
found under similar circumstances off Raa (near Hel-
singborg) at the beginning of March, 1883, and for-
Avarded by Dr. Trybom to the Royal Museum. Mobius
and Heincke mention two specimens from Eckernforde
Bay, the one 2 m. long and 34v/2 kilo, in weight, the
other weighing 45 kilo., one from Travemunde Bay,
measuring 17 dm. and Aveighing 1 4 3/4 kilo., and one
from Kiel Harbour, 16 dm. in length and over 15 kilo,
in Aveight. According to Mela the Conger has even
penetrated into the Gulf of Finland, to the Nyland
coast. It is plentiful on the English and French coasts
and, above all, in the Mediterranean. In the Black
Sea it is rare off the Crimea, but is often seen in
Constantinople.

The true haunts of the Conger lie in 20 — 50 fa-
thoms of Avater, but it ascends between the tide-marks,
and has been found at Ioay water on dry land. It
shares the preference of the common Eel for a noc-
turnal life, and in the daytime sIioavs the same procli-
vity for hiding among Aveeds and stones, in crevices
and in the sand. Off the mouths of rivers it lies in
Avait for the migratory fishes on their upAvard and
dowmvard journeys. It has great poAvers of endurance
and a robust appetite. Its dentition shows that it is
one of the most formidable predatory fishes. With the
pointed, though smaller, front teeth it seizes its prey,
and Avith the sharp edges formed by the close-set,
larger jaAv-teeth it mangles the victim. Its strength

CONGER.

1041

may be gathered from the fact that it has been seen
to tear asunder lines as thick as a man’s finger. It
does great damage to fishing-tackle, and lays the fisher-
man’s take under heavy contribution. It bites holes
in the nets and drags out the fish. It creeps into
lobster-pots, makes short work of the catch, and if the
material of the pot is flexible, bores its way out, tail
first, through one of the chinks. Like the common Eel,
it is sensitive to changes in the weather. Immediately
before a gale it is restless, and many Congers are cast
ashore in stormy weather. It seems to suffer especially
from cold. In winter it is often found half-dead at
the surface, with air-bladder strongly distended. The
cold apparently deprives it of command over the disten-
sion of this organ.

The rapacity of the Conger often embroils it with
other pirates of the deep, and it falls a victim to por-
poises, dolphins, Sharks, and large Rays. In the Medi-
terranean one of its bitterest enemies is its own relative,
the large Murmna, and the combats of these antagonists
were famed even among the ancients. Against the
Mursena its ordinary stratagems, such as suddenly tak-
ing refuge in holes, or firmly coiling its tail round
stones or in crevices, are of no avail; and as a proof
of its tenacity of life the old story relates that the
Murama bites off its tail, but that it survives the
wound.

The diet of the Conger comprises all kinds of fish
— principally belonging to the Clupeoid and Gadoid
families, and especially Rocklings — lobsters, crabs, and
cuttles; and it shows no mercy to its own species.
The Conger is consequently taken in numbers on the
hook, Avith a bait of Herring or other fish; but the
bait should, if possible, be fresh. Yet it does not
disdain decomposing flesh, and it has been found con-
cealed within the dead bodies of large animals.

The spawning-season occurs in Avinter. Buckland
estimated® the number of eggs in a female at over
fifteen million, perhaps a liberal computation ; but Day
mentions a femaie, Aveighing nearly 7 kilo., that had
died in the Southport Aquarium, and the ovaries of
Avhich Aveighed rather more than 3 kilo., containing
over six million eggs. In fecundity the Conger is thus
by no means inferior to the common Eel. The actual
spawning has never been observed; but the young have

been diligently studied in recent times, especially by
the Italian Facciola6. He distinguishes between six
different Leptocephalus forms belonging to this species,
more or less distinct transition forms Avhich compose
an unbroken series from the most degenerate larva?,
Leptocephalus incequalis , or from the longer knoAvn
Leptocephalus Morrisii, to the easily recognisable fry
of Conger niger. The first-mentioned larva? are sharp-
nosed, transparent, slender, but deep behind the middle
of the body, are furnished Avith comparatively large,
subulate jaw-teeth, and have the beginning of the dorsal
fin situated in the posterior third of the length of the
body, the vent in the hindmost fifth thereof. Lepto-
cephalus Morrisii is blunt-nosed, Avith prominent tip
of the snout, terete, though transparent, and of more
uniform depth, has lost all the jaAV-teeth or the greater
number thereof, and has the beginning of the dorsal
fin situated at a distance from the tip of the snout
measuring about 28 % of the length of the body, the
vent at a distance from the same point measuring about
38 % of the said length. The ascertained fry of Con-
ger niger are brown on the back, and at first sIioav
two longitudinal bands of chestnut-broAvn. As Ave have
already remarked, however, the average size of the last-
mentioned, least degenerate forms may be less than
that of Leptocephalus incequalis.

The subsequent growth of the Conger is apparently
very rapid. Jackson, the manager of the Southport
Aquarium, added to the collections a number of Con-
gers weighing 2 — 3 lbs. (0'9 — 1*36 kilo.), Avhich he
assumed to be about a year old. Five years aftenvards
(in 1880) one of these specimens died, and Avas sent
to Buckland. It measured 6 ft. 5 in. (1,956 mm.)
and weighed 90 lbs. (40'8 kilo.).

In aquaria the Conger is easily kept alive, but its
ravenous hunger renders it a dangerous comrade to its
fellow-captives. “We find these congers,’’ Avrote Jack-
son, “are the most voracious creatures Ave keep; they
attack and devour even dogfish, and these of a size
that one Avould think beyond their powers of SAvalloAving.
We have quite given up keeping the piked dogfish with
them, and even the topers must be big ones, or down
they go. The only safe things are our big sturgeons,
monk-fish, skate, and the huge turbot Ave have reared
from little ones (these latter continue to grow). Con-

a Nat. Hist. Brit. Fish., p. 387.

b' ll Natnralista Siciliano, Anno XII, 1893, p. 194.

1042

SCANDINAVIAN FISHES.

gers swallow their prey head first, as a rule, but when
committing an act of cannibalism they swallow their
small brother often, if not always, tail first. If the
youngster is a bit too big, you may often see him
three parts swallowed, and when the big one is quite
exhausted the other will wriggle out none the worse,
except that it is scratched by the big one’s teeth. We
have had this happen so often with the same fish that
at last it has got as ragged and full of scratches as it
could hold; some day, however, one of the big ones
has given a mighty gulp, and once let its jaws close
over the head of the little one, and we see it no more”.
In the fisherman’s boat the Conger struggles violently

to escape; it bites at everything within its reach, and
with the aid of its prehensile tail casts itself overboard,
unless stunned in time. Even when cut off from the
body, the head of the Conger has been known to in-
flict a dangerous bite.

In Scandinavian waters the Conger, on account of
its small numbers, cannot repay any special fishery.
Where it is more plentiful, large quantities are taken
on hand-lines and long-lines; but its flesh is inferior
and full of bones. It is eaten boiled, or the flesh is
dried and ground to a powder, which is used as an
ingredient in soups. The tough skin may be employed
in the same manner as that of the common Eel.

CHONDROSTEOUS FISHES.

1043

PISCES CHONDROSTEI

(STURGEON-FISHES).

Fishes with endoskeleton principally cartilaginous, but with dermal ossifications repre-
senting several of the internal bones of the Teleosts, with shoulder-girdle suspended from
the head, with maxillary and palatine arches free from the skull or united therewith
merely by a mobile connexion, with branchial cavity, which is situated under the skull
and common to the free branchial arches, more or less entirely covered by the opercula,
with fully heterocercal caudal fin, and with paired fins unilaterally rayed. Nostrils lateral,

set just in front of the orbits.

We have briefly mentioned above (p. 1) the most
important anatomical points wherein the Teleosts differ
from other piscine orders, and we have now to examine
the relations obtaining between those typical fishes
which in general organization are by no means inferior
to the Teleosts, but which never attain so high a degree
of skeletal differentiation or ossification as the latter,
and have therefore borne the general name of carti-
laginous fishes. As a rule, though Avith important ex-
ceptions, Ave find that in these forms the ossification
deficient in the endoskeleton is compensated in the
dermal system, Avhich is strengthened or protected Avith
hard, thick (Ganoid) scales, spiny plates, or scutes. In
Artedi" they Avere included, together with the Cyclo-
stomata, in a single order, Chondropterygii, which Avas
retained under the name of a series by Cuvier6, in
contradistinction to the “true fishes” (Teleosts). Agassiz
divided this order into two: the Ganoidei c, among
which he further ranged, on account of their hard
dermal groAvths, several Teleosts (the Glanoinorphs,
Plectognates, and Lophobranchs), and the PlacoideC ,

among Avhich he placed the Cyclostome fishes too.
Bonaparte e introduced the first reform, and removed
the Cyclostomata to a separate subclass, Marsipobranchii ,
as opposed to the subclass of the Chirmeras, Sharks,
and Rays, Elasmobranchii, and MulleiC eliminated
from the order of the Ganoids the said Teleosts. The
delimitation of the Ganoids from the Teleosts, hoAvever,
Avas no longer based exclusively either on the enamelled
scales of the former or on the less advanced calcifica-
tion (ossification) of their skeleton, but mainly on the
characters adduced above (p. 1 ) and derived from the
more intimate fusion (chiasma) of the optic nerves after
their emergence from the brain, and the prolongation
of the heart, at the transition to the common branchial
artery, into a muscular conus arteriosus , furnished in-
ternally with several roivs of valves. Thus defined the
Ganoids still comprised, in addition to the Chondrostei,
the multitude of tishes, possessing more complete jatv-
bones, but noAv for the most part extinct, in Avhich the
piscine type has evolved by manifold processes a Avealth
of forms rivalling that of the Teleosts, but of which

“ Gen. Pise., p. 64.

b R'egn. Anim ., ed. 1, tom. II, p. Ill; ed. 2, tom II, p. 128.
c Rech. Poiss. Foss., tom. II.
d „ „ „ tom. III.

e Selacliiorum tabula analytica , Rom® 1839.
f Abh. Akad. Wiss. Berk 1844, p. 147.

1044

SCANDINAVIAN FISHES.

only two types survive, inhabitants of the tropical re-
gions of Africa (the Bichir, Polypterus) and America
(the Bony Pikes, Lepidosteus). Lutken a restricted the
scope of the Ganoids to the last-mentioned division
(Muller’s Holostei), but ranged them as well as the
Sturgeon-fishes (Muller’s Chondrostei ) in the category
of distinct subdivisions among the Teleostean Physostoms.

The free gill-arches alone, with the branchial la-
mellae projecting beyond their free, outer (convex) mar-
gin, are enough to distinguish the Sturgeon-fishes from
the true cartilaginous fishes and approximate them to
the Teleosts, with which they are also allied by several
other points in their organization, e. g. the covering of
the head with membrane-bones that distinctly represent
the several component bones of the Teleosteous cranium,
and the structure of the generative organs. Their in-
clusion within the order of the Teleosts is indeed for-
bidden not only by the above-mentioned characters
( cliiasma and conus), but also b}^ the structure of the
extremities, wherein the Sturgeon-fishes have preserved
a great portion of the primordial cartilaginous radialia,
as well as by the undivided chondrocraniuin and the very

partially transformed notochord. But Lutken is, beyond
doubt, fully justified in his opinion that the Sturgeon-
fishes have far more in common with the Teleosts than
with the true cartilaginous fishes. Doderlein* 6 too com-
bined them together with the Teleosts and the true
Ganoids into an order, Teleostomi, characterized by the
completely ossified skeleton or at least by membrane-
bones on the head and shoulder-girdle, by the free
(joined to the cranium by articulations, sutures, or
ligaments) hyomanclibular and palatoquadrate arches',
and by the unarmoured pectoral fins'7. Among the
Teleostoms they are distinguished by their fully hetero-
cercal caudal fin, with the tip of the spinal column ex-
tending quite or nearly to the end of the upper caudal
lobe, and by their unilaterally rayed' paired fins. In
Doderlein they rank, with these characters, as a sub-
order ( Heterocerci ). In the Scandinavian fauna, which
contains neither Lung-fishes ( Dipnoi ) nor Ganoids, it
may suffice to have drawn attention to the relation of
the Sturgeon-fishes to the Teleosts and the true carti-
laginous fishes, which is adequately expressed by Mul-
ler’s interpretation of them as a distinct order.

Fam. ACIPENSERlDfiE.

Body fusiform , with, five rows of large osseous bucklers. Four barbels in a transverse row on the under surface
of the snout, in front of the protrusile mouth. No branchiostegal rays.

In modern times the Chondrosteous order contains
only two families, one of which, the American and
Chinese Polyodontidce, with almost or quite naked body-
and without barbels, is principally distinguished by its
comparatively large mouth, open even laterally (on the
sides of the head), and with upper jaw incapable of
protrusion, the anterior extremity of the palatine arch
being firmly united by ligaments to the under surface
of the skull. Another important difference is that the
Polyodontidce are without pseudobranchiae, those of the
Sturgeons being exceedingly well developed, even func-
tional as gills, with at least superiorly free branchial
lamellae, on the inside of the opercula.

On opening the pharyngeal and branchial cavities
of a Sturgeon, the most striking difference from all
the Teleosts meets us in the roof of the palate. This
(fig. 284, mx — mpt) is a continuous, convex disk, with
free margins both in front and behind, and capable,
together with the toothless jaws and fleshy, tumid lips,
of protraction forwards and downwards and of retrac-
tion. On each side of the deep fold that hangs, covered
with the thick mucous membrane of the pharynx, be-
tween the free hind margin of the palatine roof and
the under surface of the skull (parasphenoid bone) or
posterior, fixed palatine roof with its continuation, the
roof of the pharynx, we find an aperture. This hole

a Om Ganoidernas Begroendsning og Inddeling, Vid. Meddel. Naturh. For. Kbhvn, 1868, sep.

6 Steinmann and Doderlein, Elemenie der Paldontologie, pp. 540 and 556.
c As opposed to the Lung-fishes.

d As opposed to the primeval and extinct Placodermi (Pterichthys, Cephcilaspis, etc.).
e Not lobate, or but slightly so, in contrast to the strongly lobate fins of the so-called Crossopterygii.

STURGEON-FISHES.

1045

leads into a canal which runs upwards, skirting ihe
anterior margin of the great upper suspensorium Oto-
mandibular, Jim) common to the whole apparatus of
the palate and jaws, and opens (tig. 28G, spir) on the
top of the head, a little behind the eye, at the outer
margin of the osseous plate covering the squamosal
part (squ) of the skull, and foreshadowing the squa-
mosal bone. The upper opening is called the spiracle,
and the canal, which as well as the former is entirely
absent in the tishes we have hitherto considered, but
which in all higher animals, man included, has its
homologue in the external and internal auditory meatus
and the tympanum, is here furnished in its lower
(inner) part, just above the internal aperture, with a
row of branchial lamella?, not respiratory, however,
for they are supplied with arterial blood. The entire

to the suboperculum and interoperculum. In the Sterlet
these lamella? are for the most part free; in the com-
mon Sturgeon they coalesce throughout the greater part
of their external margin with the operculum, only their
tops being free. In one of the Ganoids (Lepidosteus)
Muller" has shown that the upper part of this row of
gill-lamina1 loses its functional importance as a respi-
ratory organ, and it is thus fully homologous with the
pseudobranchiae of the Teleosts. Behind the pharyngeal
cavity the branchial cavity of the Sturgeon shows the
same structure as that of the Teleosts, four gill-arches,
each with two rows of branchial lamella? and short,
scattered gill-rakers; and the fifth arch (fig. 284, hbr5
+ cbr5), the lower pharyngeal of the Teleosts, furnished
with one row of gill-rakers, is here, too, destitute of
branchial lamella?, and has no gill-slit behind it, being

P.5

Fig. 284. Forepart of the endoskeleton in a Sturgeon. Partly after J. Muller and Parker.

B, basal angle; 6’, notochord; cbr , — cbr., first — fifth ceratobranchial cartilages; cehy, ceratohyoid; costl — cost8, first — eighth ribs; csp, spira-
cular cartilage (according to Parker); cle, dental part of mandible; etl, lateral ethmoid; hbr] — bbr-, first — fifth hypobranchial cartilages; hhy,
hypohyoid; hm , hyomandibular cartilage, with the upper part covered by a parostotic disk; M, articular part of the mandible (hind part of
the Meckelian cartilage); mx, maxillary; rn.pt, metapterygoid cartilage; N, neural arches; Na, nasal cavity; obsp, orbitosphenoid ; opt, foramen
of the optic nerve; pi, palatine; pop, preoperculum (according to Parker); Ps, neural spines; psp, parasphenoid ; pt, pterygoid; qu, quadrate
cartilage; 11, rostral cartilage; spl, symplecticum ; stliy, stylohyoid; tr, foramen of the nervus trigeminus', vg, foramen of the

nervus vagus', vom , vomer.

canal is analogous to a gill-slit, between the palato-
mandibular and hyoid arches, the latter of which fur-
nishes with its upper parts (the hyomandibular, fig.
284, hm, and symplecticum, spl ) a suspensory apparatus
to the former as well. The gill- slit next behind (between
the hyoid arch and the first branchial arch proper) has
its branchial lamella? set in a large, but single, arcuate
row on the inside of the operculum, throughout the
hind margin thereof, as well as on the inside of the
plates situated below the. operculum and corresponding

firmly coalescent. with the hind wall of the branchial
cavity or anterior side of the scapular arch.

In an adult state the mouth of the Sturgeons is
entirely toothless; but their larva? have teeth, in form
and distribution not unlike those of the Sharks, in their
corneous structure resembling those of the Lampreys,
- both in the upper and lower jaws and on the cerato-
branchial bone ( cb)\ ) of the first branchial arch proper.
These teeth are developed soon after the palato-mandi-
bular arch, originally continuous and arcuate, has brok-

Abh. Akad. Wiss. Berlin 1844, p. 133, taf. II, fig. 1 and taf. V, fig. 6.

1046

SCANDINAVIAN FISHES.

en up into two parts, an upper, which has medially
grown together from the sides to form the framework
of the palatine roof and an analogue to the quadrate
bone (fig. 284, qu), and a lower (the Meckelian carti-
lage), which forms the framework of the lower jaw (de
+ M). At the age of three months these teeth dis-
appear. Meanwhile the palato-quadrate cartilage has
differentiated into two firmly coalescent, but distinguish-
able parts, the posterior (mpt) answering to the meta-
pterygoid of the Teleosts, the anterior corresponding
principally to their pterygoid, proper and quadrate (qu),
but also acquiring at its anterior margin special ossi-
fications, homologous with the palatines (pi) and the ento-

Fi°\ 285. Ossifications on the under surface of the skull in Acipen-
ser sturio. J/4 of the natural size, psp, parasphenoid; pa, its ascend-
ing sphenoid process; vom, vomer; scr , vomerine scutes.

pterygoids ( pt ). There further appears on each side of
the anterior margin of the palatine arch a freer bone,
the maxillary (mx), which extends from the anterior
tip of the palatine disk (pterygoid bones) to its lateral
extremity (the knob by means of which the quadrate
bone articulates with the mandible), leaving between
itself and the palatine disk a fissure, through which
the levators of the lower jaw find a passage. On the
outside of the above-mentioned juncture between the

maxillary and the quadrate lies a triangular ossifica-
tion (pop), most pointed above, which lias been inter-
preted by Parker as representing a preoperculum; and
on the outside of the Meckelian cartilage (M) is deve-
loped the dental part (de) of the lower jaw.

In the skull itself there appear ossifications the
largest of which (psp) answers to the parasphenoid of
the Teleosts, and extends from the nasal region not
only under the skull, but also, divided into two lateral
plates (fig. 285), under that part of the spinal column
which has not been completely divided into separate
vertebras, but in the form of a cartilaginous mass, con-
tinuous below, composes a backward prolongation of
the cranial cartilage. This parasphenoid is covered
underneath, throughout the greater part of its length,
by the hard mucous membrane alone; but between the
nasal and orbital regions (in the ethmoidal region) it
pierces the downward projection (fig. 284, B, the so-
called basal angle, which may also be observed in the
Sharks) of the chondrocranium. Within this it meets
and wedges itself into the vomer (vom in figs. 284
and 285) in front of it, which advances under the
rostral cartilage (fig. 284, B), and is continued in its
turn by several vomerine scutes (fig. 285, scr), evi-
dently belonging to the skin. In the ethmoidal region
(the lower anterior part of each orbit) the lateral eth-
moids (fig. 284, etl) are developed; above the foramen
of each optic nerve (in the arched roof of the orbit)
appears an orbitosphenoid bone (fig. 284, obsp), behind
this a smaller osseous disk, corresponding to the ali-
sphenoid, and behind the orbit, round the orifice for
the nervus trigeminus (tr), an osseous disk answering
to the petrosal. But all these bones, according to Par-
ker", are superficial growths external to the cartilage
( parostoses ), foreshadowing the cartilage-bones (ectosto-
ses and entostoses) present at the same points in the
Teleosts and the higher vertebrates. Similar foresha-
dowers appear in the form of osseous scutes (scale-
growths) in the skin on the top of the head and outside
the shoulder-girdle. At the middle of the occiput, but
firmly united to the first dorsal scute, lies a trefoiled
(posteriorly broad, with a narrower lobe projecting in
front) osseous plate, evidently answering to the supra-
occipital (here called the suprdoccipital scute, fig. 286,
ocs) of the Teleosts. Further forward lie two larger
scutes, usually the largest on the whole head, whose

Philos. Trans. Roy. Soc. London, vol. 173 (1882), p. 175.

STURGEON-i'ISIIES.

1047

place is occupied in the skull of the Teleosts (see, for
instance, the cranium of the Eel, p. 1014, tig. 268, B ,
par) by the parietals ( parietal scutes , tig. 286, par).
Their hind extremities are parted by the said project-
ing lobe of the supraoccipital scute; they are sometimes
contiguous throughout the rest of their length (tig. 286,
B), and in other cases their anterior ends are separated
(tig. 286, A) by an osseous plate (et) fitting in between
them from in front. Before them and partly on the
sides of their anterior extremities lie a pair of plates
( frontal scutes, fr), often but slightly inferior to them

A

smaller plates. All these variations may be observed
in the same species, e. g. in the common Sturgeon.
The inconstancy of form depends upon the comparatively
lax connexion between these scutes and the skeleton,
the looseness increasing towards the snout, which is gene-
rally covered with numerous, more irregular, smaller
plates, except on the sides, which are armoured in adult
specimens of the common Sturgeon with large scutes in
a definite row. Outside each parietal plate lies a squa-
mosal scute (by others called the temporal scute, squ),
and in front of this, above the orbit, on the outside ot

Fig. 286. Upper surface of the head iu the Sturgeon (Acipenser sturio). of the natural size. Two figures showing variations iri this

species; both specimens taken at Christianstad.

dy, first dorsal scute; et, ethmoidal scute; ett, ectethmoidal scute; fr, frontal scute; mt , mastoid scute; n, anterior nostril; o, eye; ocs, occi-
pital scute; par, parietal scute; ptob, postorbital scute; ptp, posttemporal scute; spho, sphenotic scute; spo, supraorbital scute; spir, spiracle;

squ, squamosal scute; tp, temporal scute; op, operculum.

in size. These correspond in situation to the frontal
bones of the Teleosteous skull, and are sometimes close-
ly applied to each other, but usually separated at least
in front (fig. 286, B), sometimes throughout their length
(fig. 286, A), either by an elongated ethmoidal scute (et),
which may even extend, as mentioned above, between
the anterior extremities of the parietal scutes, or by
several (up to 5 or more and sometimes two-rowed)

each frontal plate, a curved row of three scutes, the
posterior representing a postfrontal bone ( sphenotic scute,
spho), the anterior a prefrontal bone ( ectethmoidal scute,
etl), and the middle one a supraorbital bone ( supra-
orbital scute , spo). The lower end of the sphenotic scute
touches the top of an angular plate, the form of which
calls to mind the preoperculum of the Teleosts, but
which here composes the posterior and inferior limits

Scandinavian Fishes.

132

1048

SCANDINAVIAN FISHES.

of the orbit, and is hence known as the postorbital scute
( ptob ). Behind each squamosal plate we sometimes find
two plates, sometimes only one“, touching the outer
sides of the parietal, occipital, and first dorsal scutes.
Where the plates are two in number, the anterior is
evidently homologous with the mastoid of the Teleosts
( mastoid scute, mt ), and the posterior, which has re-
ceived the name of the temporal scute ( tp ), is the upper-
most plate in the curved row covering the scapular arch
of each side. It rests on (covers) a backward prolonga-
tion of the mastoid angle of the chondrocranium, just
as the first dorsal scute is placed on a similar process
springing from the supraoccipital part of the chondro-
cranium. We have already mentioned (p. 635) Parker’s
comparative investigations wherein lie traced the homo-
logues of the human clavicle in different vertebrates
from the very stage where they are mere dermal growths

even in its cloven form, with the posttemporal bone of
the Teleosts. It stands out, however with equal dis-
tinctness as the first plate in the upper lateral row of
body bucklers, and forms the superior part of the hind
limit of the branchial cavity, but lies as a tegumentary
bone on two cartilages, one of which (the upper) is
the top of the above-mentioned process issuing from
the mastoid part of the head, while the other (fig. 287,
ssc) is the uppermost, segmented part of the shoulder-
girdle proper, a part which has disappeared in the
Teleosts, where the dermal plate has also been trans-
formed into a portion of the endoskeleton. The median
part of the hind limit of the branchial cavity consists
chiefly, sometimes entirely, of a large plate, the clavi-
cular scute (cl), answering to the clavicle of the Tele-
osts. Often, however, there is inserted, close behind
the upper part of this plate, a smaller one, the s upra -

Fig. 287. Left scapular scutes and cartilages of a Sturgeon (Acipenser sturio). l/2 of the natural size. A, seen front within ; B, from without.

After Parker.

The cartilaginous parts of the shoulder apparatus are: ssc, pars suprascapular is b ; sc, pars scapularis; cr, pars coracoidea; ecr, pars epicora-
coidea; per, pars prcecoracoidea', fnc, fenestra coracoidea ; /. cs, fenestra coraco-scapularis. The scales (plates) transformed into tegumentary
bones for the shoulder-girdle are: ptp , posttemporal scute; cl, clavicular scute; id, interclavicular scute; scl , supraclavicular scute.

(fish-scales); and it was principally from the Sturgeon
that he traced his comparisons. On the outer and
posterior sides of the temporal scute, or in the same
relative position to the mastoid scute, when this is
confluent with the former, lies a vertical plate, more
or less deeply forked above, on which Parker con-
ferred the name of the posttemporal scute (figs. 28G
and 287, ptp), and which is evidently homologous,

clavicular (scl), corresponding to the supraclavicle of
the Teleosts. The lower part of the clavicular scute
curves inwards, over the ventral side, but the greater
part of the external scapular arch on each side consists
at this point of the interclavicular scute (id), the homo-
logue of the interclavicle remarked above among the
characters of the Hemibranchs. The inner parts of
these plates are perfect bones, and as in the Teleosts,

“ This variation too may be observed within the limits of the same species, for example in the common Sturgeon.
h Above the suprascapular part, below the top of the posttemporal plate (ptp), lies the tip of the prolongated mastoid cartilage,
not shown in the figure.

ST D RGEON-FISHE S .

1049

in a manner that reminds us especially of the Trunk-
fishes (see above, p. 619), both the clavicle and the
interclavicle are continued inwards by an entire osseous
disk, forming the posterior wall of the branchial cavity.
Within (above) the anterior extremities of the inter-
cdavicles, where these bones meet in the median line of
the belly, we find, situated under the conns arteriosus ,
and forming, as it were, a special protection for the
same, a fiat, heart-shaped bone. Its original dermal
structure is indicated merely by a small, terete protu-
berance, even externally visible, and it is apparently
to be regarded from a morphological point of view as

R 12 3 4

Fig. 288. Left pectoral fin of a male Sturgeon (Acipenser sturio )
1.845 mm. long and taken at Lulea on July 18th, 1893. '/2 of

the natural size. Seen from above.

R, radiale of the first (composite) ray, answering to Gegenbaur’s
mesopterygium; 1 — 4, the four true radialia, the last of which (4)
answers to Gegenbaur’s metapterygium.

representing the sternum of the higher vertebrates
{sternal scute).

The internal parts of the shoulder-girdle — omitting
the already mentioned suprascapular part (s.sc) — form
a. continuous mass of cartilage; but in adult Sturgeons
this mass is seen to consist of three several parts, the
same as we have seen above in Glanomorphs, Cyprino-
morphs, and Thrissomorphs, namely an upper part, an-
swering to the scapula (sc), a lower part, answering to

the coracoid bone (cr), and an inner anterior part, cor-
responding to the precoracoid (per). At the line where
the scapula and coracoid meet (at f. cs), the pectoral
fin is articulated, in the same manner as normally meets
us among the Teleosts; but the articulary surface of the
large first ray (spinous ray) glides partly on a projec-
tion of the clavicle, an articulation that calls to mind
the pectoral fin of the Sheatfish. In the basal structure
of the fin we find (fig. 288) the same four radialia
(brachial bones, 7 — 4) as in the Teleosts, increasing in
length backwards (downwards). But these are cartila-
ginous and divided into two joints (an inner and an
outer row, cf. above, on the Herring, p. 951); and at
the hind inferior margin of the fin there are several

Fig. 289. Left ventral fin and pelvic bone of the same Sturgeon as
in the preceding figure, th of the natural size. Seen from above.

for , foramen obturatorium (?); mtp, metapterygium; pit, processus
iliacus(?); pp , processus pubicus (?); 1 — 7, radialia.

supplementary radialia, the inner ones branching off,
however, from the outermost (hindmost) radiale, called
by Gegenbaur the metapterygium. A similar, but still
greater superfluity of cartilaginous radialia appears in
the basal parts of the ventral fins (fig. 289). The num-
ber of radialia is greater in the Sterlet than in the
Sturgeon (fig. 289, 1 — 7), being in the former at least
9, in the latter 7; but in the former the first three,
in the latter the first four radialia (basalia.) articu-
late at the proximal (inner) end with a common
cartilage, answering to the pelvic bone of the Tele-

1050

SCANDINAVIAN FISHES.

osts“; whereas the others (the posterior six) articulate
in the Sterlet, according to von Rautenfeld, each with
a separate cartilage, parallel to the pelvic bone, but
evidently of the same nature as the radiale, from which
it has been separated by constriction. Similarly a small
piece of cartilage is cut off from the distal (outer) end
of most of the first-mentioned radialia, so that three
rows of these bones are formed, the innermost row con-
taining the pelvic bone and, behind this, the inner ra-
dialia. In the Sturgeon, on the other hand (fig. 289),
the posterior (3) radialia are distinctly inclined at
angles to each other, and so arranged that only one
of the innermost parts ( mtpt ), common to them all, ar-
ticulates with the pelvic bone, and with the top of this
part are jointed two cartilages, one belonging to the
fifth radiale, the other forming a common base for the
last two (6th and 7th) radialia. All the radialia thus
lie on one side of an imaginary axis, drawn through
the inner margin of the pelvic bone, and continued
outwards by the hindmost (innermost) radiale. The
pelvic bone is now* * * * * 6 interpreted as being formed by the
coalescence of the inner parts of the anterior radialia;
and at the inner end of each pelvic bone we find in
adult Sturgeons*' a constriction setting off the part (pp)
which, according to Wiedersheim, is the true rudiment
of the pelvis of the Selachians and the higher verte-
brates. The difference in the number of the radialia in
the Sterlet and Sturgeon is the expression of a conti-
nued reduction, which has finally brought about their
disappearance in by far the greater number of the Te-
leosts. On the other hand, the similarity in structure
of the original basal parts of the pectoral and ventral
fins, their composition of articulated radialia, covered
at the distal ends by the bases of the true (secondary)
fin-rays, is the expression of the common origin of
these fins. They have been produced by the differen-
tiation of a. fold running along each side of the belly,
now persistent only in the Lancelet, in the same man-
ner as the dorsal and anal fins have originated from
differentiations in the region of the embryonic vertical
fin. The one-sided arrangement of the radialia — set

more or less distinctly at angles to a more developed
basal part, which is originally situated at the hind
margin of the fin — is best explained by a comparison
with the transformation of the caudal fin from diphy-
cercy to heteroeercy. Fishes d in general began by making
their caudal fin - — their earliest organ of locomotion -
heterocercal, for the attainment of a more highly deve-
loped musculature, concentrated on one side, to steer and
accelerate their movements. The same alteration was ex-
tended to the other vertical fins, dorsal and anal, the
anterior margin and its basal parts being strengthened
and developed into organs for cutting and stemming the
water or into weapons of offence and defence, while the
posterior parts grew more mobile with more numerous
divisions, but with the outer (distal) joints arranged in
series on one side of the more or less confluent basal
portions (Gegenbaur’s metapterygium). Having once
been established among the vertebrates, this manner of
development spread to the lateral fins, to the fore
and hind limbs of the highest vertebrates. These too
became unilateral.

Among the remaining skeletal peculiarities most
characteristic of the Sturgeon-fishes is the persistency
of the notochord with only slight alteration, without
undergoing such coarctations or constrictions as attend
the development of perfect vertebrae. It is enveloped,
however, by a comparatively thick sheath (perichord,
fig. 290, Cs and Ee ), in and upon which there develop
cartilages representing both neurapophyses ( N ) and
haemapophyses (77), as well as upper spinous processes
( Ps ), which parts, however, remain separate. Between
the apophyses lie strengthening disks (inter cal aria, Ic),
and the luemapophyses are prolongated into transverse
processes, bearing cartilaginous ribs, and also grow
inwards (Fo) under the notochord, where they sur-
round in the caudal region both the aorta and the
caudal veins, in the abdominal region only the former.
Above the spinal cord, which is enclosed on each side
by the apophyses, runs another similar canal, formed
by holes through each vertebra, and containing an
elastic, longitudinal, tendinous band (El).

a Cf., however, our remarks (above, p. 635) on the pelvic bones of the Hemibranchs, which bones are evidently dermal growths of

the nature of interspinal plates. The normal pelvic bones of the Teleosts, on the other hand — as appears from their form — are homolo-

gous with interhcBinal bones (supporting bones of the anal and caudal fins), and we now see that the pelvic bones of the Sturgeons are in-

termuscular growths, and have the same origin as the supporting bones, of the vertical fins, ontogenically being confluent constrictions of the

supporting cartilages of the ventral fins.

6 Cf. Wiedersheim, Das Gliedmassenskelet der Wirbelthiere, Jena 1892, p. 70; Grundriss Vergl. Anat. Wirbelth., Jena 1893, p. 184.
c As in Polyodon and Scaplrirhynchops , see Wiedersheim, 1. c.

d Smitt, Ur de hogre djurens utvecklingshistoria (lectures for 1873), Stockholm 1876, p. 230.

STURGEON -F I S HES .

1 05 1

Of the system of the lateral line (dermal sense-
organs) we find in young Sturgeons pores in the plates
of the lateral line and in the temporal and mastoid
plates, as well as a connecting canal (a junction com-
mon in the Teleosts too between the two supratemporal
canals, cf. S])t in tig. 104, p. 368) across the occiput,
between the mastoid and supraoccipital plates. On each
side of the head the main canal runs forward under
the squamosal plate, to branch at the upper posterior
corner of the eye downwards to the sphenotic and post-
orbital plates, and forwards, under the inner margin
of the supraorbital and ectethmoidal plates, in the der-
mal bridge between the nostrils. The system is most
distinct, however, on the snout, especially on the under
surface thereof, where the four filaments of touch (bar-
bels) of the Sturgeon a, re also situated. Here we find
partly small, simple pores (ordinary lateral line pores),
partly agglomerations of these, forming wheel-shaped
figures that cover large muciferous cavities. These ca-
vities (the ScMeimsdcke or Nervensdcke and Gallert-
rohren of Leydig, the tubes muceux a ampoules of Du-
meril) occur in the common Sturgeon, being most
distinct in young specimens, 1) below the eyes and on
the upper surface of the snout in a row along each
margin, forward from the nasal cavity, 2) on the under
surface of the snout, sometimes throughout its extent.
The small, simple pores are set on the under surface
of the snout in a single or double row, curving in-
wards from the lower posterior angle of each sub-
orbital plate, and thus answering to the suborbital
branch of the Teleosts, but running forwards, outside
the outermost barbel, and discernible in young speci-
mens to the very tip of the snout.

The variations in the form and arrangement of the
body bucklers belong to the description of the several
species; but common to all the Sturgeon-fishes and
most of the Ganoids are the so-called fulcra , imbricated
splints, resembling spinous rays, at the anterior mar-
gins of the vertical fins. The posterior fulcra of the
several fins, however, more and more assume the form
of true fin-rays, and thus clearly show a transition be-
tween these growths. The fins of the Sturgeon-fishes
a.re furnished, like those of the Teleosts, with secondary,
partially ossified and sagittally (in the longitudinal di-
rection of the body) divided rays, originally all simple
and articulated. But with age these rays become more
and more confluent and, in the anterior part of the
fins, hardened, as in most of the Physostoms, so that

the foremost rays are spine-like, though with distinct
vestiges of the original articulations.

In the internal structure of the Sturgeon-fishes we
have already remarked the most essential respect where-
in the heart differs from that of the Teleosts. In a
male Sturgeon measuring 1,845 mm. from the tip of
the snout to the end of the upper caudal lobe, the
length of the abdominal cavity is about 700 mm. The
thick-walled oesophagus runs in this specimen for a
distance of about 9 cm. close under the spinal column,
and is internally furnished here anteriorly with 5, poste-
riorly with 8 series of hamate or (behind) valvular
flaps, exchanged furthest back for shallow, flat folds
of the mucous membrane. Where these folds terminate,
the mucous membrane of the stomach, which is some-
what wider, but has thinner walls, commences, and
about 4 cm. further back opens the short, but wide,
pneumatic duct of the large and thick-walled air-bladder.

Fig. 290. Transverse section of the anterior part of the spinal co-
lumn in Acipenser ruthenus. After Wiedershkim.

Ps , upper spinous process; El , elastic tendinous band; N, neurapo-
physis; SI, fibrillar tissue (belonging to the skeletogenous layer);
M, medulla; P, inner membrane (phi) of the myelon; Ic, intercalare;
C, notochord; Os, inner chordal sheath; Ee, outer (elastic) chordal
sheath; H, hsemapophysis; Z, basal part of a haemal arch; Fo , cross-
pieces projecting mediad (inwards), and ventrally (interiorly) covering
the main trunk of the arterial system (aorta, Ao).

The stomach now assumes a more intestine-like form,
and runs upwards to the left, to a distance of about
23 cm. from the diaphragm, where it turns forwards
to the right and downwards, to a point (tig. 291, v)
distant about 3 cm. from the diaphragm, where it
again bends back and is thickened. At each crook its
inner surface is furnished with thick, terete folds of
the mucous membrane, 8 at the former bend, 6 at the
latter; but between these points the inside is smooth,
and the last-mentioned folds are continued on the inner
surface of the pylorus (pi/l), which is directed back-

1052

SCANDINAVIAN FISHES.

wards, has very thick walls, and is bounded from the
retral duodenum ( duod ) by a strong, annular valve.
Just behind this valve the upper wall of the intestine
is pierced with a large hole (app), wide enough for
the insertion of a finger, and t wo somewhat smaller
holes, outlets of the large pancreas ( per ), which is
flesh-coloured with yellowish brown spots, and has been
formed by the coalescence of the ctecal pyloric append-
ages", being therefore coursed by numerous, tubular
passages ( cap ). This gland is of a flattened elliptical
shape, about 12 cm. long, 8 cm. broad, and 2 1/3 cm.
thick, with sharpened edges. It is situated in the
circle formed by the above-mentioned gyrations of the
intestiniform stomach; and in the present case it oc-
cupies rather more than two-thirds of this circle. The
duodenum (duod) runs back to a distance of 37 cm.

depressions on the inside. A similar, though somewhat
finer network appears on the mucous membrane lining
the rectum (isp), which is about 50 cm. lorig.'' Here
we find an apparatus peculiar to the Sturgeon-fishes,
Ganoids, Lung-fishes, and Selachians. The valve is con-
tinued by a spiral, in the present case with eight coils,
which extends almost to the termination of the rectum.
The spiral consists of a thick raised margin (sp), which
shows the same depressions as the network of the mu-
cous membrane, though more sparsely, and which runs
along the inside of the upper rectal wall, being .con-
tinuous therewith; but at eight points it detaches itself
from the roof, and is attached instead by a backward
membranous spiral (msp) to the rest of the intestinal
wall. About 2 1/2 cm. before the vent the central cord
of the spiral terminates in a tubercular swelling (tub),

Fig. 291. Viscera of a male Sturgeon (Acipenser sturio ) 1,845 min. long, taken at Lulea on July 18tli, 1893. About 3/J0 nat. size.

Seen obliquely from below.

abd, right ( abs , left) wall of the abdominal cavity opened and folded back; a pm, inner mouth of the left Mullerian duct (M) ; app, largest
of the pancreatic orifices visible in the opened beginning of the duodenum (duod)', cap, section of one of the pancreatic canals; duod, duo-
denum; h, front part of the liver; hd, right lobe of the liver; isp, spiral intestine, opened; l, spleen; M, left Mullerian duct; msp, spiral
membrane, the lateral membrane of the spiral valve; per, pancreas, sectioned and with one (the ventral) half somewhat raised to show the
canals (cap) within it; pijl, pylorus; sp, spira, central cord of the spinal valve; tel, right, ts, left testis, with the posterior half of the latter
cut loose and laid back behind the left wall of the abdominal cavity; tub, tubercular swelling of the spira ; v, termination of the procurvated
stomach; ves, dotted line indicating the outlines of the posterior part of the air-bladder, otherwise concealed; i if, gall-bladder;

* — **, mesorchial thickening of the right testis.

from the diaphragm, where it abruptly bends forward
to a point distant about 20 cm. from the same, and turns
back with equal abruptness, soon passing into the straight
(spiral) intestine, from which it is divided by an an-
nular valve. Throughout the course of the duodenum
the mucous membrane forms a uniform network of

and the extreme end of the rectum has tive longitu-
dinal ridges with rather sharp margins on its inside.
The liver (// — ltd) of the present specimen is of a. bluish
black colour0. The left lobe is about 21 cm long, the
right lobe (hd) 1 7 1/2 cm. The gall-bladder (vf), about
36 mm. long and 23 mm. broad, lies embedded in

a Cf. their structure in the Tunny, see above, p. 99.

b We here adopt Rathke’s division of the intestinal canal. According to others (see, for instance, Wiedersheim, Grundr. Vergl. Anat.
Wirbelth., p. 412), the greater part of what we here coll the rectum, so far as the spiral valve extends, should be regarded as a part of
the middle intestine.

c According to Kroyer it is of a ruddy yellowish brown.

ST U I iG E O N -El S H E S .

1053

the lateral anterior margin of the right lobe; and the
gall-duct ( ductus choledochus communis ) opens on the
right side of the above-mentioned pancreatic orifices.
The spleen (/), about 1372 cm. long and Sl/2 cm. broad,
is pointed behind and forked, being divided from in
front, throughout the greater part of its length, into
two anteriorly pointed lobes, the lower of which lies
under and to the left of the above-mentioned first crook
of the stomach, behind and projecting a little way over
the pancreas, while the other lobe is laid to the right
of the mesenterial fold joining the stomach to the intes-
tine. The testes ( ts and td) of the present specimen,
which was taken in the middle of July, are very tumid,
and had nearly attained their annual ripeness. They
extend throughout the greater part of the length of the
abdominal cavity, the left being, however, longer than
the right. Throughout their length they are divided
into irregular lobes by transverse incisions and shal-
lower constrictions. The length of the left testis (ts) is
55 cm., its greatest breadth 7Vg cm., and its greatest
thickness 4 cm. The corresponding dimensions of the
right testis (td) are respectively 49 cm., 6 cm., and 3
cm. So oblique is the relative position of each testis to
the other that the hind extremity of the left only ex-
tends to a point distant 11 cm. from the anal aperture,
a distance which in the case of the right testis is re-
duced to 7 cm.“ Their outside (facing the wall of the
abdominal cavity) is convex, their inside flatter; the
former is covered with an extremely thin membrane,
the latter with a thicker continuation of the mesorchium,
the peritoneal membrane that holds each testis suspend-
ed partly (in front) from the wall of the abdominal
cavity, partly (further back) from the air-bladder, partly
(behind the air-bladder, which terminates about 1972
cm. in front of the vent) from the kidneys. Posteriorly
the mesorchium is thickened (* — **), on the left testis
for a distance of about 10 cm., on the right for about
8 cm. Behind this thickening the testes are free from
their suspensory membrane, the left only for a, very
little way, the right for a distance of 672 cm. On the
inside, in front near the middle of the breadth or even
near the lower, free margin of the testicle, behind
near its upper (suspensory) margin, runs a wide ef-
ferent duct (vas deferens ), the radices of which ramify
in all parts of this side. The efferent duct is skirted

by a sharp-edged, rather thin, adipose band, with free
margin and in some places 1 cm. broad, at other
points narrower or even merging into the mesorchium.
Air blown into the efferent duct spreads partly into
its ramifications, partly, in the hind portion of the
testis, into large lacunae between the testicular mem-
brane and the stroma. Mercury injected into the
duct penetrates through canals in the thickened part
(* — **) of the mesorchium into the kidneys. It is
thus evident that the sperma may find egress in this
direction and through the ureter; but another passage
seems also to be afforded. Both male and female Stur-
geon-fishes possess peritoneal funnels, not unlike those
we have seen above in female Sal monoids, especially
in the Smelt and Capelin. But here these funnels are
developed into independent ducts (71/), distinct from
the true organs of generation, densely ciliated, and
opening behind into the ureter. They are thus fully
homologous with the so-called Mullerian ducts (ovi-
ducts) of the Ganoids, Lung-fishes, Selachians, and the
higher vertebrates. In the males of most vertebrates
they are more and more reduced, the testes being fur-
nished instead with deferent canals through the pro-
nephros; but in the Sturgeon-fishes this transformation
has not advanced so far, the males having as large,
as open, and apparently quite as functional Mullerian
ducts as the females. In the above-mentioned male
Sturgeon the inner (abdominal) aperture (apm) of the
left Mullerian duct lies about 31 1 2 cm., that of the
right 321/2 cm., in front of the anus; and the former
duct measures 15 cm. to its posterior aperture, which
opens into the ureter. Frequently*, at least in male
and female Sturgeons not yet in spawning condition,
the last-mentioned aperture is closed; but to judge
by the perfect development of the Mullerian ducts in
our specimen, it would seem quite probable that they
may be functional in breeding fish of both sexes, even
if the sperma of the males may also find its way into
the ureter by another mode of passage.

On each side of the anus the Sturgeon-fishes are
commonly furnished with a. so-called abdominal porec,
an aperture b}^ means of which the abdominal cavity
communicates with the surrounding water. It is often
wanting, however, as in the specimen just described.
Its function is somewhat doubtful, though probably

a In another Sturgeon the relations were reversed; the left testis extended further, even in a backward direction, than the right.
h Cf. Jungersen, Bidr. Kundsk. Konsorg. Udv., Disp. Kbhvn 1889, p. 101.
c Cf. Weber, Morphol. .Tahrb., P>d XII (1886), p. 366.

1054

SCANDINAVIAN FISHES.

that of a respiratory organ. It is a relic of the seg-
mental organs found in lower animals, and recurs in the
Selachians and "Ganoids, as well as in the Salmonoids,
Enchelymorphs, and Mormyroids among the Teleosts.

fhe family of the Sturgeon-fishes contains only
two, not very dissimilar genera, S c a p hirhynchops and

Acipenser. The former, of which only one species is
known, with spatulate snout, long but shallow (depressed)
peduncle of the tail, with the upper lobe of the caudal
fin prolongated into a filamentous appendage, and further
characterized by the absence of spiracles, belongs ex-
clusively to the southern regions of North America.

Genus ACIPENSER.

Peduncle of the tail terete (not depressed ) and at least

meet from each

The Sturgeons have met with the same variety of
systematic treatment as the Salmons, and probably from
the same cause. They are anadromous fishes, breeding
in fresh water — a few, it is true, permanent inhabitants
thereof — but else living in the sea. Their geographical
range is about the same as that of the Salmons, ex-
tensive enough to offer a great variety of physical
environments. They also show an inconstancy of form
fully comparable with that of the Salmons, and the
methods of their classification have been equally diver-
sified. In 1870 Dumeril described" 81 species of this
genus, among them 62 from America. In the same
year Gunther adopted in his Catalogue only 19 spe-
cies, 9 American. Jordan and Gilbert6 (1883) re-
cognised only 5 species in America, 2 from the Pacific
coast, 3 from the Atlantic; but the specific rank even
of these may be called in question.

The characters hitherto employed in the definition
of the species are subject to considerable variations.

It has long been known that the large body bucklers
of young Sturgeons are set closer together, and have
a longer, more pointed, and usually more hooked, cen-
tral spine, than those of older specimens. Their num-
ber was indeed recognised by Gunther as a valid
character, but varies in the common West European
and American Sturgeon, for example, between 11 and
13 in the dorsal row and between 26 and 34 in the
upper lateral row. According to Jordan and Gilbert
these variations extend between 11 and 14 in the

twice as deep as the lateral caudal plates , which do not
side of the body.

dorsal row and between 27 and 36 in the upper lateral
rows, according to Kroyer between 10 and 14 in the
former and between 26 and 31 in the latter. Fatioc
counted 15 plates in the dorsal row. Of the small
plates in the skin between the large bucklers Gunther
remarks, in the case of the common Sturgeon, that in
very young examples (which thus would be referred
to the genus Huso of Dumeril) the skin is provided
with very small rough points; in older ones these ossi-
fications are broader, rough, substellate, and more (as
in the genus Acipenser of Dumeril) or less (as in An-
taceus of Dumeril) regularly arranged in oblique se-
ries d. According to Milner’s observations e of the
American fresh-water Sturgeon, the large bucklers in-
crease in size until the fish has attained a length of
about 63 or 64 cm., but afterwards diminish, partly
owing to the detrition of the spines, partly by resorp-
tion of their margins, and both at the dorsal line and
the ventral margins they partially drop off, or at least
become indistinct. Simultaneously with this process
may be observed a shortening of the snout; and the
far greater relative length of the snout and its more
pointed form in young Sturgeons give the head an
appearance quite different from that of older specimens.

The external sexual characters are not marked,
and but little has been observed on this head. In the
Seuruga (Slier g, Acipenser stellatus ) of the Black Sea and
its feeders, which species is characterized by a very long
and slender snout, the females, according to Heckel

a Nouv. suites a Buffon , Hist. Nat. Poiss., tome II, pp. 87 cett.,
b Bull. U. S. Nat. Mus., No. 16, pp. 85, cett.
c Fne Vert. Suisse , vol. V, part. II, p. 491.

d The same observation had already been made by Nilsson ( Skand . Fna, Fisk., p.
systematic significance.

c U. S. Comm. Fish and Fisher., Rep., part. II (1872 and 1873), p. 70.

702), though he did not expressly point out its

STURGEONS.

1055

and Kner", have a shorter snout, a weaker spine in
the pectoral fins, smaller dorsal and lateral scutes,
with longer hooks than in the males. Of the Sterlet
(Ac. ruthenus), however, the same writers /j remark that
the females have a longer, thinner, and more upturned
snout and a flatter forehead. In the common Sturgeon
(Ac. sturio ) we have found no perceptible external
difference between the sexes, save that the ventral fins
of the males are not removed so far back during
growth as those of the females, the length of the pec-
toral fins even in old (more than 1 m. long) males
being at least J/5 of the distance between the ventral
fins and the tip of the snout. The males thus seem
to represent the characters of the early stages of growth.
Of the Hausen too Meckel and Kner'; state that the
females attain the greatest size. The materials within
our reach as well as the recorded observations, how-
ever, leave much to be desired in our knowledge of
the external differences of sex.

The form of the Sturgeons is also influenced by
their environments. “Some forty years ago”, wrote
Grimm'' in 1883, “the sterled penetrated through the
canals into the North Dvina, and finding the condi-
tions favourable to its existence (for ex. cold water,
which is so necessary for it) it not only settled down
and multiplied, but acquired some peculiarities in its
exterior (a short, blunt snout and an arched back)
and also a fine flavour, for which in Petersburg it is
prized more than the Volga sterled. I must remark
that even in the system of the Volga the sterled is
much finer in the north (for instance in the river
Sheksna) than in the southern parts, and the further
south one goes, the less tasty the sterled becomes”.
Among the Sturgeons, as among the Salmons, certain
forms' are sometimes found landlocked in fresh water
and unavoidably debarred from exchanging at regular
intervals, in obedience to the natural instinct of the
genus, fresh-water life for a marine existence. It is

only to be expected that such forms should suffer
alteration.

The actual number of species within the genus
it is thus impossible as yet to state ’with certainty.
In Europe 7 or 8 species are supposed to occur, most
of them belonging to Russia and the Danubian coun-
tries. In Scandinavian waters only one species of the
genus is found, the common Sturgeon of Western
Europe. King Frederick I. of Sweden attempted to
plant the Russian Sterlet in Edsviken (near Stockholm)
off Ulriksdal, an experiment which at first seemed to
promise success, but eventually proved a complete failure.

The Sturgeons have been highly esteemed from
time immemorial for their flesh, their roe, and the
gelatine extracted from their air-bladder. Gregarious
and at certain seasons extremely sluggish, they are
an easy catch; and at other times, when they are more
active, their great timidity drives them blindly into
the fisherman’s snares. Some of them are among the
colossi of the piscine world: the Hausen, for instance,
is stated to have attained a weight of more than 1,500
kilo., and specimens weighing 550 kilo, are still taken
from time to time. A single fish of the latter weight
commands at Astrachan a price of about £22 ■ . The
Sturgeon-fishery is consequently of great importance
and, especially in Russia, a productive source of food
and income. According to DanilovskG the annual
take of Sturgeons in European Russia may be approxi-
mately valued at eight million roubles (£1,250,000).
Of this sum about five millions are represented by
the flesh, about two millions and a quarter by cavi-
are, about 600,000 roubles by isinglass, and about
100,000 roubles by vjeziga, the notochord (chorda
dorsalis ), which is principally employed in the pre-
paration of stock for soups and as an ingredient in pies''.

Acipenser* 1 is classical Latin, and was introduced into
ichthyology, as a generic name, by Artedi. In Pliny
the Sturgeon is called AttilusJ and Mario.1 Among the

"■ Susswasserf. Ostr. Mon., p. 345.

6 l. c., p. 339.
c 1. c., p. 569.

d Fish., Hunt, in Russ. Wat., p. 24.
e E. g. Acipenser rubicundus in America.
f Grimm, 1. c., p. 32.
g See Grimm, 1. c., p. 26.

h For a description of the Sturgeon-fishery in Russia, see Brehm, 7 hierleben, Gr. Ausg., 1879, Abtli. Ill, Bd. 2, p. 358.

1 Gesner supposed that the word had arisen by letter-changes from the Greek O^VQQVyygg.

■’ Hist. Mundi , lib. IX, cap. XIII.
k 1. c., lib. IV, cap. XV.

Scandinavian Fishes.

133

1056

SCANDINAVIAN FISHES.

Greeks the genus bore the names of Ellopsa, Antaceusb,
and several others, the application of which is, however,
disputed. In mediaeval times the name of Sturio was
coined from the Teutonic Stoer, which according to W OR-

mitjs c is the same as the Scandinavian stor (great), but
according to Gesner'' should be derived from the Teutonic
storen (to root, stir up the bottom), a reference to the
method practised by the Sturgeons in procuring their food.

THE STURGEON (sw. storen).

ACIPENSER STURIO.

Plate XLVI, fig. 1.

Dorsal bucklers mesocentric ( with the boss uppermost in the middle , and sloping forwards and backwards). Num-
ber of plates in the upper lateral row as a rule less than 37e. Length of the snout about l/2 (53J — 42 %) of
that of the head , which occupies about 27 — 20 % of that of the body. Width of the mouth , which , when pro-
truded, is square with rounded angles, at most about 2/3 of the breadth of the snout, at the barbels. Barbels
terete, simple ( not fimbriated g), and shorter than the distance between them and the anterior margin of the mouth.
Base of the dorsal fin less than 1/12 of the length of the body. Length of the pectorals less than V 4 of the

distance between the ventrals and the tip of the snout.

V.

R. hr. 0; D.
26—27(30); C.

30—40(44); A.
26 — 29 super.
80 — 100 infer.

23—26(30);

P. 36—40(42);

Syn. Acipenser sive Sturio , Sohonev., Ichthyol. Slesv. Hols., p. 9.
Sturio, Willughb., Hist. Pise., p. 329, tab. P. 7, fig. 3.
Acipenser corpore tuberculis spinosis aspero, Art., Ichthyol.,
Gen., p. 65; Synon., p. 91; Lin., Fna Suec., ed. I, p. 101;
It. Scan., p. 187.

Haae-Storje (Sturio), Str6m, Sondm. Beskr., pt. I, p. 286.
Styria, Olafs., Reise Isl., pt. II, p. 711.

Acipenser europceus, Lin., Mus. Ad. Frid., p. 54, tab. XXVIII,
fig- 2.

Acipenser Sturio, Lin., Syst. Nat., ed. X, tom. I, p. 237 ;
Mull., Zool. Dan. Prodr., p. 39; Bl., Fisch. Deutschl.,
pt. Ill, p. 89, tab. LXXXVLII; Retz., Fna Suec. Lin.,
p. 309; Ekstr., Vet. Akad. Handl. 1831, p. 104; Nilss.,
Prodr. Ichthyol. Scand., p. 109; Bdt, Ratzeb., Medic.
Zool.., vol. II, pp. 17 et 352, tab. Ill, fig. 1 et M — S;
Parn., Mem. Wern. Nat. Hist Soc., vol. VII, p. 403; Bonap.,
Fna Ital., Peso. (tom. Ill, 2), tab. 129, fig. 1; Yarr., Hist.
Brit. Fish., vol. II, p. 475 et Suppl. (Richards.) II, p. 7;
Kr., Damn. Fisk., vol. Ill, 2, p. 747; Nilss., Sic, arid. Fna,
Fisk., p. 699; Hckl, Kn., Siisswasserf. (Jstr. Mon., p.362;
Mgrn, Finl. Fiskfna (disp. Helsingf. 1863), p. 70; Malm,
Gbgs Vet. Vitt. Samh. Handl., Ny Tidsf., H. VIII (1863),
p. 102; Steind., Stzber. Akad. Wiss. Wien, Math. Naturw.
CL, LIII, 1 (1866), p. 204; v. Bemm. in Herkl., Bouwst.
Fna Nederl., tom. Ill, p. 318; Lindstr., Gotl. Fisk., Gotl.
L. Hush. Sallsk. Arsber. 1866, p. 24 (sep.); Hum., Hist.

Poiss. (N. Su. a Buff.), tom. II, p. 184; Gthr, Cat. Brit.
Mus., Fish., vol. VIII, p. 342; Coll., Fork. Vid. Selsk.
Chrnia 1874, Tillsegsh., p. 205; 1879, No. 1, p. 102; N.
Mag. Naturv. Chrnia, Bd 29 (1884), p. 115; Malm, Gbgs,
Boh. Fna, p. 604; Winth., Naturh. Tidskr. Kblivn, ser. 3,
vol. XII, p. 55; Fedders., ibid., p. 93; Mor., Hist. Nat.
Poiss. Fr., tom. I, p. 471; Bncke, Fisch., Fischer., Fiscliz.
O., W. Preuss., p. 191; Id. in M. v. d. Borne, Handb.
Fiscliz., Fischer., p. 181; Doderl., Man. Ittiol. Medit ., pt.
II, fasc. I, p. 6; Jord., Gilb., Bull. U. S. Nat. Mus., No. 16,
p. 85; Mela, Vert. Fenn ., p. 362, tab. X; Mob., Hcke, Fisch.
Osts., p. 149; Day, Fish. Gt. Brit., Irel, vol. II, p. 280,
tab. CL; Lillj., Sv., Norg. Fislc., vol. Ill, p. 488.
Acipenser Lichtensteinii, Bl., Schn., Syst. Ichthyol., p. 348,
tab. 69; Bdt, Ratzeb., 1. c., pp. 21 et 352, tab. II, fig. 1.
Acipenser oxyrhynchus, Mitch., Trans. Lit., Phil. Soc. N. York,
vol. I, p. 462 (+ 7 A tip. sturio, p. 461 = Acip. brevi-
rostrum, Lesueur, Trans. Am. Phil. Soc., vol. I, p. 390);
Dek., N. York Fna, pt. IV, p. 346, tab. LVIII, fig. 189;
Stor., Mem. Am. Acad. Arls, Sc., vol. VIII, p 431, tab.
XXXV, fig. 4; Dum., 1. c., p. 106.

Acipenser latirostris, Parn., 1. c., p. 405, tab. XXXIX.
Acipenser huso , Thomps., Ann. Mag. Nat. Hist., vol. XX, 1847, p.

172; Sundev., Ofvers. Vet. Akad. Forh. 1853, p.228. NecLiNNE.
Acipenser Thompsonii, Ball, Proc. Irish Acad., No. 25, p. 21;

Tiiomps., Nat. Hist. Irel, vol. IV, p. 245.

Acipenser liospitus, I\r., 1. c., p. 780.

Acipenser sturioides , Malm, Gbgs Samh. Handl., 1. c., p. 108;
Gbgs, Boh. Fna, p. 605.

“’EXXoxp, Aristot., Zool., lib. II, capp. 13 et 15.
h Herodot., Hist., lib. IV, cap. 53.
c Cf. Cttarleton, Onomast . Zoic., p. 152.
d De Aquat., p. 932.

e The specimen described by Richardson, (in Yarr., Brit. Fish., ed. 2, Suppl. II, p. 21) from the Free Kirk College of Edinburgh,
with 38 plates in the upper lateral row on the right side and 40 on the left, has been shown by GOnther to belong to the American spe-
cies Acipenser rubicund, us, which may thus be assumed lo have strayed into European waters. But Day (Fish. Gt. Brit., Irel., vol. TI, p.
279) questions the Scottish origin of the specimen. Malm counted in a Swedish specimen 38 plates in the said row.
f In a young specimen — probably an exceptional case — 57 %, according to KrOyer.

9 In old Sturgeons, however, the barbels are sometimes broader, compressed, and more or less distinctly fimbriated.

STURGEON.

1057

The ordinary size of the Scandinavian Sturgeon is
about 1V2 — 2 m. In the island-belt of Soder mankind,
according to Ekstrom, small specimens 3 — 6 dm. long
are commonest. The young Sturgeon, 18 cm. long,
figured by Linnaeus, is still preserved in the Royal
Museum; but whether it was taken in Scandinavia, is
uncertain. Among the specimens measured by Kroyer
the three smallest are 16 — 32 cm. long; but he does
not state whether they are Scandinavian. Malm’s small-
est specimen was 39 cm. in length. On the other
hand, rather large specimens, measuring up to 3 in.,
are more frequently found; and Lindstrom records the
find in Gothland of a dead, stranded Sturgeon which
was stated to have been 14 Sw. feet (415 cm.) long.
According to an old statement in Schonevelde, the
Elbe Sturgeon has been known to attain a length of
18 feet (5 2/3 m.)“.

The body of the Sturgeon is of an elongated fusi-
form shape or, when the fish is spent, clavate, thickest
at the occiput. It tapers forward in a highly charac-
teristic manner, due to the pyramidical form of the
snout, and also deviates behind from the ordinary pis-
cine type in its oblique, prolongated, and heterocercal
caudal fin. The greatest depth of the body, which is
usually deepest just behind the occiput, is about 78 —
Vio of its length, and the greatest thickness about
710 — V9 °f the same. The least depth, just in front
of the caudal fin, is only about 3 % of the length, but
increases with age (during the growth of the Sturgeon
from a length of 16 cm. to one of 185 cm.) from 11
to 16 % of the length of the head, or from 23 to 28 %
of the length of the pectoral tins. The fusiform shape
of the body is considerably modified, however, especially
during youth and in lean specimens, by the large shield-
rows, which render it pentagonal in section. The shield-
rows belong to the dorsal line, the lateral line, and
the side margins of the belly; but on the under sur-
face of the tail, between the vent and the anal fin, the
ventral rows coalesce, at least partially, into one, which
is again divided, however, behind the anal fin; and on
the back of the tail, behind the dorsal fin, the shield-
row of the dorsal line commonly, but often irregularly,
breaks up into two rows, which may also be pre-
indicated in front of the dorsal fin by one or two pairs
of plates, smaller than the rest, being fitted in before

a Belon (Nat., Div. Poiss., p. 89) also mentions a specimen
at Montargis to King Francis I.

b I. e. of a texture that distinctly calls to mind the plates o

its fulcrum. In shape and relative position the buck-
lers show considerable variations, individual, it is true,
appearing even in specimens of equal size, but evidently
also expressing the alterations of growth. In young
Sturgeons they are more densely set and deeper, with
sharper and more pointed, recurved spine, which is
furnished with small spines on its sloping hind margin.
With age the spines are obliterated, the plates become
shallower (flatter), and separate more and more from
each other in the several rows. Yet it applies to these
alterations of growth, as to so many others, that they
appear more distinctly in one individual than in an-
other, and have consequently occasioned the designation
of the same species by a plurality of names. As a
rule the spines are most persistent on the posterior

body bucklers. The plates of the dorsal line are most

curved, showing in young Sturgeons an acute-angled
transverse section; the plates of the lateral lines are
least curved, especially in the anterior parts thereof;
the plates of the ventral line, even during youth, are
most remote from each other. The surface of the
scutes is scabrous and both radially and concentrically
striated with cavities and ridges, calling to mind the

scale-texture we have seen above in the Eels. In the

dorsal and lateral lines the anterior scutes are shorter
(comparatively broader) than the posterior. The form
of the plates further varies from the rhomb, which pre-
ponderates in the ventral lines and the anterior part
of the dorsal line, or hexagon (rhombs with truncated
anterior and posterior angles), which appears in the rest
of the dorsal line, to the semicircle or triangle, as shown
in the lateral lines. The foremost (properly the only6)
fulcrum supporting the anterior margin of the vertical
fins is linguiform in the dorsal and anal tins, with the
narrower end directed up the fin-margin: at the upper
and lower margins of the caudal fin the tongue is pro-
longated to a lanceolate form. The inconstancy in the
number of plates in the dorsal line before the dorsal
fin and in the lateral lines has been mentioned above.
Behind the dorsal fin 4 pairs are set, as a rule, in front
of the caudal fulcrum; but on each side of the posterior
part of the dorsal fin the small dermal plates are usually
enlarged with age, so that a more or less regular row
of 2 or 3 plates, smallest in front, appears on each side
at this point, forming a. forward continuation of the

18 feet long.

: the body.

A Sturgeon of this size was said to have been presented

1058

SCANDINAVIAN FISHES.

dorsocaudal row. The plates of the ventral lines vary
in each row within the preabdominal region, as a rule,
between 10 and 12; but Krdyer once found only 8, and
Liel.jeborg adduces as many as 14. Behind the vent
there generally lie 2 or 3 pairs in front of the anal
fulcrum, or the posterior of these pairs coalesces into
one plate. Behind the anal tin are set 3 — 5 (in excep-
tional cases only 2) pairs of distinct plates in front of
the lower caudal fulcrum ; but to the posterior part of
the anal fin the same remarks apply as to the corre-
sponding part of the dorsal. In the preceding pages
we have also considered the transformation of the re-
maining dermal covering from small scattered spines
and their more or less distinct agglomerations, simul-
taneously with their adoption of a blunter, more tuber-
culate form, into the diamond-shaped dermal scutes,
arranged in oblique transverse rows, with small rows of
tubercles between them, that are especially prominent
on the dorsal sides above the lateral lines. On the tins
and their rays the small, sharp spines are persistent;
but on the upper part of the caudal tin, on each side
of the continuation of the spinal column within this
tin, elongated, diamond-shaped, and smooth bony plates
are developed, arranged in oblique rows running from
below and in front upwards and backwards.

The hind part of the head is rather terete, con-
stricted above at the gill-openings and eyes, with more
or less depressed interorbital space. A striking diffe-
rence from the head of the Teleosts is that the opercula
do not entirely cover the gill-openings, the gills of the
Sturgeon being consequently laid bare to some extent
behind. In front the head is depressed and of varying
length, tapering in a pyramidical form, with the superior
surface of the snout more convex than the inferior and 1
the other two sides narrowing sometimes to an edge.
In young Sturgeons the relative length of the snout, as
we have mentioned above, is considerably greater than
in old, varying with age from about 14 % at least to
about 83/4 % of the length of the body, or from 53 %
(according to Kroyer sometimes 57 %) to 42 % of that
of the head, so that the entire length of the head is also
comparatively less in the latter. The postorbital length
of the head, on the other hand, relatively remains al-

most unaltered during growth, being about V]0a °f the
length of the body. The eyes are small and somewhat
oval. Their longitudinal diameter, which shows relative
diminution during the growth of the fish, varies between
10 and 7 % of the length of the head, between 19 and
14 % of the length of the snout, or between 37 and 22 %
of the interorbital width6. The interorbital space, which
is convex, but medially more or less depressed, grows
narrower in proportion to the general growth, varying
between about 7l/2 and 6V2 % of the length of the body,
but wider in proportion to the length of the head,
varying between about 27 and 33 % thereof.' In front
of the eyes lie the large nostrils — comparatively larger,
however, in young Sturgeons than in old — the posterior
in each pair being the larger, an obliquely or even a
transversely set ellipse, the length of which is some-
times greater than the diameter of the eyes, and situated
low down; the anterior more extended in the longitu-
dinal direction of the body, and set nearer to the level
of the forehead. They are separated by a thin (in old
specimens rather broad) dermal ridge. The shortening
of the snout during the growth of the fish is accom-
plished at the expense of its anterior part (the rostral
cartilage), the distance from the anterior nostril to the
tip of the snout being sometimes reduced with age
from 47 to 28 % of the length of the head.

The under surface of the head, like that of the
belly, is rather plane. Behind the perpendicular from
the centre of the eyes is situated the comparatively
small mouth0. Owing to its structure, which we have
described above, it may be protruded (in a downward
direction), so as to form a square tube, rounded, how-
ever, at the corners (fig. 292). The toothless jaw-margins
are rounded. The lips are tumid, on the upper jaw
rather uniformly, but with a median indentation, at the
corners of the mouth in a bulging form, on the lower
jaw in the form of two fleshy folds, which are closely
applied to each other from the sides, but distinct, the
jaw itself being medially naked. Half-way between the
tip of the snout and the mouth — sometimes, in young
Sturgeons, distinctly nearer to the latter, sometimes, in
old, nearer to the former d — hangs the transverse series
of barbels characteristic of the Sturgeons. These are 4

a Varying, according to our measurements, between about lO1/^ and 9 1 9 /.

b For the vertical diameter of the eyes the corresponding percentages are respectively 8 and 5'/2! 16 and 12, 30 and 16.
c One of the most reliable distinctions from the Hausen ( Acipenscr huso ) and the Osse'tr ( Tok , Ac. Gulden stcidtii), which have a
broader mouth.

d The distance between the barbels and the tip of the snout measures 64 — 44 % of that between the mouth and the same point.

STURGEON.

1059

in number and of fairly equal length, sometimes ex-
tending, when laid back, rather near, but never quite
to the anterior margin of the mouth, seldom indeed to
the hollow in the ethmoidal cartilage for the reception
of the latter. In the median line of the under surface
of the snout there lies in front a series of tuberculated
bony plates in the skin, paired or, at the very begin-
ning, set in several irregular rows. These rows termi-
nate behind, between the middle barbels, in a plate
which does not belong exclusively, however, to the skin,
but is merely the prominent, though sometimes concealed,
head of the vomer (fig. 285, p. 1046), whose hind ex-
tremity, as we have described above, is embedded in
the cranial cartilage and meets the parasphenoid bone.
Thus we have here a most excellent anatomical example
of the morphological transition from der mo-ossifications
to parts of the endoskeleton.

The plate-armour of the head we considered above,
when we selected the Sturgeon as our type of the ge-
neral conformation of these parts within the family.
There remains only the opercular apparatus, which ex-
ternally resembles that of the Teleosts, but is charac-
terized by the absence of rays in the thick branchiostegal
membrane. The gill-openings extend right across the
sides of the body, the branchiostegal membranes being
united below to the isthmus in a line with the upper
angles of the gill-openings, and separated from each
other by a distance hardly greater in young Sturgeons
than the diameter of the eyes, but which widens in
older specimens until it is at least half of that between
the upper angles of the gill-openings. Each branchio-
stegal membrane contains three bones, in a row one
below another. The uppermost is the operculum, which
is naked throughout the greater part of its extent, and
shows the same radial striation and thimble-like granu-
lation of the surface as the dermal scutes. In form it
resembles a sector of a circle, somewhat greater than a
quadrant, and with the inferior radius incurvated. At
the anterior part of its lower angle, and united by a
firm suture to its inner surface, is inserted the more
elongated suboperculum, which lies across the body,
and in old Sturgeons is mostly covered by the skin, but
on the outside has a high ridge, above sending out
through the skin a few granulated spines. The ridge
is continued on the lowest bone, explained as an inter-
operculum, which has a more quadrilateral form, but
is irregularly incised, with radially disposed ridges on
the outer surface.

The dorsal fin lies far back, farther in old Stur-
geons than in young. The distance between its begin-
ning and the tip of the snout increases with age from
about 62 to 69 % of the length of the body. Its last
ray is so small in old Sturgeons that its trapezoidal form
approaches to the triangular. The upper posterior mar-
gin is concave. The length of the base varies between 6
and 8 % of the length of the body, and the height is
about the same, somewhat greater or less. The first three
or four rays in old Sturgeons are very short, thick, and
hard, resembling fulcra. As a rule, the first seven or
eight ra}'s are simple, gradually increasing in length,

| and the ninth or tenth ray is the longest in the fin.
The anal fin is opposed to the posterior part of the dorsal,
and resembles the latter fin, but has a shorter base
— varying between about 3 and 4 '/2 % of the length of
the body — and consequently a more pointed form. The
form and covering of the caudal fin we have already
noticed. The upper lobe (the very tip of the tail), which

Fig. 292. Head of the Sturgeon, with mouth protruded like a proboscis-

After Benecke.

in adult specimens (more than 2 dm. in length) has all
its upper rays, except the outermost 3 — 5, transformed
into hard fulcra, is nearly twice as long as the lower.

The pectoral fins are set low, almost in the plane of
the belly, and horizontally. They are obliquely pointed
in form (fig. 288, p. 1049), with rounded inner posterior
angle. Their length varies between about 12 and 11 %
of that of the body, or between about 22 and 1 8 '/2 % of
the distance between the ventral fins and the tip of the
snout, affording an easy distinction between the present
species and the Sterlet, which has perceptibly longer pec-
toral fins. The first ray is considerably stronger than
the rest; but distinct traces in its structure mark it as
the result of a coalescence of several (8 — 10) simple and
articulated rays. The ventral fins are inserted behind
the middle of the body, at a distance from the tip of
the snout measuring about 54 — 59 % of the length of the
body. Their form (fig. 289) is less pointed and more
trapezoidal than that of the pectorals. Their length dur-

1060

SCANDINAVIAN FISHES.

ing youth is about 55 %a, in old specimens about 63—
65 % of that of the pectoral tins. At the base of the
anterior margin are set two short and thick supporting
rays, similar to fulcra, and the following 6 or 7 simple
rays gradually increase in length out to the tip of the tin.
Both in form and structure the ventral tins are thus not
unlike the vertical. The preabdominal length measures
about 33 — 37 %, the postabdominal about 12%, of the
length of the body. The latter (the distance between
the anterior margins of the ventral and anal fins) is,
however, not fully entitled to the above name, for the
vent lies rather far in front of the beginning of the
anal tin, the distance between them being 5 — 7 % of
the length of the body.

The coloration of the Sturgeon is simple, its salient
features calling to mind that of the Codfishes6. The dor-
sal side is of a reddish or bluish gray, gradually paling-
down the sides of the body, though a distinct boundary-
line, about half-way between the plates of the lateral
lines and those of the ventral margins, divides this colour
from that of the belly, which has a bluish tinge above,
sometimes with a silvery lustre, and below is pure white.
Above the ventral tins, however, runs a broad stripe, in
an obliquely forward direction, joining the bases of these
tins to the dorsal coloration. In young Sturgeons Kro-
ner found, “between the plate-rows of the dorsal and
lateral lines, blackish spots, giving this part a chequered
appearance”. The outer parts of the tins are of a more
or less pronounced ash-gray, becoming paler towards
their bases. The pectoral tins are darkest, but share,
as well as the dorsal and caudal, iii the coloration of
the back; the ventral and anal tins are palest. The iris is
of a brassy yellow; the black pupil has a dash of green.

The Sturgeon is a salt-water fish with the migratory
instinct of the Salmons, an anadromous form that as-
cends from the Atlantic into the rivers of North Ame-
rica and Europe in order to spawn. From the Atlantic
its range extends into the Baltic and the Mediterranean
including the Adriatic Sea. Whether it penetrates into

the Black Sea", is doubtful. Neither Heckel and Knee
nor Nordmann found it there''. In the Baltic, on the
other hand, it makes its way to the head of the Gulf
of Bothnia, according to a statement by Wi degree", even
up the Tornea Elf. Now and then it is met with off
all parts of the east coa.st of Sweden as well as on the
Finnish coast, and occasionally it ascends into the Swe-
dish lakes7; but it is common in the Baltic proper only
to the south and in the German rivers falling into those
waters6'. It is common in all Danish waters, says Win-
tiier; but as the Danes do not ply any special Stur-
geon-fishery, it is not seen very often. The case is the
same throughout the west coast of Sweden, especially
off Mount Kullen and in Laholm Bay; and in the Gota
Elf it has been found at Lilia Edet, 33 miles above
Gothenburg (Nilsson). In Norway too the same ob-
servation has been made: the Sturgeon is not taken
anywhere in numbers, but solitary specimens are fre-
quently met with throughout the coast-line, both in the
sea and the river-mouths, even east of the North Cape
(Collett and Storm). Off’ Iceland it is rare. It is
common enough in the rivers running from the east
into the North Sea. In Great Britain and Ireland it is
not much commoner than in Sweden; but into the
French rivers with a westward course, especially into
the Garonne, the Sturgeon ascends more frequently and
in greater numbers (Moreau). Its most constant habitat-
lies, however, in the Mediterranean and Adriatic as well
as the rivers flowing into these seas. On the east coast
of North America its range extends from Cape Cod to
Florida. A fresh-water form, considered to be a distinct
species'1, inhabits the Mississippi Valley, the Great Lakes
with their feeders, and the basin of the Albany River.
On one occasion this American Sturgeon is supposed to
have extended its wanderings into Scotch waters, and
Gunther* * was of opinion that the young Sturgeon from
Bohuslan, described by Malm under the name of Aci-
penser sturioides, should be referred to this species. So
wavering are the specific distinctions within the genus;

° Sometimes 50, according to Kroyer.

* The name of ovio/.og was thus applied by the ancient Greeks both to Codfishes and the Sturgeon.
c Grimm, 1. c., states that it occurs there.

d In the Black Sea and the Danube lives a species very nearly allied to the Sturgeon, the Ossetr ( Toks , Acipenser Guldenstadtii) , which
has a broader mouth. Cf. below, as to the difference between the American fresh-water Sturgeon and the present Sturgeon.
c Landtbr. Akad. Hnndl., I8:de delen (1858), p. 181.

/ According to Lilljeborg a specimen has been found in Lake Limmar (Roslagen, just south of Norrtelje).
g In the Baltic Provinces of Russia the Sturgeon is rare, according to Seidlitz, Fauna Baltica, p. 89.

h Acipenser rubicund, us = Ac. maculosus. According to Milner it has a broader mouth than our Sturgeon. No other distinction of
any significance seems to be given between them.

1 Cat. Brit. Mils., Fish., vol. VIII, p. 339, note.

STURGEON.

1061

and the suspicion readily suggests itself that the Ame-
rican fresh-water Sturgeon is a landlocked form of the
common Sturgeon, and stands in the same relation to
the latter as Salmo trutta to S. salar.

Of the life and habits of the Sturgeon we possess
but scanty information. Most of its existence is passed
at the bottom, where it is beyond the reach of all cu-
riosity. To judge by its behaviour in aquaria, it is a
sluggish, though petulant creature; and it usually glides
at a gentle pace over the bottom, shooting out and
drawing in its mouth; but of a sudden, awaking, as it
were, from its drowsy laziness, it darts through the
water with the speed of an arrow. It is extremely
tenacious of life, and in a cool place may be kept alive
out of the water for twenty-four hours or even longer.
“It seems a spiritless fish”, says Pennant®, “making
no manner of resistance when entangled, but is drawn
out of the water like a lifeless lump”. By the asser-
tion of its great muscular strength, however, it some-
times gives formidable proof that it is by no means
insensible to danger and sufferings, and a blow dealt
by the tail of a large Sturgeon is enough to fracture
an arm or a leg. Besides it is far from always so in-
different to its fate as to abandon itself to the toils
without more ado. The fishermen frequently complain
that it breaks loose from their nets. When the Stur-
geons are found in rivers Avhich they have ascended in
order to spawn, and when they are distressed by the
pressure of the tumid generative organs, they are slug-
gish from natural causes; but “in the sea, where they
are sometimes stated to be sterile, they are evidently
not ripe for the time being, being in a certain sense
virgin, and consequently far more active. Herr Dec-
ker lias communicated a proverb current among the
Sturgeon-fishermen of the North Sea, which runs, Leap-
ing Sturgeons and dancing girls are hard to hold fast"1'.
The Sturgeon also resembles the Salmons in its habit,
when migrating, of now and then flinging itself entirely
out of the water.

The Sturgeon feeds in the manner just described
on the worms of the bottom, crustaceans ( Amphipoda ),
and mollusks, but also, like several Cyprinoids, on the
decomposing animal substances to be found in the mud.
The stomach of a male which was taken at Dalaro on

the 18th of July, 1890, and which is represented in
our figure (Plate XL VI, fig. 1), was filled at the an-
terior crook with a score or two of Isopods (. Idothea
entomon). That the diet of the Sturgeon consists prin-
cipally of Herring, Mackerel, Cod, and Salmon, the last
of which it chases up the rivers, is a statement which
originated with Lacepedec, and was dictated by his
conception of the Sturgeon as distinctly a predatory
fish, using its barbels as a lure to entice its prey. The
Sturgeon, no doubt, does not disdain a fish or two,
when it can procure a meal of this kind — its larger
congener, the Hausen, is notorious for its pursuit of
small fishes, especially a variety of Bleak, in the Rus-
sian rivers — but so far as we know, it contents itself
with victims of insignificant size, such as Sand-Eels,
whose manner of life renders them a suitable prey. We
also know** that it is sometimes taken on Haddock-lines
or Cod-lines, the bait of which must thus have attrac-
tions for it. From the last-mentioned circumstance we
may also conclude that its haunts in the sea extend from
the littoral zone down to a depth of some twenty-five
fathoms; but how deep it can descend, is unknown to us.

The spawning-season of the Sturgeon occurs in
spring and early summer, from April to the end of
July. Like the Salmons, it then repairs to running
water, preferring rivers with wide mouths, deltaic
streams, or estuaries where the salinity of the water
gradually diminishes up the channel. That it spawns in
Sweden, admits of no question, though no special lo-
cality where the operation takes place is known. In
the middle of July we once received from Lulea a ripe
male, the internal organs of which are described above,
and at the same time of year we obtained a spent male
from Dalaro. In the island-belt of Soderuianland Ek-
strom secured fry 3 — 4 dm. long, “during autumn, in
the deeper watercourses”. Under ordinary circumstances
the Sturgeon does not ascend so far up the' rivers as
the Salmon. Its timidity too probably leads it to avoid
the Swedish rivers where timber is rafted. That it
should spawn at the mouths of the Swedish rivers or
even in the Gulf of Bothnia, on the shores of the island-
belt, is by no means incredible. Milner describes how
its nearly related congener, if not a member of the
same species, the fresh-water Sturgeon of . America,

" Brit. Zool. (1776), vol. Ill, p. 110.

b Henking, Deutsch. Fisch. Ver., Mitth. Sect. Kiist., Hochseefisch., 1893, p. 21.
c Hist. Nat. Poiss., (1798), tome I, p. 418.
d Krgyer, 1. c., p. 775.

1062

SCANDINAVIAN FISHES.

congregates near the shores and at the months of the
rivers, and how the dish may be seen in the evening leap-
ing from the surface, throwing their bulky forms entirely
out of the water. “At Pier Cove, Mich., on the 11th of
June, 1871, schools of sturgeons were at the edge of the
shore iu a few feet of water, and men from the vici-
nity were in the habit of wading out and drawing them
ashore with gaff-hooks. Mr. Portman, of Benton Har-
bor, has seen the sturgeon at this season lying in num-
bers on a shallow clay ledge at the edge of a stream,
several of them lying flat on their backs, with their
bellies upward, rolling and splashing in shallow water,
with apparent enjoyment” With regard to the spawn-
ing of our European Sturgeon, to the best of my know-
ledge, no personal observations have yet been recorded.

The eggs are at first yellow. As they approach
ripeness, this colour turns to black. Eggs of both co-
lours are often found interspersed in the same female.
The diameter of the ova rises to about 2 mm., and
their number in a single gravid fish runs to several
millions. They are said to be hatched six days after
deposition ; and the fry soon afterwards set out on their
journey to the sea, whence the young Sturgeons do
not return until capable of taking part in the opera-
tions of reproduction.

The Sturgeon-fishery is naturally pursued on the
most extensive scale in the countries where the fish
annually ascend the rivers (for instance the Po) in large
shoals. But the value of the Sturgeon is greatest just
before the roe begins to ripen ; and off the river-mouths
of Northern Europe important sea-fisheries are carried
on for Sturgeon in this condition, especially in the
German Haffs and on the west coast of Germany. The
tackle employed consists of coarse-meshed nets, with a
mesh 14 — 18 cm. square, and with or without sinkers.
The net is always loosely attached to the head-line,
so that in certain parts the meshes run free along
this rope, the Sturgeon thus entangling itself in the
net more and more at every struggle. Besides Stur-
geons the catch often includes seals and dolphins, Sal-
mon and other large fishes. The captive Sturgeons
are generally handled with no great tenderness. A
rope is passed in through the mouth and out at one
of the gill-openings. Another rope is made fast round
the tail. The fish is then towed at the end of these

a Of. Dean, Note on the Spawning Condition of the Sturgeon ,

1 The caviare principally imported from Russia into Sweden is
caviare of the Sterlet is scarce in the latter country.

lines beside the rowboat, and tethered, on the fisher-
man’s return, to a stake or an anchor. The Sturgeon’s
tenacity of life often exposes it to considerable suffer-
ings before it finds a purchaser.

The purposes to which the Sturgeon is applied in
domestic economy, and the value set upon it, vary
widely in different localities. The flesh, in many pla-
ces where its excellent qualities are appreciated, has
been and still is highly esteemed. There is an old
saying which declares that a skilful cook can convert
the flesh of the Sturgeon into ham, beefsteak, roast
lamb, or feathered game. The ancient Romans decked
the Sturgeon with flowers, and the dainty fish appeared
at table to the strains of music. At present the Stur-
geon is eaten less commonly fresh than smoked. The
carcase is cut up, and the strips of flesh are cured in the
said manner. Most generally known and prized, how-
ever, is the roe, which is usually sold in Sweden under
the name of Elbe caviare \ To be fit for consumption,
especially if it is to be kept any length of time, the
roe should be not quite ripe. The ovaries are opened,
and the roe is stirred in a sieve, through which it is
carefully pressed, so as to rid it of the ovarian mem-
branes and blood-vessels. This caviare requires no fur-
ther preparation but salting, though it is often com-
pressed in small bags, especially for transportation to
other markets, in which form it is, of course, drier,
but is preferred by many palates. Vjeziya (the noto-
chord), which finds a great demand in Russia, is not
prepared from the Sturgeon in Sweden.

The Sturgeon also yields — though on a far less
important scale than the Hausen — an excellent fish-glue
for industrial purposes. This gelatine properly consists
of nothing but the innermost membrane of the air-
bladder, washed and dried, in leaves which are cut to
a convenient size. In this condition, its purest form,
it is both tasteless and without smell, and may be used
not only as a glue, but also as the best isinglass for
soups and jellies, as a vehicle in the administration of
medicines, and in the fining of wines. In the usual
process of preparation, however, several other parts of
the air-bladder are not removed, and the purity of the
isinglass thus suffers. A coarser glue may be extracted
from the skin of the Sturgeon, and the offal may be
boiled down into oil.

Zool. Anzeig., XVI Jahrg., No. 436 (27 Dec, 1893), p. 473.

the coarse-grained roe of the Hansen. The finer-grained, more valuable

ELASMOBRANCHS.

10G3

PISCES ELASMOBRANCHII.

Fishes with cartilaginous endoskeleton (without dermoskeleton), with the shoulder-girdle
detached from and suspended behind the head, with the maxillary (and mandibular) bones
represented merely by loose cartilages at the sides of the palate and lower jaw, with the
branchial arches entirely or partly united to the skin (gill-slits externally open or covered
by a common dermal fold), and with diphycercal or heterocercal caudal fin. Fin-rays

primary. Nostrils as a rule ventral". Air-bladder none.

These fishes were removed from Artedi’s Clion-
dr opterygiib by Bonaparte into a separate order, most
clearly distinguished by the structure of the gills both
from the preceding orders, which have borne the com-
mon name of Tectobranchii0 , as having the branchial
cavity covered by a true opercular apparatus, belonging
to the skeleton, and from the lower fishes, the Cyclo-
stonii and Leptocardii, which are without true branchial
arches. The union between these arches and the skin
is accomplished in the Elasmobranchs by the extension
of a membrane, supported by cartilaginous rods, be-
tween the latter and each of the former. With this
membrane too the several branchial lamellae- coalesce
throughout their length. In some Elasmobranchs, in
the suborder of the Chimaeras, however, the said mem-
brane extends only in part, above and below, quite to
the skin, a common branchial cavity being thus pro-
duced on each side of the body. This cavity is fur-
nished, as in the preceding fishes, with a single aper-
ture, and the rest of it is covered by a dermal flap,
extended on cartilaginous rods, which are homologous

with the branchiostegal rays of the Teleosts, and by a.
thin disk of cartilage (tig. 294, op), representing the
operculum of the preceding fishes, but evidently form-
ed here, as in the other suborder, by the mutual
confluence of the upper (posterior) cartilaginous rods
at the upper (anterior) end. Analogous cartilaginous
rods also appear, as we have mentioned, in the mem-
brane originating from each of the branchial arches,
and supplying attachments for the branchial lamellau
In the rest of the Elasmobranchs, the suborder of the
Sharks and Rays, which is far superior in variety of
form, the last-mentioned membrane, supported by its
cartilaginous rods (figs. 298 and 300, rbr and br), is
united throughout its extent, both from the hyoid and
the branchial arches, either obliquely or in a straight
outward direction, to the skin, which opens at the in-
terstices into external gill-slits, numbering 7 — 5 in the
surviving forms of this suborder.

The scapular and pelvic arches are far more de-
veloped in the Elasmobranchs than in the preceding
fishes; and starting from the Elasmobranchs, morpho-

a An exception is made by ancient (primitive) forms, such as Chlamydoselache anguineus , a Japanese fish remarkable in many other
respects on account of its primitive characters, and described by G-akman, Bull. Mus. Comp. Zool. Harv. Coll., vol. XII, No. 1, and G-Onther,
Deep Sea Fish., Chall. Exped., p. 2, pi. LXIV, LXV.
b See above, p. 1043.

c Hasse, Beitr. allgem. Stammesgesch. Wirbelth., Jena 1883, p. 12.

Scandinavian Fishes.

134

1064

SCANDINAVIAN FISHES.

legists have succeeded in explaining the equivalent
parts of the preceding fishes, where these are more
and more simplified by reduction, as well as the cor-
responding structure of the higher vertebrates. In some
Elasmobranchs (the Rays, fig. 293) the shoulder-girdle
forms a closed, flattened ring around the anterior end
of the abdominal cavity and the spinal column behind
the last branchial arch, being suspended over the con-
tinent neural spines of the anterior part of the spinal
column. The upper part of this ring consists of the
two simple suprascapular disks, one on each side of

articular condyles (for the pectoral fin), set in an hori-
zontal row. The. scapular part lies above the coracoid;
but both of them form a continuous cartilaginous disk,
and are also of a piece with the lower median port of
the ring. The last-mentioned part has been interpreted
by Parker as an equivalent of the epieoracoid bone of
the higher vertebrates, where this bone is distinct
even in the lowest mammals. In most of the Elasmo-
branchs (the Sharks and Chimaeras, fig. 294), however,
the suprascapular cartilages ( ssc ) are parted from each
other and suspended from the membranous investment

A

0

B

Fig. 293. Skeleton and shoulder-girdle of the Rays.

A: Skeleton of a Skate ( Raja nidrosiensis ), cf1, seen from the dorsal. side. 1/10. Trondhjem Fjord, March 17, 1891. Conservator Storm.
B: Shoulder-girdle of a Thornback ( Raja clavata), seen from the ventral side. 1/2. After Parker, v, part of the spinal* column; gl, the
first of the three lateral processes furnished with articular surfaces for the pectoral fin; cr, coracoid part; e. cr, ventral side of the shoulder-

girdle (epieoracoid part).

the spinal column. The sides thereof are formed by
the much more expanded scapular and coracoid parts,
which are pierced with large foramina (for the passage
of nerves and blood-vessels) and furnished with three

(. aponeurosis ) of the dorsal muscles only by a prolonga-
tion of their own investing membrane {perichondrium),
the whole girdle too being more simple and not unlike
the pelvic girdle. The cartilage of the latter (figs. 293,

ELASMOBEANCHS.

1065

A and 295) never forms a closed arch, but only a
centrally situated half-arch on each side, the two halves
being usually confluent in the median line of the belly,
but. in the Chimaeras united only by a mobile sym-
physis. The main part of the pelvic girdle — originally
produced by the inward growth and coalescence of the
inner (proximal) parts of the primary ventral radialia -
answers to the ossa pubis and ossa iscliii of the higher
vertebrates, but in most cases there appears on each
side a process ascending from the end of the pelvic
disk, especially well developed iu the Rays, and re | we-

an d the aponeurosis of the lateral muscles, in the same
manner as the suprascapular cartilage suspends the
shoulder-girdle in these fishes and in the . Sharks. A
closer morphological connexion in respect to the struc-
ture of the shoulder-girdle and pelvis exists, as we see,
between the Elasmobranchs and the higher vertebrates
than between the former and the Teleosts.

The structure of the tins points to a similar con-
clusion. In this respect, it is true, (he Elasmobranchs
approach nearer to the common parent-forms, where
all the fins were more or less differentiated sections of

Fig. 294. Anterior part of the skeleton in a ( 'J hi nicer a monstrosa , a71. 2/3 nat. size.

b, saddle-like disk formed by the coalescence of the neurapophyses of the first abdominal vertebrae; bhy, basihyoid ; ce, anterior aperture of the
ethmoidal canal; ch , notochord; chy, ceratohyoid; cp, preorbital canal, passage of the ramus ophthalmicus superior nervi trigemini to the
ethmoidal canal; da, anterior dental plate of the upper jaw; di, dental plate of the lower jaw; dp, posterior dental plate of the upper jaw;
fa, foramen for a branch of the facial nerve; fb , fibrils of the anterior dorsal and the pectoral fins; h, rudimentary haem apophyses ; hm, hyo-
mandibular; iktd, spine (ichthyodorulite) of the anterior dorsal fin; in, interneurals; If, fronto-nasal cartilage (prehensile organ of the male):
Imi, labial cartilage of the lower jaw; Ims, labial cartilage of the upper jaw; hij, first labial cartilage of the nasal capsule; ln2, second labial
cartilage of the nasal capsule; lri , first rostro-nasal cartilage; /;%>, second rostro-nasal cartilage; mb, posterior, fibril-less membrane of the first
dorsal fin; mp, mesopterygiuvi ; mt ,, first part of the metapterygium; mt2, second part of the metapterygium; N, nasal capsule; na, neurapo-
physes; occ, occipital condyles; on, orbitonasal canal, passage of the ramus ophthalmicus profundus nervi trigemini to the ethmoidal canal;
opt, optic foramen; pp, propterygium; qpg , palatoquadrate; Ii, undivided radial (interspinal) cartilage of the first dorsal fin; rb, foramen for
the ramus buccalis nervi trigemini ; s, Solgerian (spiracular) cartilage; sc, scapular cartilage; sp, tegumentary cartilage of the spinal canal
( processus spinosi superior es) ; ssc , suprascapular cartilage; sy, symplecticum; tr, foramen for the trigeminal and facial nerves; trs, foramen
for the ramus ophthalmicus nervi trigemini: 1 — 6, first (proximal) to sixth (distal) series of the pectoral radialia.

senting an os ilii. In the Chimaeras this process runs
so far up, more in a straight line with the pelvic disk,
as to unite by means of ligaments the pelvic girdle

originally continuous dermal folds — unpaired along the
dorsal and inferior caudal margins, but double (paired)'
on the ventral side, as having been divided to surround

1066

SCANDINAVIAN FISHES.

the vent- and advance along each side of the belly".
Yet this structure has undergone great development,
retaining meanwhile the axial parts of the paired tins
to such an extent that morphological comparison may
have recourse to the Elasmobranchs in quest of starting-
points for the interpretation of the arms and legs of
the higher vertebrates. From embryology Balfour* 6
and Wiedersheim" learnt — as Teacher'* and Mivart"
had previously discovered by researches in comparative
anatomy — that the primordial parts in the skeleton of
the paired tins are the basal portions of the cartilagi-
nous rays (primary radialia), which grow inwards to
form the groundwork of the shoulder-girdle and the pel-
vis, and in their outward collocation, radiating unilater-
ally from a basal part, are split in a, transverse direction

a still surviving Lung-fish The axial parts which have
been retained in the paired tins of the living Elasmo-
branchs have been named by Gegenbaur the anterior ,
middle , and posterior pteryyial parts {pro-, meso-, and
metapterygium). In some forms, as in the Rays, the
Notidanoids, and some other Sharks, all these parts
share in the articulations of the pectoral tins, in others
only two of them or even only one, as the proptery-
gium and a part of the inetapterygium in ChimceYa
(tig. 294, pp and mtt), and the metapterygium in
Scymnus. The last-mentioned peculiarity may also be
observed in the ventral tins. In the Chondrosteans and
Teleosts we have seen how these parts gradually dis-
appear during the development of the normal piscine
type. In the ventral tins of all male Elasmobranchs,

Fig. 295. Left side of the pelvic girdle and left ventral fin of a Chimcera monstrosa, 2/s nat. size.

b , undivided basal cartilage of the ventral fin, answering to the propterygium and mesopterygium ; for , obturator foramen, at the side of
which appear two fenestrae, closed with membrane, in the pelvic cartilage; pit, iliac part; pis. ischiadic part; pp, pubic part ; prp , prsepubic
part, serrated cartilage or anterior copulatory organ of the male; ptpd, pterygopodium of the male, metapterygoid structure, consisting of:
mti, inner metapterygoid cartilage; mf and mt ,, inner (proximal) and outer (distal) metapterygoid parts, the latter divided into three bran-
ches, the outer superior ( se ), the inner superior (si) and the inner inferior (it).

and multiplied within the tin itself. That the prim-
ordial arrangement, however, here as in the cliphycercal
caudal tin, was bilateral, with rays on each side of an
axial part, appears from’ such forms as Fleur acanthus,
a primitive Shark of the Permian fauna, or Ceratodus,

on the other hand, the metapterygium is prolongated
into free appendages (tig. 295, ptpd), which may serve
as organs of motion, but are properly copulatory organs.
The male Chiinseras are furnished with another ap-
pendage, possessing the same function, a mobile, sickle-

“ Cf. Smitt: Ur de hogre djurens utvecklingshistoria, pp. 240 and 241.

6 Comparative Embryology , vol. II, pp. 492, cett.
c Das Gliedmassenskelet der Wirbelthiere , Jena 1892.
d Median and paired fins, Trans. Connect. Acad., vol. Ill (1877).
e On the Fins of Elasmobranchii , Trans. Zool. Soc. Lond., vol. X.

f Appending a reference to the structure of the paired fins in Ceratodus, I expressed in 1873 the above-cited opinion, which
Wiedersheim (1. c., p. 13) calls “Die TuACHER-MivART-BALFOUR-HASWELL-DoHRN’sche Lehre” .

ELASMOBRANCHS.

1067

shaped disk of cartilage (fig. 295, prp) on each side
of the pelvis, armed with sharp teeth at the outer
(lower) margin, and analogous in position to the prm-
pubis of the Batrachians and Reptiles.

The fin-rays of most of the Elasmobranchs are
cartilaginous, distal sections (joints) of the radialia; but
in the membrane of the outer parts of the fins — some-
times, as in the dorsal fins of the Rays, also between
the tops of the true rays — are set fibrillar or corneous
rays (figs. 294 — 297, fb), such as we have seen above
in the embryonic fins of the Teleosts or in the so-
called adipose fin. In the Sharks and Chinneras similar
rays support the greater part of the fins (outside the
radialia). Well-marked calcifications indeed appear,
under the form of fin-rays almost exactly resembling
in their exterior the spinous rays we have seen among
the preceding fishes. Such rays are set in many Elas-
mobranchs before the unpaired fins (fig. 294, iktd );
and in some Rays they arm the tail with mobile wea-
pons of offence and defence. In palaeontology they bear
the name of ichtliyodorulites. Agassiz has shown", that
both as regards their attachment, which is without true
articulation, and their texture, which exactly corre-
sponds to the dentinal structure of dermal spines (pla-
coid scales) and jaw-teeth, these spines are widely dif-
ferent from the spinous rays of the Teleosts. They are
mere dermal growths; but inasmuch as they immediate-
ly overlay and cover with their bases the anterior
margin of the supporting cartilages of the fins — e. g.
the spines of the dorsal fin in the Sharks and Chimse-
ras — they should also be interpreted as most nearly
representing skeletal tegumentary bones, although they
are without true ossification.

The skeleton of the Elasmobranchs is characterized
by this very want of endoskeletal and tegumentary bone.
Calcifications of great extent appear it is true, in old
specimens, under the form af' a thin coat outside or
just within the surface of the skeletal cartilage, and in
several Sharks the intercellular mass of the vertebral
cartilages is radially or (at the centre of the vertebra)
concentrically calcified. But there is no true ossification.
The structure of the spinal column, composed of very
numerous vertebrae, displays the most sweeping altera-
tions and the most radical divergencies. In the Chi-
maeras the notochord (fig. 294, ch) remains almost un-
altered throughout the life of the fish; but around it

slender calcific rings develop, several, five or even more,
to each vertebra, whose area may be defined as the space
between the origins of two pairs of roots belonging to
the medullary nerves. In the Sharks and Rays, on the
other hand, the notochord is more or less constricted
(fig. 296, c), or even snipped off entirely at each ver-
tebra (fig. 297, c), by the formation of the hollow
double cones, contiguous at the tops, of which the body
of the vertebra is composed. The neural arches of each
vertebra are usually broken up into two or more pairs
of triangular disks, the true neurapophyses (figs. 294,
296, and 297, na) applying their base to the body of
the vertebra, and the other disks, the so-called upper
intercalaria or intercruralia (in), wedging the apex of
their triangle in a downward direction between the
neurapophyses. The spinal canal is sometimes (fig. 294,
sp and fig. 297, partly between le) closed above by
the development of unpaired terminal parts, answering
to the upper spinous processes of more developed ver-
tebra), and where the vertical fins appear, vertical disks of
cartilage sometimes afford high supports to these fins and
their radialia — sometimes produced by the coalescence
of the spinous processes, as in the Chinneras (fig. 294,5),
sometimes corresponding to the interspinal or interneural
bones of the Teleosts. In other cases this closure is
accomplished by the intercalaria (fig. 296 and fig. 297,
partly between le). Or the covering may consist simply
of a fibrous membrane. The neural arches sometimes
(e. g. in the Rays) grow downwards along the sides of
the vertebra) or even to their ventral side; but as a
rule haemal arches (figs. 296 and 297, ha), lower inter-
calaria (ih), and lower terminal parts (spinous process-
es) are developed in a downward direction, their extent
and alterations being similar to those we have just re-
marked in the corresponding parts of the neural canal.
In the anterior part of the body, behind the head, both
the vertebra) and their apophyses often coalesce into a
continuous mass of cartilage, pierced only by the spinal
cord and the spinal nerves; and in this part ribs are
wanting or replaced by a continuous fibrous membrane,
whereas costal cartilages are present further back. Such
is the case in the Rays. In the Sharks, on the other
hand, the vertebra' are more regularly differentiated
throughout the spinal column; and ribs, though some-
times wanting, are more frequently present throughout
the abdominal region. The Holocepliali (fig. 294) show

Poiss. Foss., tom. Ill, pp. 1 and 212.

1068

SCANDINAVIAN FISHES.

a coalescence of the anterior part of the spinal column
similar to that just mentioned, and are without ribs.
The ribs of the Elasmobranchs, however, differ essen-
tially from those of the Teleosts in being more deeply
embedded in the musculature (flesh), so that they are
not immediately applied to the peritoneum, but in this
respect more nearly answer to the scleral bones of the
Teleosts and the ribs of the higher vertebrates.

The skull forms a continuous capsule of cartilage,
with no other limits between the separate parts than
those indicated by the situation of the organs of sense
and by the orifices of the cranial nerves. With their
aid we can distinguish (/) an occipital region, around

their postorbital ( ptob ) and preorbital ( prob ) processes,
(4) an ethmoidal region, around the nasal capsules ( N )
with the internasal cartilage, and lastly (5) a rostral
region, with the confluent tops of the primordial tra-
becuhe (the basitrabecular tip, B) and the supporting
cartilage (Jr) of the strongly developed rostral system
of the lateral line.

The occipital region is the shortest. In the most
primitive forms the anterior end of the notochord ex-
tends into the basioccipital part", and the foremost
neurapophyses enter into the cerebral capsule. In the
Sharks the articulation between the head and the first
vertebra is commonly very little more developed than

b

Fig. 296. First dorsal fin with its skeletal parts and the subjacent part of the spinal column in a Cyclospondyle (Hasse), a Greenland
Shark (Acanthorlnnus carckarias ), 230 cm. long, from the North Sea. J/2 of the natural size.

b, basal disk of the dorsal fin; ikt, rudimentary dorsal spine, hidden under the skin; fb, dorsal fin with its bunches of fibrils; rx and r2, first
(proximal) and second (distal) rows of radialia; cr, lateral ridge of the basal disk, a projecting crest for the attachment of muscles; Ig , ver-
tical ligament of the dorsal fin: le , elastic ligament of the spinal column; in, interneural cartilages (intercalaria neuralia ); na, neurapophyses;

c, vertebrte, superficially constricted and each containing in the space between the constrictions a division of the spinal cord; dispondylic,

each vertebra answering to two neurapophyses; h , heemal ridge, divided into haemapophyses (ha) and interhannal cartilages

(ih, intercalaria hcemalia).

the foramen magnum (figs. 298 and 299, fom), and
limited in front by the nervus vagus foramen ( mg ),

(2) a temporal or labyrinthine region, bounded in front
by the true trigeminal foramen (figs. 294 and 300, tr),

(3) a frontal (orbital) region, surrounding the orbits with

in the Teleosts; but in the Rays (fig. 300) and the
Chirmeras (fig. 294), as well as in some Sharks (fig.
298, C and fig. 299), this articulation is accomplished
by true condyles ( occ ), in the same manner as in the
higher vertebrates. In the Chimseras, these condyles,

See, for example, Chlamydoselache in Garman (Bull. Mus. Comp. Zool. Harv. Coll., vol. XII, No. 1), pi. VII, fig. B.

ELASMOBRANCHS.

1069

low but broad, medially convex and laterally concave,
are set one on each side of the foramen magnum,
and a corresponding condyle, medially concave and
laterally convex, is set on the front side of the con-
fluent vertebral mass at the beginning of the spinal
column (tig. 294, b).

The temporal region surrounds the parts of the
labyrinth, and frequently shows traces (elevations),
even on the surface, of the position within the carti-
lage of the labyrinthine canals (fig. 298, a. sc, psc, and

On the outside of the lateral cranial walls there lie in
this region the more or less marked articular cavities
of the hyomandibular cartilage (km). Through the
temporal region pass the foramina for the new us glos-
sopharyngeus (fig. 298, C, gp), n. facialis (fig. 300,
fa ), and, at the boundary between it and the orbital
region, for the n. trigeminus ( tr ).

The orbital region is furnished on each side in
the most primitive forms with a posterior articular
cavity for the palatine cartilage of the upper jaw (pa-

ih ha

Fig. 297. First dorsal fin with its skeletal parts and the subjacent part of the spinal column in an Asterospondyle (Hasse), a Porbeagle

( Isurus cornubicus), 213 cm. long, from the North Sea. ]/4 of the natural size.

ra, radialia of the anterior margin; rb, basal radialia; fb, bunches of fibrils, partly inserted between the distal radialia; rmt, metapterygoid
radialia, with their distal sections (rmt2 — rintp ; Ig , vertical ligament of the dorsal fin, investing in its lower part (Igf the elastic ligament
of the spinal column ( le ) — the last ligament is partly removed to show the roof of the spinal canal, composed partly of the interneural arches,
partly of special supraspinal cartilages; in , interneural cartilages; ncc, neurapophyses ; c, vertebrae, complete, amphiccelous, with longitudinal
lateral cavities and ribbed; monospondylous, each vertebra answering to only one neurapophysis ; ha, haemapophyses ; ih, interlnemal cartilages;
the two last-mentioned cartilages together forming on each side of the spinal column a rim projecting outwards and downwards (answering

to li in the preceding figure).

hsc ) and the vestibule. On the upper side of the
skull (tigs. 298 — 300) this region is usually depressed
into a parietal cavity, where the two endolymphatic
ducts or aqucediictas vestibuli (aqv) have their orifices.

latoquadrate cartilage) on a postorbital process, and with
an anterior articular cavity for the same cartilage at
the bottom of the orbit, on an expansion of the cranial
cartilage which has been called the basal disk, and which

1070

SCANDINAVIAN FISHES.

Fig. 298. Skull of Rough Hound ( Scylliorhinus Canicula ), seen from above (A), from the right ( B ), and from behind (6T). Natural size

After Parker.

aqv, aquceductus vestibuli; asc, external prominence of the anterior semicircular canal of the labyrinth; bbr, basibranchiale (copula) ; br, car-
tilaginous rods of the branchial membranes; cbr , 5, ceratobranchials of the first — fifth branchial arches; ch, space for the notochord, filled

with cartilage; chy, ceratohyoideum; cp , preorbital canal; ctr, cornua trabecidarum in the floor of the skull, seen through the prefrontal

fontanelle; ebrx ., epibranchials of the first — fifth branchial arches; evs, extravisceral cartilages; fom, foramen magnum; fspo , supraorbital

foramen; gp, foramen for the nervus glossopharyngeus ; librx 2, hypobranchials of the first and second branchial arches (other hypobranchials

not denoted); km , hyomandibulare ; hsc, external prominence of the horizontal (outer) semicircular canal of the labyrinth; la, anterior suspen-
sory ligament, for the palatoquadrate cartilage; hni, labial cartilage of the lower jaw; Ims , labial cartilage of the upper jaw; /«, anterior

and outer nasal labials; Ip, posterior suspensory ligament, for the palatoquadrate cartilage (upper jaw) and the mandibular arch (Meckelian

cartilage); Ir, rostral labials; Mb, Meckelian cartilage (lower jaw); N, nasal capsule; occ, occipital condyles; opt, optic foramen; phbrx 5,

pharyngobranchials of the first — fifth branchial arches; prob , preorbital process; psc, external prominence of the posterior semicircular canal
of the labyrinth; ptoli, postorbital process; ptp, suspensory process of the pterygopalatine arch; qpq, pterygopalatine cartilage; R, rostral
cartilage; r. br, brancliiostegal rays; spc, spiracular cartilage; spir, spiracle; tr, foramen for the nervus trigeminus ; vag, foramen for the

nervus vagus.

ELASMOBRANOHS.

1071

projects downwards in an angle, known as the basilar
angle". But in the more recent forms of Sharks and
in the Rays these articulations have been loosened, and
are replaced by ligaments, the postorbital process and
the basilar plate having meanwhile suffered reduction/'.
The Chi mamas have undergone an opposite development,
the palatoquadrate cartilage (tig. 294, qpg) being con-
fluent with the skull — whence the name of Holoce-
phali — and forming a, firm floor to the orbit. Within
the skull, above and just behind the basilar angle, lies
the Turkish saddle (with the infundibulum , glandula
pituitaria, and canalis caroticus ), and here the septum
between the orbits is pierced by a transverse canal, in
and around which the musculi recti of the eye have
their insertions, and which is consequently homologous
with the posterior orbito-muscular canal present in most
of the Teleosts. In the orbital region the roof of the
skull (the forehead) is sometimes (in the Rays) imper-
fectly closed, being completed by a membrane (frontal
fontanelle, fig. 300, fonf). The anterior limit of the
orbits is formed by the preorbital process (figs. 298—
300, prob), which is pierced, as well as the more or
less arched and expanded roofs of the orbits (with their
supraorbital foramina, fig. 298, fspo ), by ramifications
of the ophthalmic branch of the trigeminal nerve. The
canal (preorbital canal, figs. 294 and 298, cp) that
traverses the upper part of the preorbital process for
this purpose either opens on the top of the skull (fig.
299, cpf) and soon descends again into the cartilage
of the ethmoid region (ethmoidal canal, fig. 294, re),
or proceeds uninterruptedly, without appearing on the
surface, into the said cartilage, or leads to a deep in-
cision on each side of the skull (fig. 299, ie), the limit
between the orbital and ethmoid regions. In the lower
part of the preorbital process runs another canal, the
so-called orbitonasal canal (fig. 294, on), to receive an-
other ramification of the said ophthalmic nerve. Behind
the optic foramen there often projects within each orbit
a cartilaginous rod to support the eyeball, this rod being
the so-called eye-stalk. The tip of each preorbital pro-
cess sends out in the Notidaniclce (the most primitive in
type of the surviving Sharks) a backward process,
which in the Rays (fig. 300, pob) becomes a free car-
tilage jutting out laterally and jointed by a well-deve-

loped articulation with the preorbital process and the
nasal capsule. By Parker/ this cartilage was inter-
preted as a vestige of a. preoral (situated before the
mouth) visceral arch; but it would seem to have most
in common with the preorbital bone of the Teleosts,
especially as this appears in the Cobitidce. In the
Rays it meets the prorsal basal cartilage (propte-
rygium) of the pectoral fins, and is most highly de-
veloped in the Electric Rays ( Torpedines ) and Eagle-
Rays ( Myliobatidce ).

The ethmoid region composes the anterior limit
of the cranial cavity, being the product of the strong
expansion and development of cartilage attained by the
side-angles of the embryonic trabeculae (fig. 298, ctr ).
Medially this mass coalesces into a septum, the inter-
nasal cartilage, between the large nasal capsules (figs.
294 and 298 — 300, N), each of which is pierced with
a large foramen for the olfactory nerve. On the upper
side of the skull there appears at this point, in most
of the Elasmobranchs, the large prefrontal fontanelle
(figs. 299 and 300, fonpf ), filled with a loose, mucoid
connective tissue, the backward continuation of which
passes, in the interior of the skull, into the hard mem-
brane of the brain ( dura mater). The lateral parts of
the ethmoid region are formed by the large nasal cap-
sules (N), which are of an inverted bowl-shape, arched
above, open below, and the posterior walls of which
coalesce on each side with either preorbital process.
The structure of these capsules originates, it is true,
from the side-ends of the embryonic trabeculae; but
also includes up to three so-called labial cartilages
(figs. 294 and 298, lni and Inf), originally free, which
form the margin and marginal valves of the nostrils.

The rostral region, which attains its greatest de-
velopment in the Rays and Saw-fishes — being pro-
duced in the latter into the so-called saw with its large
lateral serrations and its two pairs of longitudinal ca-
nals, the inner pair continuations of the above-men-
tioned preorbital canals — is least developed in the
most primordial Sharks, where it forms a compressed
and narrow, or depressed and broad prolongation of
the cranial floor. In most Sharks, however, the snout
is furnished on each side with a supporting cartilage
(figs. 298 and 299, Ir), originally a so-called labial

a Cf. the Sturgeons, see above p. 1045, fig. 284, D.

b An exception to this is the Hammerhead (Zygceiia), in which the postorbital process as well enters into the laterally elongated orbits.
c Skull of Sharks, Skates, Trans. Zool. Soc. Lend., vol. X, p. 224 (ao).

Scandinavian Fishes.

135

1072

SCANDINAVIAN FISHES.

cartilage, which is directed from the tip of the rostral
cartilage (B) to the roof of the nasal capsule on the
same side. These two lateral cartilages and the rostral
cartilage thus build up a three-sided pyramidiform case,
within which is enclosed the fibrous capsule wherein
the ampullai hereafter to be described have their caecal
base. In the Holocephali — where, as in the generality
of deep-sea fishes, the rostral system of the lateral line
is particularly well developed — the soft snout and the
said capsule are supported by a pair (fig. 294, lr2) of
cartilaginous rods, widely separated at the bases but
converging towards the tips, with the liases resting on
the snout in front of the nasal capsules, and by an

The morphological homologues of the jawbones of
the Teleosts and the higher vertebrates also appear in
the Elasmobranchs under the guise of labial cartilages.
The Elasmobranchs are in general raptorial fishes of a
pronounced type, with powerful teeth in the mouth; but
these teeth are set, not on the jawbones proper, but
on the predecessors, in the development of the verteb-
rate type, of the palate and lower jaw, on the palato-
quadrate cartilage (figs. 294 and 298, qpg) and the
so-called Meckelian cartilage (MU) of each side, the
predecessors of the lower jaw, which cartilages appear
even in the embryos of the highest vertebrates as gra-
dually evanescent rudiments in the structure of the

Fig. 299. Skull of a Tope ( Galeorhinus ), seen from the left ( A ) and from above ( B ). After Gegenbaur.

cro, rudiment of occipital crest; fonpf , prefrontal fontanelle; ie, incisura etlimoidalis. The other letters are explained in the preceding figure.

unpaired, thicker and firmer cartilaginous rod (by), set
on the bridge of the snout between the two anterior
openings of the ethmoidal canal (ce), a confluence of
the anterior parts of the preorbital (cp) and orbitonasal
(on) canals. The males also possess a similar carti-
laginous rod (If) above the preorbital process, free and
mobile, transformed to a, prehensile organ set with
teeth, and probably serving to hold the female during
copulation. The history of evolution has not yet elu-
cidated the morphological significance of these carti-
laginous rods in the Holocephali , but from this point
of view they should probably be ranged, one and all,
beside the labial cartilages of the Sharks.

mandible. The relation of the palatoquadrate cartilage
to the skull we have already considered. At the pos-
terior (quadrate) extremity it articulates with the
Meckelian cartilage, which here functionates as a lower
jaw, so that the two pairs of cartilages form a ring
around the gape. But these cartilages, especially the
Meckelian, are closely united .behind by ligaments to
the hyoid arch, an opening (the spiracle, fig. 298, spit)
being left, however, between the quadrate (qpg) and
the hyomandibular (hm) cartilages, and the anterior
margin of this spiracular canal being furnished either
with a more or less developed cartilaginous disk (spc),
answering to the metapterygoid hone of the Teleosts,

ELASMOBRANCIIS.

1073

or, as in the Rays (fig. 300, nipt), by a more exten-
sive cartilage, stretching from the lower part of the
hyomandibular bone to the postorbital region. Outside
this pharyngeal and masticatory apparatus there lie as
a rule three (sometimes — as in several Sharks [fig.
298, 1ms and Imi] — only two, sometimes — as in the
Rays — none) pairs of labial cartilages, the two largest
and most constant meeting at each corner of the mouth.
Where they are all present, two of them ( Ims ) lie, one
before and below the other, on the outside of the palato-
qu ad rate cartilage, the third (Imi) outside the Meckelian
cartilage. The former two answer to the intermaxil-
laries and maxillaries, the latter is homologous with
the mandible.

i'he position occupied in relation to the said ske-
letal parts by these cartilages recurs in a number of
cartilaginous rods (tig. 298, evs ), first described by
Rathke", which lie in the skin between the gill-open-
ings, forming the outer support of the above-mentioned
cartilaginous rods in the branchiostegal membranes (fig.
298, br). GegenbauiC called the first-mentioned rods
outer gill-arches, and considered them, as Cuvier had
partly done0, to lie vestiges of the more complex
branchial arches of the Marsipobranchs. Parker'* called
them extraviscerals or extr abranchials, and saw in them
homologues of the shoulder-girdle (the scapulae and
coracoid bones), as being extracostal growths. Where
their development is highest, as in the majority of the
Sharks, they consist of two parts, a dorsal and a ventral,
which meet and are applied to each other throughout
a greater or less portion of their extent. In several
Sharks and in the Rays they either are rudimentary, or
have only the ventral parts developed (tig. 298), or are
entirely absent. The angular, hooked or forked ventral
ends of these “outer gill-arches” are joined below, from
one side of the body to the other, by an horizontally
set, fibrous membrane; and in this manner is formed,
under the hypobranchial (lowest, tigs. 298 and 300, hbr)
and copular (bbr) parts of the true branchial arches, a
special chamber, a prolongation of the pericardial cavity,
to receive the heart with the conus arteriosus and,
within an anterior division, the truncus arteriosus.

The hyoid arch shows an entirely different com-
position and suspension in the Sharks and Chiinaeras
on the one hand and in the Rays on the other. In
the former the true middle part ( ceratohyoic/eum , tigs.
294 and 298, cliy) is suspended from the lower ex-
tremity of the hyomandibular cartilage (hm), and these
parts are united below, from one side of the body to
the other, by the first copula ( basihyoideum , bhy). In
the Rays the hyoid arch consists on each side, as a
rule, of 3 or 4 parts, the lowest ( hypohyoideum , fig.
300, hby), as well as the lowest part of the first branchial
arch ( hyp abranchiate ), meeting the first copula ( bbi\ ).
Among the remaining parts of the hyoid arch the
ceratohyoid (chy) and epihyoid (ehy) are the most con-
stant; the uppermost part ( sthy ), answering to the sty-
lohyoid of the Teleosts, is either cartilaginous or re-
presented by a ligament, and suspends the arch from
the upper posterior angle of the hyomandibular cartilage.
The true branchial arches consist as a rule of four
parts on each side — the hypobranchial (hbr), the cera-
tobranchial (cbr), the epibranchial (ebr), and the pha-
ryngobranchial (phbr). But between the lowest parts
(the hypobranchials) there appear in the Sharks, though
their occurrence is irregular, unpaired pieces (copular
parts), sometimes five in number. Only the hindmost
and largest (bbr), under which the heart is situated,
is constant, and this is always present in the Rays
as well, where it attains a still greater development.

Gill-rakers are usually wanting in the Elasmo-
branchs; but two remarkable exceptions are formed by
the largest Sharks known, the North Atlantic Basking
Shark and the Whale-Shark (Rhinodon typicus) of the
Cape of Good Hope and the Indian Ocean east of Africa.
Like the whales, these Sharks live on small animals, and
like the Teleosts that feed in the same way, they are
furnished with a filtering apparatus (“gill-grating”,
Steenstrup, see tig. 332, p. 1145), which separates the
food from the water that pours from the mouth at
the same time and finds an outlet through the gill-
slits. This filtering apparatus is composed in the Bask-
ing Shark0 of tine, setiform, and dense gill-rakers, up
to 6 inches long, the microscopical structure of which

a Anat. Phil. Unters. Kiemenapp., Zungenb. Wirbelth. (1832), p. 83.
b Unters. Vergl. Anat. Wirbelth., 3:tes Heft, p. 164.
c Lemons, ed. 2, tom. VII, p. 307.
d Trans. Zoo], Soc. Loud., vol. X, p. 212.

e Gunnerus, Trondhj. Selsk. Skr., part. Ill (1765), p. 46; Foulis, Proc. Best. Soc. Nat. Hist. 1854, p. 203; Steenstr., Overs. D.
Vid Selsk. Forh. 1873, No. 1, p. 47, tab. II.

1074

SCANDINAVIAN FISHES.

Avas shown by Hannover to be the same as that of
(the teeth and) the dermal spines.

Among the peculiarities of the internal organs in
the Elasmobranchs we have already considered the
structure and position of the heart. A further charac-

vity; and as the latter, being pierced by the abdominal
(anal) pores, one on each side of the anal aperture, is
in a certain degree open to the surrounding water, this
element may find a passage to the heart itself®. The
intestinal canal is comparatively simple (tig. 301). The

phh\

tv mpt opt pob

Fig. 300. Skull of a Thornbaek ( Raja clavatd) seen from above (A) and from the right (B). Natural size. After Parker.
ehy , epihyoid; fa , foramen for the nervus facialis; fonf , frontal fontanelle; fonpf, prefrontal fontanelle; hhy, hypohyoid; nipt , metapterygoid;

pob , preorbital; sthy, stylohyoid. The other letters are explained in fig. 298.

teristic of these tishes is the communication afforded
to the pericardial cavity by means of ttvo holes behind,
in the diaphragm, Avith the abdominal (peritoneal) ca-

Avide, but sometimes very short oesophagus ( d ) passes
into a stomach ( e ) usually of greater length, and this
terminates behind in a more or less prolongated blind

a Monro, Struct., Physiol. Fish., Edinburgh, 1785, p, 20, pi. II, 22, 23, pi. XVIII, 10, 11, 12. Blainville, Ann. Mus. Hist. Nat.,
tome XVIII (1811), p. 111.

ELASMOBRANCHS.

1075

sac, but bends forward in a pyloric part (to /). From
the pylorus the duodenum runs straight back, and
passes into the spiral intestine (Ji), which is continued
by the short rectum. The mucous membrane of the
intestinal canal differs in the various sections much as
in the case of the Chondrosteans. In the oesophagus
it forms longitudinal or, in addition, transverse folds,
or is reticulated; or it may be studded with more or
less hard papillae, pointed excrescences, in considerable
number. The mucous membrane of the stomach is
usually coursed by strongly developed (but few) longi-
tudinal folds. The pyloric part, where there are also
longitudinal folds of the mucous membrane, is some-
times without internal limits, but sometimes bounded,
both at the beginning and end, by an annular valve".
As we have remarked above (p. 1052, note 5), the
spiral intestine is usually regarded at the present day
as a section of the duodenum. The spiral fold in its
interior is very highly developed in the generality of
the Elasmobranchs* * 6, with reticulated mucous membrane
and with transverse ridges on most of the coils. The
rectum is short, and its mucous membrane smooth.

The liver (fig. 301, c) is large and oily, usually
consisting in the Sharks of two lobes, in the Rays of
three. The gall-bladder lies free, or is embedded in the
liver. The spleen ( x ) lies at the posterior part of the
stomach. To the right thereof (vertically below / in
the figure) lies the pancreas, which is usually of less
volume. Into the beginning of the rectum there opens
a digitiform gland (i), which was first remarked by
Monro0 under the name of vermiform appendage or
coecal sac'7, but whose glandular, botryoid structure,
around a central efferent duct, was first clearly de-
scribed by Leydig0.

There is no air-bladder. The kidneys (fig. 301, m)
are situated as in the Teleosts. They taper in front
and expand behind, sometimes so greatly that they
seem to be confluent. Their general shape adapts itself

to the external form of the body. Thus they are
elongated in the Sharks, broader and shorter in the
Rays. The urinary bladder is usually double, forming
a dilatation of each ureter. The anterior part of each

Fig. 301. Abdominal viscera of a male Shark. After Rymer- Jones.

a, heart; b, gill-openings; c, c, c, lobes of the liver, the left removed,
the middle ( lobulus Spiegelii ) and right retained; cl , oesophagus, pass-
ing into the stomach ( e ), which is continued by the duodenum (/),
into which the gall-duct (g) opens; h, spiral intestine; i, glandnla
retroanalis ; 1c, urogenital cloaca; l , pterygopodium ; rn, left kidney;
n, left testis; o, vas deferens; p, dilatation of the vas deferens ( vest -
aula seminalis ); r , urogenital papilla; s, abdominal (peritoneal) pores;

x, spleen.

“ In the Basking Shark Blainville (1. c., p. 100, pi. 6, fig. 2, D) hence gave to this section the name of the third stomach. In

the Picked Dog-fish I find no valve, either between the stomach and the pyloric part or between the latter and the duodenum.

6 In a Picked Dog-fish I find 14 coils; Kroyer found 16 or 17. The last coil terminates in a broad, labiated fold. In several
Sharks, however, the spiral fold is less developed. In Alopias vulpes and Carcharias glaucus it is described as very short. In Thalassorhi-
nus vulpecula and Zygoma tudes no stair-like coils are formed along the inside of the intestine, but a long and broad membrane curves in
cornet-shaped folds, one within the other, and lies like a packet within the intestine, with only one margin attached to the wall thereof. At
the free margin of this membrane runs, according to Duvernoy (Ann. Sc. Nat., Zool., 2 ser., tome III, p. 274, pi. 10 and 11), a power-
fully muscular venal trunk ( vena mesenterica ), which is continued by the vena porta and impels the blood and chyle into the latter.

c Struct. Physiol. Fish., Edinb. 1785, PL II, 8; PI. III. E; PI. XI, D; PL XVIII, 16; PL XIX, 14.

d Bursa cloacce , Retz., Ohs. Anat. Chondropt., p. 24. Glandnla retroanalis, Costa, Fna Regn. Nap., Chimeridei (1852), p. 36,

tav. II, fig. 1 , g.

e Beitr. mikr. Anat., Entwickel. Haien unci Rochen (1852), p. 56, § 38.

1076

SCANDINAVIAN FISHES.

kidney (the Wolffian body) is separated in the males
from the posterior part, and is applied, as an epidi-
dymis, to the anterior part of each testis, receiving
the efferent duct thereof.

In the structure of the generative organs the Elas-
mobranchs are even more closely approximated than
the Chondrosteans to the higher vertebrates. These
organs are generally more restricted in extent, whence
it naturally results that the offspring is far less nu-
merous than that of the Teleosts. The efferent duct
(vas deferens, tig. 301, o) of the male organs (n) passes,
as iu the higher vertebrates, first through an epididy-
mis. Before opening into the cloaca, it widens into a
dilatation (seminal vesicle, p), the posterior extremity
of which is applied to the corresponding part of the
seminal vesicle on the other side of the body, and then
finds outlet, in common with the other seminal vesicle

their slimy secretion. A large gland of this kind is
situated on the under side of the ventral fins. In most
cases the said organs are besides furnished at the tip
with cards of teeth or with spines.

The copulation between the sexes that is practised
with the aid of these organs by the Sharks has been
observed in the case of the Nurse Hound ( Scylliorliinus
stellaris ) by Bolau at the Hamburg Aquarium, and is
illustrated by the appended figure (302). Of the copu-
lation of the Rays Hollberg writes* 6: “During the
season of propagation they repair to the surface of the
sea and, for the purpose of depositing their eggs, to
the shores. At these times they are taken with the
harpoon. The female is frequently attended by several
males, who endeavour by lifting and tossing her with
their snouts to turn her over on her back. The first
to succeed in this applies and presses his ventral side

Fig. 302. Male Nurse Hound clasping the female during copulation. After Bolau, Zeitschr. f. wiss. Zoo]., Bd. XXXV, p. 322.

and the ureters, into a special cloaca ( k , the urogenital
cloaca, Moreau, Hist. Nat. Poiss. Fr., tome I, fig. 26,
p. 246), which projects on the wall of the anal cloaca
in the form of a papilla (r) only requiring the erectile
tissue to represent a true penis. In the Chimaeras the
urogenital aperture of the males lies externally on the
ventral side, at the anterior extremity of an oblong
and compressed, but posteriorly tumid protuberance
behind the vent.

The external copulatory organs of the males we
have mentioned above. The most important, the so-
called pterygopodia of Petri", consisting of the trans-
formed hindmost radialia of the ventral fins ( metapte-
rygium in Gegenbaur; fig. 301, ?; fig. 295, ptpd), are
composed of several parts and furnished with a longi-
tudinal channel, into which several glands discharge

so closely to hers that, if only one of them be har-
pooned, both may be drawn out of the water without-
being parted.” On the occasion described by Bolau
the right pterygopodium of the male was inserted into
the sexual aperture of the female. The male flung his
tail upwards and forwards along the right side of her
body and bent his own in the opposite direction, though
with the dorsal side under, so that the top of his head
pressed his tail to the back of the female above her
pectoral fins. In the last-mentioned circumstance we
may perhaps find a clue to an explanation of the man-
ner in which the male Chimaeras employ the mobile
prehensory organ situated on their forehead. The co-
pulation of the Chimaeras has never been observed; but
as the mouths of the two oviducts, which open in the
females of these fishes beside each other and externally,

a Zeitschr. wiss. Zool., Bd. XXX, p. 296.

6 Gbgs Wett., Witt. Samli. N. Handl., Ill Delen (1819), p. 16.

ELASMOBRANCIIS.

1077

in the skin of the ventral side, between the anus and
the urethral aperture, are equally tumid during the
breeding-season, it seems highly probable that both the
pterygopodia functionate simultaneously. The situation
of the urogenital opening in the males, relatively to
the base of the pterygopodia, is such that the semen
may apparently be received almost immediately in the
above-mentioned channel on the pterygopodia.

The ovaries (fig. 303, n) occupy the same position
as the testes — in the anterior part of the abdominal
cavity they are attached by means of their mesoarium
(peritoneal fold) to the spinal column — but are sepa-
rated from their efferent (the Mullerian) ducts, which
unite before them, under the oesophagus and just be-
hind the diaphragm, in a curve, at the middle of which
they have their common opening (</). To this opening
the eggs thrown off by the ovaries are conducted by
a, current produced by the ciliary motion of the investing
cells of the serous tissue lining the interjacent organs.
The said current may be observed even shortly after
the death of the fish by opening the belly, removing
or lifting aside the liver, and strewing finely powdered
charcoal on the parts. The oviducts (Mullerian ducts)
bend backwards, one on each side, and are each fur-
nished with two dilatations. The anterior (a) of these
is glandular, its walls being traversed by ramified ducts
that secrete a nidamental substance to envelop the eggs;
while the posterior (r) is a uterine pouch within which
the impregnated ova either undergo the earliest stages
of their development, or remain until the young are
fully developed and capable of free motion. In the
latter case, when only the right ovary as a rule is
functional, the glandular dilatation of the oviducts is
little developed, but the uterus all the more so, its
walls being lined with vascular folds or even with a
placenta uterina, into which the vitelline sac of the
embryo is fitted by means of warty placental growths
(cotyledons), an arrangement that would be fully ana-
logous to the viviparous development of the mammals,
if the yolk sac were replaced by the foetal membranes
of the latter. The oviducts open either into the com-
mon cloaca (.s, .s), on each side of the urethral aper-

ture ( t ), or, as mentioned above, at special orifices in
the skin.

The primeval type of the Elasmobranchs has rami-
fied in two essentially different directions of evolution,
on the one hand to the Holocephali, on the other to
the Plagiostomi.

Fig. 303. Female organs of a Shark. After Owen.

a, ventral wall, cut open and laid back; b , sections of the shoulder-
girdle and pelvis; c, heart in its pericardium; h, hindmost part of
the rectum and the ascending glandula retroanalis; n, right ovary;
o, nidamental dilatation of the left oviduct ( glandula nidamental is);
q , common aperture of the two oviducts; at its bottom the orifice of
the right oviduct (to the left in the figure); r, uterine division of
the left oviduct (the right oviduct lies unopened to the left of the
figure); s, posterior mouths of the two oviducts; t, urethral papilla,
at the tip of which a bristle is inserted into the urethra. On each
side of Ibis bristle may be seen an abdominal pore (the mouth of a
peritoneal canal).

1078

S C AND I N A V I A N FIS II ES .

EL ASMOBRAN CHII HOLOCEPHALI.

Elasmobranchs with one gill-opening' common to all the gill-slits on each side of the body. No spiracles. Skin
smooth, without placoid scales (spines''). Notochord unaltered, not constricted by the formation of vertebrae,
only with superficial rings of cartilage, considerably exceeding in number the vertebral spaces (polyspondyli).
Palatoquadrate part not divided from the skull, but furnished like the lower jaw with dental disks.

That course of development from the primordial
fishes' which is represented in the present age by the
Holocephali , the suborder of the Chiimeras, has been
arrested as regards the structure of the spinal column
at a lower stage than the Plagiostoms. It is probable
too that the structure of the palate in the Holocephali
is a relic of the said ancestral types; but only the
history of evolution can decide whether the union be-
tween the palatoquadrate parts and the skull, a pecu-
liarity which we shall again meet with in the still
more primitive Marsipobranchs, is here of a primary
or a secondary nature. Equally indispensable is the aid
of the history of evolution to a determination whether
the absence of spiracles is original or rather the con-
sequence of reduction. Solgek has described in Chi-
mcera monstrosa small cartilages (tig. 294, s) which
occupy in relation to the lower jaw the same position
as that assumed in relation to the palatoquadrate car-
tilage by the spiracular cartilages (supports of the spi-
racular gills) common in the Plagiostoms. It may
reasonably be assumed that these so-called Solgerian

cartilages admit of interpretation as vestiges of aborted
spiracular gills. On the other hand, the structure of
the branchial cavity in the Holocephali more clearly
indicates a higher degree of differentiation, a step in
the direction of the Chondrosteans and Teleosts.

The Holocephali are marine and deep-sea fishes
with a wide geographical range. How great is their
geological age, can hardly be stated at present with
certainty, for ichthyologists have supposed** that remains
of these fishes have been discovered even in Devonian
deposits. It is certain, however, that they lived in the
Jurassic period, when its very earliest stratum (the
Lias) was formed; and during the latter part of this
period they attained a size far surpassing that of mo-
dern Holocephali. Townsend, for instance, found in
the Portland chalk at Great Milton (near Oxford) an
under tooth of Chimcera ( Ischyodon ) Townsendii e that
measured 1 1 cm. at the symphyseal margin, a dimen-
sion which in a Chimcera monstrosa 75 cm. long is
represented by 14 mm.

The suborder contains only one family.

Fa m. CHIMiERXDiE.

Body of a compressed clavate form , most nearly resembling that of the Macruroids. Dentition of the mouth made
up of two upper pairs and one under pair of dental plates , most similar to those of the Lung-fishes {Dipnoi).
Twof dorsal fins , the anterior , above the pectoral fins , triangular and armed with spines , the posterior low and
long; a small , sometimes scarcely distinguishable anal fin, situated far back; caudal fin diphy cereal or heterocefical .

Of this family only four* *7 species survive at the pre- which, the Antarctic Callorhynchus, contains a solitary
sent time. These have been ranged in two genera, one of species and is characterised by the more obliquely for-

0 blog, tvhole, and VM'fiaifi, head.

b In the young of Callorhynchus (a genus from the Pacific Ocean and the Antarctic regions) spines have been found in two longitu-
dinal rows, partly on the forehead, partly on the back, between the first and second dorsal fins (see Dum., Hist. Nat. Poiss., Suites a Buffon,
tome I, p. G94, pi. 14, fig. 4), an indication that the skin of the primitive forms was clothed with spines.
c Pisces aspondyli, PIasse, Nattirl. Hyst. Elasmobr., Allgem. Tbeil, p. 31.
d Of. Gunther, Introd. Study Fish., p. 349.

e Agassiz, Rech. Poiss. Foss., tome III, p. 343, tab. 40, fig. 20.

•f Sometimes the posterior dorsal fin is so deeply sinuous that three dorsal fins have been counted.

9 Gill has besides described a Chimcera phnnbea from the Atlantic coast of North America; but there would hardly seem to be any
specific distinction between this form and our Chiinsera.

HOLOCEPHALI.

1079

ward direction of the upper front teeth, by a special upward direction of the tip of the tail, the upper part
dermal appendage at the tip of the snout, and by the 1 of the caudal fin being extremely small (heterocercal).

Genus CHIMERA.

Front teeth in the upper jaiv of a vertical or hut slightly oblique forward direction; tip of the snout without
special appendage; tip of the tail prolongated straight hack, in a line with the spinal column , the caudal fin
narrowing behind and being of about the same height and extent above as below.

In the Iliad (VI, 179) Homer depicts the slaying
by Bellerophon of the Xi'/uatget, a, fire-breathing mon-
ster shaped in front as a lion, behind as a dragon, in
the middle as a goat. In the seventeenth century
Kentman sent to Gesner a drawing of a fish from
Denmark which he called Meeraff {Simia marina)', and
when Linnaeus found the same kind of fish from Bo-
huslan in the Royal Museum at Ulriksdal, he called it
Monkey-fish or Monster-fish ( Chimcera ). “Monster is
the name given to this fish, which is so unlike all
others and as it were a medley of all fishes. At certain
seasons when this strange fish dieth and is cast ashore,
the common people behold in him a miracle and ima-
gine that they see laces, points, topknots, and other

finery, which they believe have so displeased the great
God that He hath seen good to warn them with signs
and wonders, which belief news-men, no less wise than
they, speed the whole world through to the edification
of all”". Thus the name of Chimcera was introduced
into ichthyology; and though the monstrosity formerly
seen in the structure of the genus has disappeared in
the light of modern researches into its relations to the
other Elasmobranchs, the life led by these fishes is sfill
in great part a mystery to us.

Three species are recognised within the genus,
one from the west coast of North America, one from
the North Atlantic and Japan, one from the coast of
Portugal.

THE NORTHERN CHIMERA (sw. hafmusen).
CHIMiERA MONSTROSA.

Plate XL VI, figs. 2 and 3.

All the vertical fins contiguous or nearly so; the anal lobe distinct just in front of the beginning of the lower caudal
lobe and below the end of the second dorsal fin. Tip of the tail prolongated into a finless, whip-like appendage.
The tips of the pectoral fins, when laid back, extend to the anterior margin of the ventral fins or still further.
Pterygopodia ( posterior appendages of the ventral fins) of the males forked throughout the greater part of their length.

Syn. Meeraff ( Simia marina ) sec. Kentmann in litt. : Gesn., De
Aquatilibus , p. 878 (fig. in p. 877). Galens Acanthias
Clusii Exoticus, Willughby (ed. Ray), p. 57, tab. B, 9,
fig. 6. Iis-Galte , Str6m, Sondmors Beslcriv., Tom. I, p. 289.
Chimcera monstrosa, Lin., Mus. Ad. Frid., part. I, p. 53,
tab. XXV; Syst. Nat., ed. X, vol. I, p. 236; Fna Suec.,
ed. II, p. 197; Gunn. ( Hav-Katten ), Trondhj. Selsk. Skr.,
vol. II, p. 270, tabb. V et VI; Olafs. ( Iiavinus ), Reise 1st.,
pp. 360 et 598; Ascan. (Solvhaaen, argentea ), Icon. Rer.
Nat., fasc. II, p. 6, tab. XV ; Mull., Prodr. Zool. Dan.,
p. 38; Bl., Naturg. Ausl. Fiscli., part. I, p. 60, tab. CXXIV;
Lacep. ( Chim . arctique), Hist. Nat. Poiss., tom. I, 392, tab.
19, fig. 1; Retz., Fna Suec. Lin., p. 308; Donov., Brit.
Fish., tab. CXI; Rtsso, Tclithyol. Nice, p. 53; Fmng, Brit.
Anim., p. 172; Faber, Fiscli. 1st., p. 41 (+ Chim. cristata,

“ Mus. Ad. Frid., part. I, p. 54.

p. 45); Nilss., Prodr. Ichthyol. Scand., p. 112; Val. in
Gaim., Voy. 1st., Groenl., tab. 20; Bonap., Iconogr. Fna Ital.,
Pesci, tab. 130; Schleg. in Sieb., Fna Japan., p. 300, tab.
CXXXII; Ekstr., Gbgs Vet., Vitt. Samh. Handl., Ny Tidsf.,
vol. I (1850), p, 21; Costa, Fna Regn. Nap., Chim., tabb.
1 — 7; Kb., Danin. Fisk., vol. Ill, p. 784; Nilss., Sleand.
Fna, Fisk., p. 705; Dum., Hist. Nat. Poiss. (su. a Buff.),
tome I, p. 686; Gthr, Brit. Mus. Cat., Fish., vol. VIII,
p. 349; Coll., Fork. Vid. Selsk. Chrnia 1874, Tillaegsh.,
p. 206; Cederstr., Ofvers. Vet. Akad. Fork. 1876, No. 4,
p. 67; Malm, Gbgs, Boh. Fna, p. 605; Winth., Naturk.
Tidskr. Kblivn, ser. 3. vol. XII, p. 56; Mor., Hist. Nat.
Poiss. Fr., tome I, p. 455; Mela, Vert. Fenn ., p. 363, tab.
X; Day, Fish. Gt. Brit., Brel., vol. II, p. 286, tab. CLI;
Lillj., Sv., Norg. Fna, Fisk., vol. Ill, p. 511.

136

Scandinavian Fishes.

1080

SCANDINAVIAN FISHES.

Chimcera borealis , Shaw, Gener. Zool., vol. V, part. I, p. 365,
tab. 157.

Chimcera arctica, Hollb., Bohusl. Fisk., fasc. II, p. 1 cum
tab. (Gbgs Wett., Witt. Samh. N. Handl., part. IV).

Chimcera mecliterranea, Risso, Eur. Me'r., tom. Ill, p. 168.

(?) Chimcera plumb ea, Gill, Proc. Philos. Soc. Washington, Dec. 22,
1877; vide Jobd., Gilb., Bull. U. S. Nat. Mus., No. 16, p. 54.

The Northern Chimeera attains a length of about
a metre". The greater part of this length — 2/3 or
more — is occupied by the prolongated tail6, which is
thread-like at the tip. The greatest depth, just behind
the pectoral tins, measures 1/s — 1f1 of the length; and
the greatest breadth (thickness), just in front of the said
point, is about 1/2 — 3/5 of the greatest depth. Where
the first dorsal fin is situated, the dorsal line is straight;
from this part it slopes at an angle of about 45° to-
wards the snout, but very slowly towards the tip of
the tail. The inferior profile of the head ascends from
behind about as much as the upper profile descends or,
when the mouth is open, a little more; but the belly
is usually rounded and tumid, the inferior caudal profile
(behind the ventral fins) thus ascending in a very gra-
dual curve. The body is, however, so loose and slippery
that no constancy in these respects can be expected.

The length of the head, which is conical and be-
hind laterally compressed, decreases as usual during
growth, varying between 15 and 11 % of that of the
body in specimens 7 — 9 dm. long. Its most charac-
teristic features, the position of the mouth and nostrils,
as well as the structure of the soft and somewhat trans-
lucent. snout, have already been described. The longi-
tudinal diameter of the large and ovally rounded eyes
in the specimens just mentioned measures about 1/3
(37 — 31 %), and their vertical diameter about 1jr> (22
— 18 %), of the length of the head. They are set high,
their superior margin lying but slightly below the frontal
profile, but somewhat obliquely, sloping in a forward
direction; and whereas the length of the snout in front
of them measures about l/2 (48 — 51 %) of the length
of the head, the postorbital length of the head is not
much more than 1/3 (35 — 38 %) of its entire length.
The elongated form is not restricted to the orbits, but

is shared by the pupils. At the anterior margin of
the eyes the forehead of the females is somewhat tumid,
and in the males the corresponding prominence is me-
dially concave; but behind the concavity articulates the
singular hooked organ (fig. 294, If) whose probable
function as a prehensile organ during copulation we
have already pointed out. It is a curved rod of
hard cartilage, the base of which is widened and pro-
longated, gliding by means of a longitudinal articulary
groove on the median edge of the forehead, and the
top of which is furnished on the under surface with
a card of 40 — 50 pointed, recurved teeth. As the
form of the articulary surface shows, the hook is only
slightly erectile; but when it is depressed, the card
of teeth works against the anterior part of the said
prominence.

On the external surface of the snout the Northern
Chirmera acquires one of its most characteristic singu-
larities, consisting in the ramifications of the lateral
line. The lateral line itself extends, as usual, along
the sides of the body, in front rather high, behind
nearer to the middle of the sides, until it descends just
behind the beginning of the lower caudal lobe to follow
•the base thereof. It is without true pores, but instead
has a fissure-like opening throughout its length, this
being due to the structure of its wall, as first described
by Leydig0. Instead of piercing through scales, which are
wanting in these fishes, the wall of the lateral line is sup-
ported by half-rings'* set beside each other and branched
at the tops. In this form the lateral line advances on
the head, where it runs upwards and inwards6 on the
occiput, and just behind the mouths of the aquceductus
vestibuli sends out a transverse canal /, a line of commu-
nication between the two sides of the body, with a
backward offshoot in the middle. Forward from this
transverse canal a supraorbital branch5' runs on each
side of the head, above the eye and laterally along the
bridge of the snout, to the side of the tip of the snout,
where it bends downwards and backwards, though only
to meet and join in a curve on the under surface of the
snout the supraorbital branch of the opposite side. A

a Day and Lilljebobg give 4 feet (1,200 mm.). Our largest specimens are females about 95 cm. long. The true length of the

body is in many cases doubtful, for it is difficult to see whether the tip of the tail is entire or broken.

6 Our specimens corroborate Lilljebobg’s statement that the males have the comparatively longest tail; but Iyroyer’s measurements

show the reverse, and rather indicate that young Chimasras have a comparatively longer tail than old.
c Arch. Anat., Physiol. 1851, p. 251, taf. X, figs. 2 and 9.
d Of an osseous structure, according to Leydig.

e This ascending portion is called by Garman (Bull. Mus. Comp. Zool. Plarv. Coll., Cambr., vol. XVII, No. 2) the occipital canal,
f Aural canal, according to Garman.
g Cranial canal and rostral canal , according to Garman.

NORTHERN CHIMvERA.

1081

more immediate continuation of the lateral line proper
consists of a suborbital branch", running obliquely
downwards and forwards along the posterior margin of
each orbit and thence obliquely upwards and forwards
along the suborbital margin, to a point about twice as
far from the tip of the snout as from the eye, where it
forms an ^-shaped curve, first downwards and back-
wards6, then downwards and forwards, afterwards join-
ing the supraorbital branch, just before the junction of
the latter with the corresponding canal on the other side
of the body. Where the suborbital branch first alters
its course, on the cheek, below the lower posterior cor-
ner of the orbit, it sends out a branch downwards and
backwards, an opercular canal c, which becomes narrower
and narrower until it disappears externally, being con-
tinued, however, by a row of narrow, transversely set,
sharp-edged pores, right across the branchiostegal mem-
brane. Just before the origin of the opercular branch
from the suborbital, the latter sends out another branch
in a downward direction, a malar canal'6, which soon
forks into a maxillary6 and a mandibular-' canal. The
former runs along the cheek, below the middle thereof,
to a line with the anterior margin of the nostril, where
it divides into two branches, an upper anterior (maxillo-
rostraF) and a lower posterior (maxillo-nasal6). Each
of the last-mentioned canals crosses the under surface of
the snout to meet and join the corresponding canal on
the opposite side of the body; but the maxillo-rostral
branch also sends out, upwards and forwards, in the
median line of this under surface, an unpaired canal1,
which unites it to the above-mentioned junction between
the supraorbital branches. The mandibular branch cross-
es over behind the lower jaw in the same manner as
the opercular branch across the isthmus. All these ra-
mifications- of the canals belonging to the system of the
lateral line mark the head and in particular the snout
with an extremely singular design, all the more striking
as their anterior parts, on the cheeks and snout, become

coarser, with more distinct fissure, and at certain points
are widened and deepened, thus acquiring a moniliform
appearance. Between the dilatations the inside of the
canal is pierced by the inward ramifications of the caecal
ducts described by Costa'. In addition to these canals
appertaining to the true system of the lateral line the
snout is furnished with the numerous muciferous sacs
(ampullae) described by LeydigC From the many pores
that partly follow, in single rows, the above-mentioned
canals and the sharp fold which the skin forms in a
curve just before the nostrils and above the lateral por-
tion of the upper lip, partly are scattered, in groups or
more isolated positions, between the said canals, these
long ampullae take their origin, in the form of ducts
directed upwards and forwards (as they approach the
very tip of the snout, backwards). The posterior am-
pullae'terminate carnally under the skin, between it and
the large fibrous capsule, filed with a gelatinous mass,
that occupies the interior of the snout, resting on the
cartilaginous rods which we have observed above in the
rostral region of the Holocephali. The anterior ampulhe
on the under surface of the snout, as well as the above-
mentioned ducts issuing from the anterior parts of the
branches of the lateral line, penetrate within the said
capsule. In the gelatinous mass are also embedded the* *
large and numerous ramifications of the fifth pair of
cranial nerves', which innervate the ampullae. The
skin itself is marked on the back of the snout with
a network (thimble-like pattern) of pits, and within
each of these may be seen in miniature the same
rotiform figure as we remarked above in the dermal
covering that invests the muciferous sacs of the Stur-
geons.

The mouth is comparatively small. It presents, in
combination with the large preoral nostrils, an appearance
seldom exemplified in the preceding fishes. The dermal
fold which bounds in front both the mouth and the
nostrils has a, great resemblance, it is true, to the lower

° Orbital and suborbital canals , according to Garman.

b In Plate XL VI, fig. 3 of the present work there is an error, this branch being connected with the supraorbital canal.
c Jugular canal , according to Garman.

d Sometimes the opercular and malar branches issue in a common canal ( orbitonasal , according to Garman) from the suborbital branch.
e Angular canal , according to Garman.

f Oral canal , ,, ,, ,,

3 Subrostral canal, ,, ,, ,,

h Nasal canal, „ ,, ,,

* Median canal, ,, ,, ,,

3 Fna Regn. Nap., Pesci, Chimceroidei, tav. VII, figs. 1—4.
k Arch. Anat., Physiol. 1851, p. 253, taf. X, fig. 1.

1 According to Ewart, to whose investigations we shall return below, the nerves which innervate both the ampullae and the other parts
of the cephalic system of the lateral line are homologous with an anterior division of the nervus facialis of the higher vertebrates.

1082

SCANDINAVIAN FISHES.

suborbital (preorbital) margin of the Teleosts; but it can-
not be fully homologous therewith, for here the nostrils
are situated below it. The nostrils, which lie just in front
of the median upper lip, are externally — each being-
bounded laterally by the lateral upper lip of that side —
simple, round, and rather large apertures, separated by
a thin septum. On raising the lateral upper lip we find,
however, that each nasal cavity besides possesses a pos-
terior opening, a dermal fold, bent into several curves
and internally supported by a cartilaginous disk (fig.
294, ln2), being turned inwards from the outer margin
of the nasal cavity and forming the greater part of the
posterior limit of the anterior nostril. To meet the inner
edge of this dermal fold — though without being con-
tiguous with it — a longer fold rises from the inner
margin of the nasal cavity, a true narial passage being
thus present, though not fully closed behind3. Exter-
nally the nasal cavity is covered by the lateral upper
lip, so that the posterior nostril opens within the lip,

Fig. 304. Jaw-teeth of the left side in the Northern Chimarra ( Cliimcera
monstrosa). After Agassiz. A, seen from without; B , from within.
da , anterior dental plate of the upper jaw; dp , posterior dental plate of
the same; di, dental plate of the lower jaw. Cf. fig. 294 (p. 1065).

inside the cavity of the mouth. The Chimaera can
thus respire even when the mouth is closed.

The lips are thick and fleshy, studded outside and
at the margins with small protuberances (papillae), the
upper lip, as we have already hinted, being divided
into three parts, a quadrangular middle part (median
upper lip) and two larger wing-shaped lateral lobes (la-
teral upper lips). The median upper lip as well as the
underlip is double. The former partially covers the two
anterior dental plates (fig. 294, da) of the upper jaw.
The underlip is entire in front, but furnished behind
with a broad, pendent flap at each corner of the mouth.
The dentition of the mouth consists of two pairs of
palato-dental plates, one pair (figs. 294 and 304, da)
before the other (dp), at the margins of the upper jaw,
and a single pair of mandibular plates (di). The inter-
maxillary (prepalatine) plates (da) are quadrangular and

most distinctly grooved, with 6 rounded ridges; the lateral
maxillary (palatine) plates (dp) and the mandibular plates
(di) are more triangular, undulate and nodose on the
inside. The tongue is small, but free at the tip, and is
densely set with papillae. The front of the branchial arches
is also studded with similar papillae; but soft gill-rakers,
about 12 on the first branchial arch, are besides present.

The three free gill-arches are complete (with bran-
chial lamella; on both sides), but both before and behind
them lies a half gill (single row of lamellae), the former
coalescent with the operculum, the latter with the hind
(abdominal) wall of the branchial cavity. The gill-
openings are about equal in depth to the length of
the orbits, and are set far down, though separated by
an isthmus the breadth of which is about equal to the
depth of the orbits.

The general form of the fins we have already no-
ticed. The triangular first dorsal begins just behind the
head, the length of which varies between 80 and 90 %
of the distance from this fin to the tip of the snout.
The spine at its anterior margin lies exactly above the
base of the pectoral fin and sometimes (in the males) is
only a little shorter than the head; sometimes (usually
in the females) only 4/5 as long. It is rather sharp in
front and is marked anteriorly on each side with a
distinct groove. Both its posterior margins are armed
superiorly, usually somewhat more than half-way clown
the spine, with pointed, descending prickles, and its top
is usually free from the remainder of the fin to a point
a little further downwards. Behind the spine the fin is
supported by fibrils, gathered into about a dozen sepa-
rate bundles, which resemble fin-rays. The posterior
margin is somewhat concave, and the tin-membrane is
prolongated to the vanishing point backwards along the
dorsal margin and more or less near to the second
dorsal fin. The latter tin begins at a distance from the
tip of the snout measuring a little more or less than
twice the length of the head and extends back about
half-way along the rest of the body, measured from the
commencement of the fin. Its height is almost uniform
and only about 1/i or 1/5 of that of the first dorsal.
Just behind its termination the upper caudal lobe begins.
The lanceolate caudal' fin is made up of two almost si-
milar lobes, one over and one under the tail, whose tip
narrows uniformly to a filament, and of which they
occupy the anterior half or third, before they gradually
and, at last, imperceptibly disappear. The anal fin con-

a Answering to one of the stages in the development of the nostrils of the higher vertebrates.

NORTHERN CHIMiERA.

1083

sists of a small lobe, obliquely triangular and posteriorly
somewhat elongated, below the end of the second dorsal.

The paired fins are lobate (with fleshy, broad,
brachial base, so far as the radialia [fig. 294, 1 — 6 and
fig. 295, 1 — 2} extend and form the rounded basal disk)
and obliquely pointed, more or less sickle-shaped, the
pectorals more than the ventrals. The former are set
vertically on the lower part of the sides, just behind the
gill-openings, and are very large. Their tips extend,
when laid back, at least to the insertions of the ventrals,
in the males usually beyond the said points. The ven-
tral fins are set about half-way between the tip of the
snout and the beginning of the caudal fin, the length
of the head being about 2/5 of their distance from the
tip of the snout. They are only about half as long as
the pectorals, measuring in the females about 3/i, in the
males about 88 %, of the length of the head. Before
them lies in the males, within a dermal sac, opening at
an oblique, slit-like aperture, the tongue-shaped, flat,
but somewhat twisted anterior copulatory organ (fig. 295,
prp), armed at the inner margin with 6 or 7 pointed,
curved teeth, and articulating with the fore end of the
pelvis. Behind them and on their inside are situated the
posterior copulatory organs (pterygopodia) of the males,
which are trifid at the tip for about 2/3 of their length.
The three branches are equal in length, but differ in
thickness, the lowest (lower inner) branch (fig. 295, ii)
being invested only with a thin dermal covering, where-
as the two upper branches have a thick skin, with
numerous prorsal denticles on one side. When at rest,
however, the lower inner branch (ii) lies so close to
the upper inner (si) that the apparatus seemingly con-
sists of only two sections.

The Northern Chimsera, when alive, is a brightly
coloured fish of a beauty more striking than agreeable.
The back is reddish brown, lighter or darker; the sides
are for the most part silvery, shading above into blue;
the lower parts of the body white. But under the sil-
very lustre of the sides the dorsal colour spreads in
curious figures, oblong spots, arranged in longitudinal
rows, or irregular, sinuous, and indefinite patches (clouds),
a kind of marbled pattern being thus formed. The sil-
very lateral line is sharply marked by its brown edges.
The top of the head partakes of the dorsal coloration,
but in front is crossed with the retiform design which we
have already noticed; its under surface shares in the

white of the belly. The iris has either a golden or a
silvery sheen; the pupil a greenish lustre. The unpaired
fins are of the same colour as the back; but a black
border extends throughout the length of the caudal fin,
more or less far forward along the second dorsal, and
along the upper posterior margin of the first dorsal.
The paired fins too are similar in coloration behind to
the back"; the anterior (under) surface is lighter, with
rays of an ashy gray. The cavities of the mouth arid
pharynx are more or less black, but the tongue and
the branchial arches yellowish.

Of the internal organs Retzius and Hollberg (1. c.)
have given an exhaustive description, and we shall here
merely refer the reader to our above remarks on this
head, adding that the bluish black intestinal canal is
short and almost straight, with only three coils in the
spiral intestine, the heart extremely small, and the bi-
lobate liver, especially the right lobe, very large and
oily. The well-developed spleen is of a triangular fusi-
form shape and lies beside (under and behind) the pan-
creas, between the inferior edges of the lobes of the liver.

The Northern Chimaera has its proper home in the
depths, some hundreds of fathoms below the surface,
but frequently ascends to a higher level, to a depth of
40 or 30 fathoms, where it is occasionally taken on
Haddock-lines. It often falls a victim also to the not
unusual fate of deep-sea fishes, being carried involun-
tarily to the surface and cast ashore dead or in a help-
less condition by storms. It has a wide geographical
range in the North Atlantic and the Mediterranean as
well as in Japanese waters. It occurs besides, according
to Dumeril’s statements from the Museum of Paris, off
the Cape of Good Plope. Within the limits of the Scan-
dinavian fauna it 'is known from East, Finmark to the
west coast of Jutland and the Sound. It is most com-
mon on the coasts of Norwegian Nordland and Bergen
(Collett). Hollberg remarked that it was common in
Bohuskin during the Herring-fishery of last century; but
it afterwards became rare, and each time a Chirmera was
seen there, the revival of the said fishery was eagerly
expected. Off Mount Kullen it has been caught on se-
veral occasions, and in the Sound it has been met with
between Landskrona and Hveen (Nilsson). From the
Belts and the Baltic it is unknown. Off the coasts of
Iceland it is rare (Faber). It is equally rare in its oc-
currence off the Shetland and Orkney Islands and on the

“ See the special figures to fig. 3 in our plate, which represent the form and colour of the left pectoral and ventral fins, seen from
behind, in the female.

1084

SCANDINAVIAN FISHES.

Scotch coast, oft’ Banffshire (Fleming and Day). In the
Mediterranean, according to Giglioli", it is not uncom-
mon oft’ Nice, Genoa, and Palermo; but during 21 years
Costa could only procure 16 specimens from the Bay
of Naples. On the east coast of North America it is
found from Cape Cod northwards (Jordan and Gilbert);
but Fabricius does not include it among the fishes of
Greenland. According to Schlegel it is rare off South-
ern Japan, but common to the north, especially in
autumn, when it pursues the Herring-shoals to the
very heads of the inlets.

The life and habits of the Northern Chimaera are
otherwise little known. Its intestinal canal has been
found to contain the remains of fishes6, testaceans, crus-
taceans, Echinoderms, and worms. A few fragments of
seaweed among the contents have, no doubt, been swallowed
accidentally with the food. The dentition of the mouth is
evidently intended both to cut up flesh and crush shells.
The spawning-season of this fish is unknown; but in the
month of February, according to Lilljeborg, each ovi-
duct of a female nearly 1 m. long contained in the uterine
dilatation an egg ■with the parchment-like shell almost
fully developed and 122 mm. long. Esmark received
from Christiania Fjord another egg, which has been
described by Collett. It was of a lustrous brown, but
empty, and had probably been washed ashore. The thick
end was cylindrical c, the small end prolongated into a
filamentous appendage. The whole egg was longitudi-
nally fringed all round with a fin-like membrane'6, con-
taining rays directed obliquely outwards towards the
thick end. A similar, though rayless membrane followed
the middle of the egg, at right angles to the plane of
the former membrane, along the whole of one side and
the narrower part of the other. The length of the egg
was 163 mm., exclusive of the thread at the small end,
which appendage measured 42 mm. Its greatest breadth
was 42 mm., exclusive of the fin-like membrane on each
side, which was 4 mm. broad. During the English ex-
pedition on board the Triton a young male 4 1 /2 in. long,
which appeared to have just been hatched, was taken, ac-
cording to Gunther (Deep Sea Fish., Chall. Exped., p.
12), at a depth of 505 fathoms.

The economical importance of the Northern Chimaera
cannot be considerable, for as a rule only solitary spe-
cimens are met with, and the flesh is worthless as food;
but the oil that flows from its liver has been highly
esteemed in medicine. “Since olden times”, wrote Holl-
berg in 1821, “the islanders of Bohusl&n and especially
of Norway have appreciated the value of the fat prepared
from the liver of the Chiimera, as an excellent external
remedy for stiffness of the joints, gout, rheumatic pains,
glandular swellings, cataract, etc. The fishermen of the
northern island-belt of Bohuslan frequently offer this fat
for sale in the stomachs of Ling, for it does not fail to
find a ready market either in chemists’ shops or among
the peasantry. In all likelihood, however, not all this
fat is procured from the rare Chimatra, though it always
bears the name of that fish.” The oil of the Chimaera
is also used internally, like that of the Ling and Cod,
to alleviate weakness and disease of the respiratory or-
gans; and it has been especially prized as an internal
remedy for the sting of the Weaver. The great oiliness
of the liver may be seen even in Chimaeras that have
been preserved entire for scores of years in the spirit-
jars of museums. On opening their abdominal cavity,
oil runs copiously from the belly.

The Northern CMmaera has many names. The
nostrils and the invariably visible front teeth, in com-
bination with the mobile snout, give it a certain resem-
blance to a grinning monkey, whence the name of simia
marina ( hafsapa ). The thread-like tail and the wriggling
movements have given rise to the name of Jiafmus (Sea-
mouse). Linnaeus called the species vidunderfisk (Mon-
ster-fish). In Norway it has borne the names of Isgalt,
Gulhd, Blanklid, Sorotte, Somus , Hdnms (the last three
= Jiafmus), Sorcev, Solvfisk, Havkruge , Havkatt, Spil-
strceng-Hyse. In the Shetland Islands it is known, ac-
cording to Day, as the King of the Herrings and the
Babbit-fish. The former name occurs even in Dauben-
ton e, and is said to be derived partly from the ravages
committed by the Chimaera among the Herring-shoals,
partly from the hooked organ on the forehead of the
males, which appendage has been compared to a crown.
The latter name has the same origin as hafsapa.

a Espos. intern, di Pesca in Berlino 1880, Sez. Ital., Catal., p. 111.
b One of our specimens had Herring-scales in the mouth.

c According to Lilljeborg’s description of the egg-shells, which had not quite reached their full development, compared with Dume-
ril’s figure of an egg probably belonging to the nearly related genus Gallorhynchus {Hist. Nat. Poiss., pi. 8, fig. 8), the thick end of these
eggs is furnished with two short filamentous appendages.

d In the egg described by Dumeril this membrane was covered with silky hairs.
e In the old Encyclopedic M ethodique.

HAYS AND SHARKS.

1085

EL ASMOBRAN CHII PLAGIOSTOMI.

Elasmobranchs with several outer branchial apertures and with more or less distinct'' spiracles. Skin com-
monly0 shagreened, with papillse and spines (placoid scales). Notochord constricted by the formation of ver-
tebrae. Palatoquadrate part mobile independently of the rest of the skull and dentigerous. Paired fins

inserted horizontally.

That course of evolution from the PaJceichtliyes
which is represented in our times by the Plagiostoms,
the suborder of the Sharks and Rays, has advanced so
far, as regards the structure of the spinal column, that
the most highly developed forms possess complete ver-
tebrae, fully divided from each other, and retaining the
notochord only in the intervertebral spaces between the
centra, which are amphiccelous or conically hollowed at
both ends. In their typical forms the two well-known
phalanxes — the Sharks, with their elongated, fusiform
or clavate body, and the Rays, whose pectoral fins, ex-
panded and coalescent with the sides of the body, give
the forepart (the head and trunk) a discoidal shape —
are even externally so unlike each other that no other
character than the outer shape seems necessary, espe-
cially in the case of the Scandinavian fauna. But inter-
mediate forms, Rays with less expanded pectoral fins
or with an unusually terete and thick tail, and Sharks
with triangular, wide-based pectoral fins and a body
more than commonly depressed, occur in foreign seas;
and during youth — in a larval stage when the Pla-
giostoms are still furnished with filamentous external
gills — the Rays have a great external resemblance to
Sharks. The latter were undoubtedly the earlier forms,
geologically speaking, and in the present age the most
imperfectly developed forms — as Hasse in particular
has pointed out with respect to the structure of the
spinal column — are Sharks.

Even from the Silurian deposits ichthyodorulites
are known which have been referred to a Selachoid
genus ( Onchus ), and in the Carboniferous seas Sharks
were very numerous: — the family of the Cestraciontidce,
which still survives in the Pacific,, already existed. And
ever since the Jurassic Period the Not.idanoid family
has survived with a structure which in the tropical
and subtropical seas of the present age represents

the lowest stages in the evolution of the Plagiosto-
mous type.

Individually too most of the Plagiostoms show
great tenacity of life. During the fishery for the Green-
land Shark in the Arctic Ocean between Norway and
Bear Island, these large fishes are hauled up from a
depth of a hundred fathoms or so and deposited on the
deck of the fishing-vessel. There they lie motionless,
partly, no doubt, owing to their sluggish temperament,
but probably stupefied as well by the sudden reduction
of the pressure in which they are accustomed to live.
Their belly is opened with a large knife, and the liver
removed for the sake of the oil which it contains; but
nothing is done with the rest of the body, unless a
fresh bait be required for the huge hook. A Greenland
Shark in this condition may be - skinned and entirely
disembowelled; but the manifestations of life do not
cease for many hours. Even after the head has been
cut off from the body and has lain some hours by
itself, it is dangerous to get one’s fingers between the
jaws, for they may easily be bitten off, and the bite is
so tenacious that one may attempt in vain to extricate
what the jaws have once grasped. Couch tells an anecdote
of a Blue Shark which had been hooked and deprived
of its liver in a similar manner. With the entrails
hanging out of the belly it was restored to the water
and followed the fishing-vessel for some time. It was
not long before the fish tried to seize a Mackerel that
had dropped from the net of the vessel. On another
occasion a Shark was thrown overboard after being
decapitated. For some hours the fish kept swimming
round about — as a boy on board expressed himself —
as if the body was looking for its head. Of the common
Skate the same author relates that its heart may be cut
out and retain contractile powers for at least 25 hours,
when he saw the auricle beating five times a minute.

° rclayiog, slanting , oblique , and Gvoya, mouth.
b Though sometimes obliterated.

c The Electric Rays and young Eagle-Rays, with smooth skin, are exceptions.

1086

SCANDINAVIAN FISHES.

PLAGIOSTQMI BATOIDEP.

Base of the pectoral fins extended backwards along the sides of the body and forwards along those of the head ,
above the five branchial apertures. Tail slender, terete ( whip-like ) or depressed. No anal fin.

All the vertical fins belong to the tail.

The most important distinctions between the Rays
and Sharks we have already mentioned in the intro-
duction to the Elasmobranchs. They are bound up with
the different habits of these fishes. Nearly all the Rays
are ground-swimmers and live in the same manner as
the Flounder-fishes, only seldom attempting rapid ex-
cursions or movements in the water. The above-men-
tioned intermediate forms — the well-known Saw-fishes
( Pristis ) and their nearest relatives, without a saw, the
Rhinobatidce, a family common principally in Indian
waters — therefore approach, in their way of life as
well, the transition to the Sharks.

Among the vital capacities of the Rays one, namely
the power of giving electric shocks, possesses a special
interest. Not only is this faculty a rare phenomenon
in the whole animal kingdom and the peculiar property
of fishes — even among these it is shared, in a singular
and hitherto unexplained manner, by representatives of
widely separated orders and families. At least two
Teleosteous families* 6 are endowed with this power, and
a third c possesses it in so slight a degree that doubt
is still felt whether this family should be included
among the electric fishes or not; but none of these
Teleosts is so nearly connected with the Scandinavian
fauna as to have induced us to describe its faculties.

That electricity is really present both in muscles
and nerves, or, to express ourselves more cautiously,
that at least under certain circumstances electricity
may be stored in muscles and nerves, has long been
known; but nowhere has this electricity such an oppor-
tunity of accumulating as in certain fishes. The earliest
and best known among these fishes are the Electric
Bays. Their family, the Torpedinidce, is fairly rich
in forms: about 20 species, distributed among 6 genera,
have been described with varying accuracy from the
tropical and temperate seas. Three of them — ■ Torpedo
nobiliana, Torp. ocellata ( narce ), and Torp. marmorata —

live in the Mediterranean and the adjoining parts of
the Atlantic, the last-mentioned species being besides
an inhabitant of the Indian Ocean.

The power possessed by the Electric Rays of ac-
cumulating electricity in their organs and voluntarily
discharging these at need has been known at least since
the time of Aristotle and Theophrastus; and Roman
physicians, when Galen lived, if not before, employed
these Rays in the treatment of gout and nervous dis-
eases. In recent times this electricity has been more
minutely studied, and has been found to be identical
in its effects with other electricity. It has the same
influence on the magnetic needle, it can decompose
chemical compounds, and it emits the electric spark.
When the Neapolitan fishermen have drawn their seines
ashore, it is usually, according to Owen, their first
care to pour a bucket of water over the catch. If the
net contains an Electric Ray, its presence is at once
betrayed by the shock transmitted along the stream of
water to the hand holding the bucket. To handle a
live Electric Ray is unpleasant enough, for the arm at
least is paralysed by the shock for a long while.

Like the still more powerful Electric Eel — which
is not an Eel, though it has the external appearance
of one, but rather a Sheatfish, an inhabitant of fresh
water in tropical South America — these Rays employ
their electric power both to kill or at least to stun
their prey and to defend themselves against their ene-
mies. The other electric fishes known to us are far
inferior in their capacity of accumulating electricity,
and probably have recourse to this faculty in defence
alone. Other fishes again scarcely have the power of
giving appreciable electric shocks, but possess true elec-
tric organs, which on account of their feebleness have
been called pseudoelectric. Such organs appear in se-
veral of our common Rays. Their strength is not
great, and it has seldom been observed; but in 1888

? Gr. ficcTog, Ray.

6 Fain. Gym.notidce (Electric Eels) and the subfamily Malapterurince of the great Sheatfish family.
c Fam. Mormyridas.

KAYS.

1087

it was tested by Sanderson and Gotsch® in the case
of the Thornback ( Baja clavata ) and found to be about
710 of that possessed by the electric organs of the
Torpedo. Strange to say, however, these so-called
pseudoelectric organs of our common Rays are situated
in an entirely different part of the body from the elec-
tric organs of the Torpedinidce. Whereas the latter
(fig. 305, E ) lie on the sides of the head and the
branchial cavities, the former are placed on each side
of the dorsal line of the tail. With this difference are
connected two other essential dissimilarities: the nerves
which run to these organs are of a totally different
origin in the Electric Rays and the common Rays, and
the arrangement of the respective organs is entirely un-
like. The electric organs of the Torpedinoids lie in

0

Fig. 305. An Electric Ray ( Torpedo marmorata) with the skin re-
moved from the electric organs ( E ), the skull (S), and the branchial
cavities ( K ). 0 , eyes; Sp, spiracles. After Wiedeksheim.

the region of the transverse muscles, the levators and
depressors of the branchial arches, and are supplied
with nerves from the fifth and tenth cranial pairs and
have their elements transversely arranged — the piles
are vertical. The electric organs of our common Rays,
on the other hand, are situated in the region of the
great caudo-lateral muscles, at the lateral edges of the
tail, close under the skin, and their nerves are spinal,
their elements arranged in the longitudinal direction of
the body. The difference thus affects both the parts of

the body to which the organs belong and the innerva-
tion of the organs. ■ Such a genetic connexion between
these organs in the different fishes as that one form of
them can have derived its origin immediately from the
other, we can therefore scarcely expect to find; but
they have one thing in common, their development
from muscular substance: — the electric organs are
transformed muscle fibres. This was first shown by
Babuchin'' in the case of the Rays, and was still more
clearly elucidated by Fritsch’s examination c of the
Electric Eels brought home by Sachs; but a special
interest, of a yet wider importance, attaches to Ewart’s
investigation s'* of the development of the electric ele-
ments in our common Rays. From these researches it
appears as if we might be able to deduce from the
structure of the electric organs an explanation of the
composition of striated muscular tissue and of the im-
portance of the different substances in this composition e.

A muscle or a portion thereof may be converted
into an electric organ at entirely different periods in
the life of the fish. In the Torpedo, for instance, this
transformation takes place at an early stage, during its
embryonic existence within the egg and almost simul-
taneously with the appearance of muscular cells, before
these are fully differentiated from other embryonic cells.
In a common Ray, on the other hand, that part of the
upper caudal muscles which is destined for conversion
into a pseudoelectric organ exhibits the same for-
mation in the larval stage as the other parts of these
muscles, with the typical composition of the muscular
fibres unaltered; and the observation of the subsequent
changes shows in a series of different developmental
stages how one constituent of the muscle fibre sepa-
rates from the other, each being destined for a distinct
purpose, the sarcoplasma probably to intercept and
store the electricity, the rhabdia to serve as a non-con-
ductor. In the common Skate {Baja batis ), for example,
this indeed takes place before the embryo leaves the
egg-capsule, but not until it has attained a length of
about 3 in. (772 cm.) and has its entire organic sys-
tem, even the muscles, typically formed. In the Starry
Ray {Baja radiata ) the development of the electric
organs does not commence until the fish measures 12

“ Journ. Physiol., vol. 9, Nos. 2 and 3.

b Centralbl. Medicin. Wiss. 1870, p. 259; Arch. Anat., Physiol. 1876, p. 501.
c Dr Carl Sachs Unters. am Zitteraal, Leipzig 1881.
d Phil. Trans. Roy. Soc. Lond., vol. 179 (1888), B, pp. 399 and 539.
e Cf. above, p. 662, on Rollett’s investigations of the muscle fibres in Hippocampus.

Scandinavian Fishes.

137

1088

SCANDINAVIAN FISHES.

cm., long after it has left the egg and in general
assumed the external form of an adult member of the
species. The course of the development is, however,

the same and simple enough. One muscle fibre after
another (fig. 306, A, 1 — 5) thickens at one end- — where
the many terminal fibrils of the motor nerve are at-

Fig. 306. Development of the electric elements in the tail of a common Skate (Raja batis). Magnified. After Ewart.

A: Horizontal section through a part of the lateral muscle of the tail in an embryo 7 cm. long. The muscle fibres are obliquely exteuded
between the intermuscular septa (s), and their transformation proceeds from in front, the hindmost of the figured fibres (1) being entirely
unaltered, the others (2 — <5) more and more club-shaped the further forward they lie. In all these fibres the muscle nuclei and the stria-
tion are still distinct, even at the expanded end; but in the most transformed (club-like) fibres the nuclei begin to pass towards the thick-
ened end. Between the fibres and the intermuscular septum lie numerous connective tissue cells, and between these nerve fibrils are

supplied to the fibres.

B: Two isolated clubs from a somewhat older embryo. The thick end is here flattened and at the top (in front) has a layer of sarco-
plasrna with muscle nuclei (a), under this a denser striation (rhabdia), within which the nuclei are beginning to disappear (6), next a layer
of sarcoplasma with nuclei and sparser striation (c), and at the bottom the aborting stem (d). a is the rudiment of the so-called electric
plate, to which the nerve fibrils (e, with nerve nuclei and connective tissue bodies) are attached; b is destined to form the so-called striated

layer, c the so-called alveolar layer.

G and D: Two different stages of development approximating to the bilboquct form. Letters as in B.

E: A fully developed electric element, with a part of the stem (</) still persistent and at the posterior end (e) still showing traces of stria-
tion; a, b, and c as in B. f, gelatinous layer of connective tissue; g, nerve fibrils; n, nerve fibre; s, septum between two electric elements;

s', part of the original transverse intermuscular septum.

RAYS.

1089

tached — and the fibre now resembles a club. The
thick end' (inner or anterior end) soon expands still
more (fig. 306, B) and is hollowed into a cup shape,
from the bottom of which the thin end (d) of the
fibre proceeds in the form of a shaft, the fibre now
having the appearance of a bilboquet. The body is
further flattened (fig. 306, C and IJ), and the shaft is
more and more reduced, until it finally bends quite to
one side (fig. 306, E) or disappears, the fibre being
now similar to a flat disk. In the common Skate the
electric elements pass through all the three stages of
development, in the Sandy Ray (Baja falsavela ) only
the first two, in the Starry Ray only the first stage.

In the electric elements of a Starry Ray, which
are club-shaped but always somewhat concave at the
top, the two constituents of the muscle fibre, the sarco-
plasrna and the rhabdia, are partly separated from each
other. They have gathered at the thick end more
densely than in the slender portion of the fibre, and
a special layer of sarcoplasma, a so-called electric plate,
is interposed at the top, immediately below that part
of the sarcolemma where the nerve fibrils are inserted.
Under this electric plate lies a densely striated mass,
consisting principally of rhabdia, but containing nu-
merous muscle nuclei, an indication that a considerable
amount of sarcoplasma is present. In the cup-like
(bilboquet-shaped) elements of a Sandy Ray the stri-
ated layer contains far fewer and more scattered muscle
cells; and under this part the granular substance (the
sarcoplasma) has formed a distinct layer with pro-
cesses and alveoli (sponge-like meshes), a so-called
alveolar layer. In the common Skate the rhabdia is
entirely separated from the sarcoplasma and contains
no muscle nuclei. Each electric element thus consists
in this species (fig. 306, E) of three plates (a, b, c ),
one above another, the outer pair (a and c) being,
however, confluent at the margin. With the addition
of the secondary growths of gelatinous (/) and fibrous
(s) tissue that extend into and fill up the spaces be-
tween the electric elements proper and also support
capillaries and nerves, we find here, in the common
Skate, an electric apparatus of essentially the same
structure as in the Torpedo and Electric Eel. On each
side of a comparatively firm plate, in section densely
striated and originally formed of rhabdia, are set layers
of sarcoplasma differing in their histological structure.
This difference may perhaps produce different kinds of
electricity. In that case we should here possess an

analogue to the well-known galvanic piles. Or per-
haps the electricity may be of the same kind in both
layers of sarcoplasma. In the latter case the electric
apparatus of these fishes corresponds to an agglomera-
tion of electrophori.

The manner in which these organs are employed
by our common Rays is not yet known. Their elec-
tric faculty cannot be doubted; but thousands of these
Rays pass through the hands of fishermen without any
recorded instance of a human being having experienced
an electric shock from one of these fishes. When a
Ray feels the resistance of the hook, and the fisherman
begins to haul in the line, it first endeavours to cling
to the bottom by clasping its pectoral fins round some
hard object or tightly pressing them to the ground;
and when it emerges from the water, it tries to defend
itself by raising the pectoral fins as a shield for its
body, bending the tail upwards and forwards, and
dealing violent blows with the latter member. Now
the tail of these Rays is so formidably armed with
large, pointed spines that these in themselves are a
sufficient warning to the fisherman against handling
the Ray with the naked hand until he has given it a,
finishing blow, and hence it may well happen that as
a rule he has no opportunity of experiencing the elec-
tric powers of the fish. Whether the electricity is
utilised in combats with other inhabitants of the deep,
in self-defence or to stun a victim, is also unknown.
We know that dolphins (toothed whales) are the worst
enemies of the large Rays, Ling and Halibut of the
smaller ones; but Saville-Kent saw dolphins seize
Rays by the tail, the very part where the electric or-
gans are situated. From observations in aquaria we
learn that the Rays secure their prey by casting them-
selves over it with a sudden movement of the pectoral
fins, squeezing it under their body, and gliding over
it, until it can be grasped with the jaws. In these
operations the tail can evidently serve merely as a rud-
der to direct the course of the fish. So too, Avhen
the Rays swim freely about in the water, propelling
themselves by undulating movements of the pectoral
fins, the tail can only steer and preserve the equipoise
of the body. Saville-Kent compares the movements
of a swimming Ray to those of a wading bird that
stretches its long legs behind it in its flight.

The system of the lateral line in the Rays requires
a special chapter, though it rather closely resembles that
of the Holocepliali. It is generally far more developed

1090

SCANDINAVIAN FISHES.

than in other fishes. Four kinds of organs belonging
to this system may also be distinguished: 1) the true
lateral line with its ramifications on the head, the
trunk, the pectoral fins, and the tail; 2) the so-called
Lorenzinian ampul lm (muciferous sac ducts); 3) the so-
called Savian vesicles; 4) the so-called pit organs.

The lateral line, the histological structure of which
has been elucidated by Leydig (fig. 307), has nume-
rous ramifications, and opens on the surface of the body
not only into simple pores, but also into long trans-
verse branches originating at right angles or obliquely
from the line itself and its ramifications. The main
branches of the system indeed find their parallels in
the ramifications we have seen above in the Chimsera;
but peculiar to the true lateral line of the Rays is the
complex network of canals on the pectoral fins. In the
genus Baja we recognise on the upper surface of the
head the transverse occipital (supratemporal or aural)
branch (fig. 308, 21), the frontorostral branch with its
divisions, the occipital (20 to l) and the supraorbital

d b

Fig. 307. Part of a branch of the cephalic system of the lateral line
in the Thornback ( Raja clavata). Magnified. After Leydig.
a, outer wall of hard membrane, removed on one side to show b,
the inner, soft membrane, on which are visible the papillae (c) and
the nerve centra (sensory spots, d), with extended ramifications of
the entering nerves (e).

(1 to 2), and the suborbital branch (7 to 8), which
runs on each side of the body between the eye and the
spiracle. When these branches have advanced nearly
to the margin of the body (at 2 and 6’), the former
quite close to the tip of the snout (2), the latter some-
what further back (8), they pierce the body to reappear
on the under surface of the head (2 to 6 and 8 to 11),
and join each other as in the Chimaera. This junction
is simple, however, in the said species and follows the
median line of the snout (median canal, Garman); here
it is double, one branch on each side of the median
line, and the immediate continuation of the frontorostral
branch (supraorbital division) bends outwards on the
under surface of the snout in a sharp crook (from 3
to 4 and 5). After the junction the suborbital branch

is continued backwards by a maxillary part (6 to 9),
that bends inwards, in about a line with the middle
of the length of the nasal valvule, to a naso-maxillo-
rostral branch (9), which indeed (at 10) joins the cor-
responding branch of the other side to form a ma-xillo-
nasal canal, but also sends out a maxillo-rostral canal,
straight to the tip of the snout, within the continua-
tion of the supraorbital branch on the under surface
of the snout. The posterior and more immediate back-
ward continuation of the suborbital branch and its ma-
xillary part answers to the opercular canal of the Chi-
maera, and has been named by Ewart the hyomandi-
bular branch (12 to 19). This branch runs here (12
to 13) on the outside of the branchial apertures (aphj,
following the direction of the series formed by the
latter, and on the under surface of the pectoral fin
bends in a great loop, first backwards and outwards
(13 to 14), then almost straight forwards (14 to 15),
to a point near its origin, where it bends inwards (15
to 16) and afterwards runs forwards (16 to 17), pa-
rallel to its commencement and to the suborbital branch,
until (at 17) it comes in a line with the nostrils, where
it turns upwards straight through the body. On the
dorsal side it now bends straight back (from 17 to 18),
and after an inward curve (at 18), where it receives
connecting ducts from the suborbital branch, it pursues
its course along the margin of the pectoral fin, and in
the posterior part of the dorsal side thereof (at 19)
joins a pleural branch (22 to 23) from the lateral line
proper. The last-mentioned part of the system follows
on each side of the body the same course as in the
Teleosts, from the temporal region (at 21) to the tip
of the tail. At the shoulder- girdle it forms an outward
bend and here it gives off on the dorsal side of the
pectoral fin both the anterior pleural branch which we
have noticed above at its junction with the termination
of the hyomandibular branch, and, in the genus Baja
and its nearest relatives, a posterior pleural branch
(24 to 25), which ramifies in the posterior part of the
pectoral fin. Just behind the mouth there lies, as a
detached portion of the hyomandibular branch, a trans-
verse canal (27), answering to the mandibular branch
of the Chimsera.

The ampullary system (fig. 309) properly belongs
to the head alone, where all its canals have their csecal
base. These organs have been known for more than
three centuries", but their true nature was first eluci-

Lorenzini, Observ. int. alle Torped ., 1678, and Munro, Struct., Physiol. Fish., 1785.

RAYS.

1091

dated by Jacobson", Retzius6, and Leydigc. We have
observed them above in the Chimsera. In the Rays
there commonly lie several symmetrical pairs of cap-
sules (fibrous investments of the basal parts of the

ampullae and of the nerve twigs supplied to them), 1) at
the tip of the snout (fig. 308, Ba), 2) further back
on each side of the base of the rostral cartilage, on
the ventral side before the nasal capsules, 3) on the

Fig. 308. Schematic figure of the distribution of the lateral line canals in a Raja , parti}- after Garman and Ewart. Right half of the

body; to the right the dorsal side, to the left the ventral.

1 — 2, course of the supraorbital branch on the forehead and the dorsal side of the snout; 2 — 3 , its backward continuation on the ventral side

of the snout; 3 ■ — 4 — 5, its outward crook on the ventral side of the snout; 5 — 6, canal by which it communicates with the suborbitnl branch;

7 — 8, course of the suborbital branch on the dorsal side of the head; 8 — 3, its backward continuation on the ventral side of the head; 9 — 10 ,
its inward ramification, which at 10 gives out a transverse canal inwards to join the suborbital branch of the other (left) half of the body,
and then advances to its termination at the tip of the snout, at 11 ; 12 — 13, backward course of the opercular (hyomandibular) branch on
the ventral side of the head and pectoral fin, in a curve outside the gill-openings ( apbr ); 13 — 14—15, its ventro-pleural bend; 15 — 16, its
inward, ventral continuation; 16 — 17, its forward, ventral course side by side with the suborbital branch ; 17 — 18, its backward course on the
dorsal side of the head; 18 — 19, its dorso-pleural curve, which at 19 joins the anterior pleural branch of the lateral line ; 20 — 26, the lateral
line proper; 21, transverse occipital branch; 22 — 23, anterior dorso-pleural branch of the lateral line; 24 — -25, posterior dorso-pleural branch
of the lateral line; 26, lateral line of the trunk and tail; 27, mandibular branch; Ra, rostral ampullae; oa, orbital ampullae; hya, hyoid am-
pullae; via, mandibular ampullae; po, pit organs; o, mouth; oc, eye; sp, spiracle.

“ Bull. Sc. Soc. Philom. Sept. 1813.

b 6fvers. Vet. Akad. Forh. 1845, p. 177, where references are given to the remaining literature on this question.

c Roche?), Haie, Leipz., 1852, p. 41.

1092

SCANDINAVIAN FISHES.

dorsal side, one pair on the outside of the preorbital
cartilage (oa) and another outside the spiracle ( liya :).
Asymmetrical capsules also occur, as in Trygon between
the anterior ends of the two nostrils. The ampullary
canals extend throughout the sides of the body, those
of the pectoral fins for example, which run to the
hyoid capsules, attaining a considerable length (fig. 309).
In the Electric Rays Fritsch" has observed a regular
alternation on the dorsal side of the pectoral fins be-
tween the ducts of the ampullae and those of the lateral
canal proper, the outer (distal) part and the orifice of
each ampullary duct lying as a rule between a pair of
the outward transverse branches of the lateral line.
To this he appends the remark that the presence of

Fig. 309. Hyoid ampulla and their opening ducts together with the
dorsal course of the lateral line in the left half of a Raja.
After Garman.

true sensory bulbs (nerve-bulbs) with the hair cells
occurring in the lateral line has not yet been demon-
strated in the nerve ends on the inner trail of the
ampullae, whence he infers that the ampullae should
be regarded as a secretory part of the system of the

lateral line. Their cavity is filled with an endolymph-
like, more or less coagulated mucus.

The third kind of organs belonging to this system,
the so-called sensory vesicles of SavE, appear only in
the Electric Rays, and are situated in one or two series
on the snout of these fishes and around the anterior
part of their electric organs. They are homologous
with the carnal bottoms of the ampulla:, but are entirely
closed, and their inner surface is usually furnished with
three sensory spots, the middle spot being the largest
and furnished with numerous hair cells.

The fourth kind (fig. 308, po) of the sense organs
now under consideration are the so-called pit organs c
(called by FritsciE spalt-papillen). These are present
in our common Rays, and are wart-like dermal growths,
open at the top and containing a spherical or some-
what more prolongated (bottle-shaped) cavity, which is
filled with sensory cells furnished Avith hair-like pro-
cesses and surrounded by supporting cells. The pit
organs or sensory follicles lie scattered in toavs along
the inferior orbital margins, in the temporal region,
and on each side of the median line of the body, be-
tAveen this and the true lateral line.

The innervation of the entire system of the lateral
line is supplied, according to Eavart, almost exclusively
from branches of the facial nerve* 6 and from the late-
ralis. The supraorbital branch (fig. 308, 1 — 6) and the
rostral ampullae ( Ha) are supplied by the superficial
ophthalmic division of the facial, the suborbital branch
and the anterior part of the occipital, the orbital am-
pullae (oa), and the pit organs (po) at the orbits by the
buccal division of the facial, and the hyomandibular
branch (12 — 19) together Avith the hyoid ampullae (hi/ a)
and the mandibular branch (27) b}r the hyomandibular
division of the facial nerve. The lateral line proper,
as Avell as the posterior part of the occipital branch,
the transverse occipital canal (21), and the majority of
the pit organs (po), is innervated by the lateralis, one
of the main divisions of the vagus.

As Fritsch has remarked, some obscurity still
obtains in the question of the physiological importance
of the entire system of the lateral line; and the division

a Die Elelctrischen Fische, II, p. 87.

6 Matteucci and Saa’i, Traite des phe'nomenes electrophysiologiques des animaux, Paris 1844.
c Trans. Roy. Soc. Edinb., vol. XXXVII (1891 — 92), p. 101.

J Stzber. Akad. AViss. Berk, VIII (1888), p. 291.

e This nerve consequently consists in fishes, according to Ewart, of both sensory and motor fibres. In the higher vertebrates it is
motor alone, the sensorial parts, according to Ewart, having disappeared. According to the older opinion, which is no doubt still maintained
by the majority of anatomists, the divisions enumerated above belong to the trigeminal group.

KAYS.

1093

of work which he assigned on the one hand to the am-
pulla? as secretory organs, on the other to the lateral
line proper and the Savian vesicles as organs of sense,
requires further demonstration. Its anatomical structure
ranges the system of the lateral line with all its modi-
fications as intermediate between the organs of hearing
and those of the senses of pressure (touch — especially
as in the worms), taste, and smell; and its physiological
significance is perhaps best expressed as yet by Ley-
dig’s assumption that it appertains to a sixth sense,
foreign to us, arid receptive of the impressions yielded
by such vibrations of the surrounding medium as are too
grave (slow) for appreciation by the organs of hearing.

The Rays probably spend the greater part of their
existence in a stationary position at the bottom, and to
facilitate their respiration, to keep the gill-slits supplied
with fresh water, they have been provided with large
spiracles on the upper side of the head, just behind the
eyes. Above each spiracle is simple, below the passage
divides into two, one branch entering the cavity of the
mouth, the other passing to the gills. To prevent the
water thus supplied from escaping through the mouth,
they have a palatal fold, usually of powerful develop-
ment, in the upper jaw. On the under surface of the
head the nasal cavities show about the same relation
to the mouth-cavity as in the Holocephali , only that no
lateral upper lip encloses them outside and in front.
Here the median upper lip instead is still more deve-
loped, and forms a broad dermal fold extending from

the anterior margins of the nostrils and between them
back to the corners of the mouth a, on each side cover-
ing a deep groove that runs from the nasal cavity to
the corner of the mouth. On the outside this groove
is bounded by a dermal ridge, which is indeed furnished
anteriorly with a more or less prominent, blunt or
rather pointed protuberance, but which does not form
any limit between the anterior and posterior nostrils.
The live pairs of branchial apertures are set on the
ventral side6, in two more or less straight lines con-
verging behind or in a curve anteriorly concave, pos-
teriorly interrupted, behind the head and between the
prorsal parts of the pectoral tins.

As the form-series of the Rays, which includes
about a hundred and fifty species, is a developmental
offshoot of the Selachian type, the forms that have most
widely diverged from the Sharks must, of course, be
regarded as the most advanced in the scale of evolution.
Among the Rays observed in Scandinavia there are two
families which are both distinguished by the exceedingly
slender (whip-like) tail and by the loss of at least one
dorsal fin. In the third family of Scandinavian Rays
both dorsal fins are persistent on the much broader and
depressed tail, which is besides fringed on the sides
with a more or less distinct dermal fold. But even
this family is more widely removed from the Selachian
type than the three remaining families, which are
strangers to our fauna: the Torpedinidce , Bliinobcitidce ,
and Prist ides.

Fam. M Y L I 0 B A T 1 1) M.

The whip-like tail without caudal fin , hut with a dorsal fin in front , behind which there generally appears a
serrated spine , with or without compensatory spine. The large pectoral fins interrupted on the sides of the head ,

hut continued on the sides of the snout by so-called cephalic fins.

This family contains the giants among the Rays,
some forms being veritable monsters of the deep. In
addition to the characters given above the members of
the family are distinguished by the elevation of the
head above the plane of the pectoral fins, the forehead
being especially high, and causing the eyes to assume
a vertical position with lateral aspect, instead of the

“ The above-mentioned intermediate forms (the Rhinobatidi
Sharks in this respect too, and have the nostrils entirely separated fr<
h Hence the name of Hypotr ernes, applied by DUmeril to the

oblique or horizontal position they occupy in the other
Rays. The skin too is smooth during youth in most
of these fishes, and in older specimens is commonly
shagreened on the tail alone or also on the bases of
the pectoral fins, but sometimes over the whole body.
The family derives its name from the form and molar
function of the jaw-teeth, which are adapted for the

and Pristidce) between the Rays and Sharks are approximated to the
the mouth.

suborder of the Rays.

1094

SCANDINAVIAN FISHES.

crushing of hard-shelled animals. These flat, plate-like
or tuberous teeth are also characteristic of different
genera and species. They are set as if in a mosaic,
fitted beside one another like the stones in a pavement
(■ dentes pavimentati ), and arranged, in the longitudinal
direction of the body, in several rows or in a quincunx,
with the pointed corners wedged in between each other
(fig. 310). Here, as in the Sharks, the largest forms
have relatively the smallest teeth. On account of the
differences that prevail in this respect, Muller and
Henle" distinguished among the forms now under
consideration two families, the Myliobatides and Cepha-
lopterce , which Gunther6 united into a single family

Fig. 310. Jaws of an Eagle-Ray ( Myliobatis aqitila ). After
Agassiz and Gunther.

under the name and with the definition given above,
but divided into two subfamilies, Myliobatina and
Ceratopterina0 .

Our knowledge of the gigantic but comparatively
small-toothed Rays of the latter subfamily is indeed ex-
tremely defective; but they have long afforded material

for fabulous narratives, and should probably be included
in the list of marine monsters that have posed as the
great sea-serpent. Their true homes are the great
oceans and the Mediterranean. As an example of their
magnitude we may cite, after Mitchill, the dimensions
of a Ceratoptera vampyrus taken in September, 1823,
in Delaware Bay, on the east cost of the United States'*.
Its length was 10 ft. 9 in. (32 3/4 dm.), exclusive of the
tail, which measured 4 ft. (12 1 /5 dm.), and its breadth
between the tips of the pectoral fins 18 ft. (nearly 55
dm.). Its weight was so considerable that three yoke
of oxen, a horse, and twenty-two men were required
to haul it ashore. The pectoral fins, as well as the
cephalic fins, which in the above instance were 2 ft. 6
in. (72/3 din.) long, may be folded over so as to meet
at the mouth; the latter fins are besides mobile in all
directions. These Rays often swim in pairs, male and
female. They cleave the water with rapid strokes, like
the flight of a bird of prey; and in pursuit of their
victims, which consist principally of Cephalopods and
fish, they display a litheness in their movements which
one would hardly credit to a Ray. Sometimes they
swim so high that their fins emerge above the surface6;
and when the cephalic fins are thus exposed to view,
the seaman compares them to horns, and hence confers
upon the fish such names as ox, cow, or calf, or even
that of the prince of darkness, which appears in the
form of Bevil-fisli. Others have compared the move-
ments of these Rays to the flitting of a bat, whence
the name of Vampire-Ray. On the Irish coast a small
specimen of a Ceratoptera has once been met with
(about 1828).

Genus MYLIOBATIS f.

Cephalic fi.ns (on the sides of the snout and at its tip) in the same plane as the pectoral. Molars in the middle of
the jaws of adult specimens much (3 — 8 times) broader than long , and larger than the lateral teeth , which are

set in several rows.

This genus too, which contains 7 or 8 species, can so great as those of Ceratoptera: one species, the Me-

boast of considerable dimensions of body, though not diterranean Myliobatis bovina, often attains, according

a System. Beschreib. Plagiostoin., Berlin 1841, pp. 176 and 184.
b Cat. Brit. Mus., Fish., vol. VIII, p. 488.

c Cephaloptera, the genus established by the elder Dumeril, has necessarily been altered to Dicerobatis, the name more recently con-
ferred upon it by Blainville, Cephaloptera having been previously employed as a generic name among birds.

d Isis, vol. XXV, 1832, p. 1063. According to Brown-Goode ( Fisher . Industr. U. S., sect. I, p. 666) this species attains a breadth
of 30 ft. (9 m.) between the tips of the pectoral fins.

e Thus we may perhaps explain the account of the “sea-serpent” as seen by Lieutenant Hayes from H. M. yacht Osborne in June
1877. See Henry Lee, Sea Monsters Unmasked, Handbooks, Intern. Fisher. Exhib. London 1883, p. 94, fig. 23.

/ C. Dumeril, in Cuv., Regn. Anim., ed. 1, tom. II, p. 137. From yvllag, millstone, and jazig, Ray.

EAGLE-RAY.

1095

to Bonaparte, a weight of 300 Italian pounds (about
100 kilo.), and the Japanese form, Gunther’s Mylio-
batis cornuta, which is probably identical in species
with the Atlantic form, sometimes turns the scale,

according to Schlegel*, at 400 pounds (about 180 kilo.).
The genus belongs properly to the tropical and sub-
tropical seas; but one species has strayed northwards
within the limits of the Scandinavian fauna.

THE EAGLE-RAY (sw. ornrockan).
MYLIOBATIS AQUILA.

Fig. 311.

Molars in the middle row within the jaius of adidt specimens 4 — 6 times as broad as long. Tip of the snout
blunt , with a small , prominent protuberance at the middle. A more or less distinct similar protuberance on the
front of the upper orbital margin in the males. Anterior margin of the pectoral fins convex , posterior margin
concave , tip someivhat obtuse. Beginning of the dorsal fin situated about three times the length of its base behind
the insertions of the ventral fins. Length of the tail at least equal to the breadth of the body between the tips
of the pectoral fins or greater. Skin quite smooth or roughened with spinulce only at the root of the tail. Colora-
tion above brownish green or yellowish gray with a bronze lustre , below of a dirty white or grayish brown. Tips
of the pectoral fins dark. Young sometimes spotted with white. Faint traces of dark transverse bands sometimes

present on the dorsal side.

Fig. 311.

An Eagle-Ray, Myliobatis aquila , o71,

'/e of the natural size.

From the Museum Adolphi Friderici.

Syn. Mercg, Aristot., Anim. Hist., lib. V, cap. V. Aigle de mer
(. Aquila marina ), Belon., Nat., Divers. Poiss., p. 85. Se-
cunda Pastinacce species, Rondel., Pise. Mar., p. 338.
Aquila, Salv., Hist, aquat. anim., p. 147. Raja corpore
glabro, aculeo longo serrato in cauda pinnata; Art.. Ichth.
Gen., p. 72; Synon ., p. 100.

Raja aquila, Lin., Syst. Nat., ed. X, tom. I, p. 232; Cuv.

( Myliobatis , ex Dum.), 1. c.; Johnst. ( Raia ), Proc. Berwicksh.
Nat. CL, Sept. 1839 (vide Yarr., Brit. Fish., ed. 2, vol. II,
p. 592); Mull., Henl. ( Myliobatis ), Syst. Beschr. Plagiost.,
p. 176; Dum., Hist. Nat. Poiss. (su. a Buff.), tom. I, p.
634; Gthr, Cat. Brit. Mas., Fish., vol. VIII, p. 489; Mok.,
Hist. Nat. Poiss. Fr., tom. I, p. 442; Coll., Forh. Vid.
Selsk. Cbrnia 1882, No. 29; Day, Fish. Gt. Brit., Irel., vol. |

II, p. 352, tab. CLXXVI; Doderl., Man. Ittiol. Medit.,
fasc. Ill, p. 234; Lillj., Sv., Norg. Fisk., vol. Ill, p. 534.
Myliobatis noctula, Bonap., Iconogr. Fna Ital., tom. Ill, Pesci,
tav. 159.

The Eagle-Ray attains a length of lV2 m. or more.
According to Bonaparte, however, it is commonly
smaller than its Mediterranean congener Myliobatis bo-
vina, which is characterized by a more pointed and
more elongated snout and more pointed pectoral fins.
The rhombic disk is remarkable for its great breadth,
which is sometimes more than twice its length. The

a Myliobatis aquila in Sieb., Fna Japon., Pisces, p. 310, tab. CXLII.

Scandinavian Fishes.

138

1096

SCANDINAVIAN FISHES.

head ■ — with the semicircular” or still more obtuse and
depressed snout — presents an appearance which was
compared by Rondelet to that of a toad’s head, by
Johnston to the print of a horse’s hoof, the great fon-
tanelle representing the impression left by the frog of
the hoof. The eyes are set so high that their superior
margin is almost in a plane with the forehead. Their
longitudinal diameter in the young is about half the
length of the snout, in old specimens less. Just behind
the eyes lie the large spiracles, the length of which is
about twice that of the former. The nostrils are ap-
proximated rather closely to each other, the dermal flap
which covers their internal parts (the nasal valvule)
being medially attached only at its base by a narrow
frenum. This valvule is trapezoidal in form, broad,
and expands behind to the truncate edge that lies, with
a shallow median sinus and fringed with papillae, close
to the anterior margin of the mouth and outside the
very thin true upper lip. The underlip too is thin at
the middle, but thickened at the corners of the mouth.
The breadth of the mouth is about 2/'3 of the distance
between it and the tip of the snout.

The body is somewhat depressed (flattened) behind
the head, but a little further back rises again to the
shoulder-girdle, where the depth is about equal to the
length of the fontanelle mark on the top of the head.
In the median line behind this point there is a low
ridge, which passes on the tail into a dermal edge in
front of the dorsal fin, which is rounded above or
obliquely truncate. Behind this tin lies the spear-like
caudal spine, with faintly convex front, carinated
back, and numerous (40 — 50) retral barbs on the
sides*.

The large triangular pectoral fins have been com-
pared to the wings of an eagle, and have given the ge-
nus its vulgar name. The ventral tins are quadrangular,,
with straight or somewhat convex posterior margin.

The general features of the highly variable colo-
ration have already been indicated.

The Eagle-Ray, which is known from Australia (Syd-
ney) — probably from Japan too — and from the Cape
of Good Hope, has its principal European habitat in the
Mediterranean and the neighbouring parts of the Atlantic.
On the north-west coast of France it is fairly common,
but even on the English coast it is rare. Further
north it is still rarer. On the 16th of November, 1882,
however, a young male0 of this species was taken among
small Herrings at Vettre Farm in Asker, 20 kilom.
south of Christiania. The Herring-seine was drawn on
a clay bottom, at a depth of 12 — 14 fathoms (Collett).

The habits of the Eagle-Ray essentially resemble
those of the Devil-fishes as described above. R is a
less marked bottom-fish than the true Rays. It appears
to fly rather than to swim, says Moreau, whether it
is traversing mid-water or lashing the surface with one
of its pectoral tins. An Eagle-Ray was kept in an aqua-
rium at Arcachon, the same author states, and whenever
it was taken out of the water, it uttered a rather loud
bellowing noise. Its flesh is of little value and is not
much eaten. On the other hand, it is greatly dreaded by
the fishermen for the sting of its spine. The tail is usually
chopped off before handling the fish. According to Mo-
reau the Eagle-Ray is viviparous. A fisherman from Ros-
coff told him that a female of this species had given
birth to seven living young just after she had been
hauled into the boat.

Fam. TRYGONIDfE.

The whip-like tail with or without either fins or spine. Pectoral fins extended without a break forward along
the sides of the head and contiguous in front of the tip of the snout.

This family has its true home in the seas and a scourge to bathers. They generally live in shallow
rivers of the tropics, where many of its members are water, concealing their body in the sand or mud, so

“ In a male 66 cm. long the length of the flattened snout in the median line before the fontanelle is half its breadth at the
same point.

6 In the specimen described by Collett, the tailless body of which measured 318 mm. in length, the true caudal spine was 82 mm.
long, but in front of it lay a compensatory spine, still covered with skin and 11 mm. long. In a male 66 cm. long, the tail of which
measured 40 cm., the former spine was 63 mm. long, the compensatory spine 55 mm.

c Length of the body 853 mm., including the tail (535 mm.). Greatest breadth (between the tips of the pectoral fins) 540 mm.
Length of the copulatory organs (pterygopodia) 78 mm.

STING-RAYS.

1097

that only the eyes and spiracles are free. On being
alarmed they usually take to flight at once — and
bathers or waders therefore stir up the bottom in front
of them with a stick, an oar, or the like, or by scrap-
ing their feet — but if accidentally trodden on, they
promptly dart their caudal spine into the foot or leg,
inflicting an extremely painful, perhaps even a mortal
wound. About fifty species are known — not all, how-
ever, armed with caudal spine — and among them are
several (7) inhabitants of the Mediterranean, which
have rendered the family notorious since prehistoric
times. A Greek myth relates how the sorceress Circe
tipped with the spine of a Sting-Ray — or perhaps of
an Eagle-Ray — the spear she gave to her son Tele-
gonus, when he was setting out to seek his father
Odysseus, and how this spine became the latter’s bane.

The family occupies an intermediate place between
the preceding and the following families. Often the
head is perceptibly elevated, the eyes assuming almost

the same position as in the preceding family, the ven-
tral tins are undivided, never deeply forked or lobed,
and the skin is sometimes almost entirely smooth; but
the extension of the pectoral fins along the sides of
the head ranges this family beside the true Rays. Se-
veral of the forms are also furnished with vertical
dermal folds, sometimes with a true caudal tin, on the
superior or inferior caudal margin, sometimes on both.
Most of them have a caudal spine and sometimes as
many as two or three compensatory spines in front of
the former. One dorsal tin is occasionally present
within this family, but there are never two.

Dumeril" divided the family into four subfamilies,
distinguished by the absence of the caudal spine ( Uro -
gymni ), or by the presence of a caudal tin {TJrolophi),
of a dorsal tin ( Trygonopterce), or by the absence of
both these fins ( Pastinacce ). To the last-mentioned
subfamily, which contains the greatest number of spe-
cies, belongs the

Genus TRYGON'.

One or more dagger-spines furnished with retral barbs on the tail. Where vertical dermal folds appear on the
tail , these are low and do not extend out to the tip thereof. No rayed vertical fins. The transverse cleft,
of the mouth almost straight or at most but slightly curved. Jaw-teeth {fig. 312 ) flattened , triangular , and set
in a dense quincuncial arrangement; in the males sharpened. Disk of the body rhomboidal , of almost equal

breadth and length.

Thus defined the genus includes about a score0 of
ascertained species from tropical and temperate regions
all round the globe. The generic name is of classical
Greek origin, and occurs in many passages of Aris-
totle’s works in its present signification, though it
was originally applied to a dove. The notorious and
dreaded Ray thus became the namesake of the symbol
of innocence, “not on account of its colour,” says Ron-
DELETfi, “for this is yellow, but because of the resem-
blance of the pectoral fins to expanded wings.” Among
the Romans the genus was called Pastinaca, a name
that Rondelet derives from the colour and terete form
of the tail, which in these Rays is like a parsnip.

“ Hist. Nat. Poiss. (su. a Buff.), tome I, p. 579.

b Adanson, Conrs d'Histoire naturelle, 1772, vol. II, p. 170. This work first appeared in print in 1825; but Geoffr. St. Hil.
adopted the name of Trygon from it in 1809 ( Descr . de I'Egypte).
c Gunther diagnoses 23 species in his Catalogue.
d De Pise., p. 332.

Fig. 312. Jaws and teeth of a Sting-Ray ( Trygon pastinaca), 9-
Natural size. After Moreau. Within the figure teeth on a magnified
scale. I, ligaments.

1098

SCANDINAVIAN FISHES.

THE STING-RAY (sw. spjutrockan).

TRYGON PASTINACA.

Figs. 313 and 314.

Tail, which occupies about half ( in the young rather more than half) the length of the body, furnished both above
and below with a shallow dermal fold, the upper, however, insignificant. Length of the disk about 80 — 85 %
(88 %) of its breadth. Tip of the snout more or less obtuse, lateral tips of the disk (of the pectoral fins)
rounded. Ventral fins rectangular, with the inner posterior angle strongly rounded.

Length of the head (from the tip of the snout) to
the occiput about 2/s (37 — 40 %), to the first gill-
opening somewhat more than 7s (34 — 36 %), of that
of the disk. Length of the snout (from the anterior

mouth. Skin of young specimens entirely smooth, of
old often tuberculated in the median line of the back
and above the shoulder-girdle. Coloration above of a
grayish or brownish yellow, or darker, of a blackish

Fig. 313. Trygon pastinaca, $. nat. size. From Mount Kullen, July 24th, 1849. Baron Gyllenstjerna. The property of the

Museum of Lund.

margin of the eyes) about V3 of the former length of
the head. Nasal valvule, which is attached by a nar-
row frenum almost throughout its length, of a rect-
angular form, about twice as broad as long", and its
breadth at the base somewhat greater than the width
of the mouth. Greatest depth of the body (the thick-
ness at the shoulder-girdle) somewhat less than half
the length of the head to the occiput. Teeth of the
upper jaw, with thimble-like indentations on their sur-
face, set in about 20 — 30 rows directed obliquely back-
wards. .Distance of the caudal spine from the root of
the tail about equal to the length of the head. Breadth
of the tail at the base about equal to the width of the

green, in front grayish, in the young sometimes spotted
with white; under surface of a dirty white.

Byn. Pastinaca, la Pastenade de mer, cm Tourterelle, ou Tareronde,
Belon, Nat., Div. Poiss., p. 82. Pastinaca marina Oxy-
rinchos, Sghonev., Ichthyol. Slesv. Hols., p. 58. Raja cor-
pore glabro, aculeo longo auterius serralo in cauda apterygia,
Art., Ichthyol,., Gen. Pise., p. 71; Syn., p. 100.

Raja Pastinaca , Lin., Byst. Nat., ed. X, tom. I, p. 232;
Penn., Brit. Zool. (ed. 1776), tom. Ill, p. 83; Retz., Fna
Suec. Lin., p. 304; Donov., Brit. Fish., tab. XCIX; Cuv.
{Trygon), R'egn. Anim., ed. I, tom. II, p. 136; Nilss. {Raja),
Prodr. Ichthyol. Scand., p. 120; Parn. {Trygon), Mem.
Wern. Nat. Hist. Soc., vol. VII, p. 440; Bonap., Iconogr.
Fna Ital., Pesci, tab. 156; Nordm. in Demid., Voy. Russ.
Mer., tom. Ill, p. 549; Mull., Hle, Syst. Beschr. Plagiost.,
p. 161; Ekstr., Gbgs Vet., Vitt. Samh. Hand!., Ny Tidsf..

Length of the nasal valvule 48 — 56 % of its breadth at the base.

STING-RAY.

1099

II. 1, p. 41; Kr., Damn. Fiske, vol. Ill, p. 1018; Nilss.,
Skand. Fna, Fisk., p. 741; Rhdt, Vid. Meddel. Naturh.
For. Kbhvn 1864, p. 276; Dum., Hist. Nat. Poiss ., tom.
I, p. 600; Gthr, Cat. Brit. Alas., Fish., vol. VIII, p.
478; Winth., Naturh. Tidskr. Kbhvn, ser. Ill, vol. XII, p.
61; Mob., IIcke, Fisch. Osts., p. 158; Day, Fish. Gt. Brit.,
Irel., vol. II, p. 350, tab. CLXXV ; DOderl., Man. Ittiol.
Medit., fasc. Ill, p. 220; Peters., Vid. Meddel. Naturh.
For. Kbhvn 1884, p. 160; Lii.lj., Sr., Norg. Fisk., vol.

III, p. 539.

Trygon lyrnma , GeOffr., Descr. Egypt., Poiss., p. 219, tab.
27, fig. 1.

Raja Sayi, Lesueur, Journ. Acad. Nat. Hist. Philad., vol. 1,
p. 42. Vide Gthr.

Trygon vulgaris (= Pastenague commune ex Cuv.), Risso, Ear.
Merid., tom. Ill, p. 160; Mor., Hist. Nat. Poiss. Fr.,
tom. I, p. 448.

Trygou Akajei, Mull., Hle, 1. c., p. 165, tab. 53; Schl. in
Sieb., Fna Japon., Pise., p. 308. Vide Gthr.

The Sting-Ray, which is common in Southern
Europe, is stated to attain rather considerable dimen-
sions, at least a length of 2l/a m. and a breadth of
IV2 m-) though the occurrence of so large specimens
has not been authenticated in European waters. Bona-

parte indeed speaks of Sting-Rays from the Mediter-
ranean weighing 10 centners; but Risso says that their
average weight in that sea is about 10—12 kilo., and
according to Doderlein the largest Sting-Rays in the
fish market of Palermo weigh 20 kilo, or more. Schle-
gel states that the Japanese form ( Trygon Akajei )
attains a weight of several centners; and the dimen-
sions given above are those of a specimen from the
Cape of Good Hope, now in the Museum of Paris.

It seems rather probable, however, that several
species are included in the above list of synonyms.
Our material is indeed scanty, consisting of two females,
which we figure here, of about the same size; but even
these show divergencies of form uncommon at least in
the other Rays. The one, which was probably the type
of Linnaeus’s Raja Pastinaca , has a less expanded disk,
and apparently comes nearer to the American Trygon
Sayi; the other, the only Swedish example of this spe-
cies on record, corresponds more closely to Bonaparte’s
and Day’s Trygon pastinaca. The most important dif-
ferences are expressed in the following percentages:

Specimen

Specimen

in the

from the

Lund Museum.

Mus. Ad. Frid.

Length of the disk in % of its breadth ..

80. 0

88.6

„ „ ,, head to the occiput . in % of the breadth of the disk

30. 0

35.3

■ • ,, „ „ first gill-opening „

27.5

31.5

Distance between the caudal spine and the root of the tail... . ,, „ „ „ „ ,, „ „

31.6

37.2

Length of the snout from the corners of the mouth in % of the length of the head to the occiput

57.2

63.4

Distance between the nostrils in % of that between each of them and the tip of the snout

75.6

59.6

According to Risso the Sting-Ray is a fish of noc-
turnal habits, hiding itself as we have described above,
and lying in wait for any prey that may approach;
but at night-time it may be taken in gill-nets. When
it attacks a fish, it flings its tail round the victim,
which it then pierces and tears with its sting; and it
defends itself in the same manner from the hand that
would grasp it. Its food consists of fish, crustaceans,
and mollusks. On the Sicilian coast, according to Do-
derlein, it approaches land and is taken in numbers
from the middle of December till the end of March,
during which season it contains eggs of but little de-
velopment. This is probably the breeding season, for
the more isolated females caught in summer are more
or less gravid. The young are born alive.

From the Black Sea and the Mediterranean the
true geographical range of the Sting-Ray extends along

the Atlantic coast of Europe hardly further north than
to England. Even there, as well as on the north-west
coast of France, it is far from common; and to the
north it becomes rarer and rarer. Now and then it is
met with off the Scotch coast. Schonevelde knew it
from Heligoland. Blocii received it from Hamburg.
Kroyer heard fishermen speak of its occurrence off
Hirsholm. Reinhardt received a specimen that had
been taken in the autumn of 1862 off Frederikshavn.
Schrader sent to the Museum of Copenhagen a speci-
men that had been caught on June 10th, 1882, on the
south side of the Skaw. From the fishing-village of
Arild, at the foot of Mount Kullen in Scania, Baron
Gyllenstjerna received, on the 24th of July, 1849,
a specimen which is now preserved in the Museum of
Lund. This example (fig. 313) has a disk 341/2 cm.
broad and 27 1/2 cm. in length from the tip of the snout

1100

SCANDINAVIAN FISHES.

to the extreme end of the margin of the pectoral fin.
Whether more Sting-Rays have been found on the coast
of Scandinavia, we are ignorant; but Retzius included
the species in his edition of Linnaeus’s Fauna Suecica;
and the Museum of Drottningholm contained a speci-
men about 6 dm. long (now preserved in the Royal

Fig. 314. Tryqon pastinaca, $, J/4 nat. size;

Museum, but with the tail broken off short), which
had certainly been in Linnaeus’s hands, but is probably
of foreign origin, as he did not personally recognise
the species as Swedish. The last-mentioned specimen
is represented in the appended figure.

, caudal spine, nat. size. From the Museum Adolphi Friderici.

Fam. RAJIDiE.

Tail depressed, with dermal edge on the sides , flat underneath , convex on the top , and furnished with tivo dorsal
fins, tvith or without caudal fin. The pectoral fins extend forward to the snout or even in front of the rostral
cartilage. Where electric organs are present, they lie on the sides of the tail. No spear-like spine on the tail.

As we have remarked above, these fishes, the fa-
mily of the true Rays, in spite of the singular deve-
lopment of the pectoral fins, deviate less than the
preceding families from the form of body typical of
the Sharks. A distinct expression of this is given by
the arrangement of the nostrils and the nasal valvule.
Each nostril is indeed continued here too by a groove
to the corner of the mouth; but the nostrils are farther
apart, and the nasal valvule coalesces throughout the
greater part of its inner surface with the bottom of
the snout, so that in many cases only its posterior
lateral corners form free dermal flaps.

The stronger flattening of the body, especially of
the head, imparts to the eyes, as we have mentioned
above, a more horizontal position; but they are pro-
tected above by an expansion of the frontal skin, and
to mitigate the excessive brilliancy of the light, the
upper margin of the iris is prolongated in most of the
species into finger-shaped processes partly covering the
pupil.

The family as a whole — it is dispersed round the
globe from the tropical to the frigid zones — is about
as varied in form as the preceding one, but in our seas
more so; and its members are often difficult to distin-

RAYS.

1101

guish. In manner of life they essentially resemble the
preceding Rays; but many of them descend to greater
depths. They are oviparous; but the embryo leaves
the egg very soon after the exclusion of the latter

from the cloaca, or even during its transmission through
the said passage.

Of the four or five genera which the family con-
tains, the Scandinavian fauna possesses only one.

Genus RAJA'k

The pectoral fins do not extend in front of the rostral cartilage. The ventral fins are deeply forked at, the hind
margin. Where a caudal fin is present , it does not extend to the under surface of the tip of the tail.

The foundation of our knowledge of the species
belonging to this genus and their mutual relations,
especially as they appear in the Scandinavian fauna,
was laid in its essential details by B. Fries6. His first
remarks had reference to the value of the specific char-
acters employed at his time within the genus.

The spiny armature of the body, he wrote, has
been the chief specific distinction employed since the
infancy of science, and it also affords beyond all question
not only clear, but also really trustworthy characters,
if only we refrain from minutiae, such as counting the
number of the spines, determining their positions and
size, as has hitherto been customary, for it is in this
very manner that most of the nominal species and the
constant confusion of the species arise. Every one who
studies the species by comparing a number of indivi-
duals, and who afterwards compares nearly related spe-
cies with each other, will find without fail that the
armature, however similar it may appear at first sight,
yet follows in each species a distinct development, char-
acteristic of the species, and expressed not only in the
form, size, position, growth, and shedding of the spines,
but also in such normal deviations from the original
specific type as age and, in part, the difference of sex
entail. For the attainment of descriptive lucidity Fries
proposed the following terminology. Smdtaggar ( spi -
nulce ) he called the prickles that cause the asperity of
the skin. They are quite small and short, either subu-
late, with a bifid or quadrifid, stellate base ( spinulce
stellares), or granular ( granuloses ). Knaggor ( aculei )
was the name he applied to the large, claw-shaped,
and usually recurved, thorn-like spines which stand as
wound-inflicting weapons partly at certain fixed spots on
the body (aculei or dinar ii), namely in rows along the spinal

column, on the rostral cartilage, around the orbital
margin, and on the dorsal side above the scapular re-
gion, partly at other undefined spots and in highly
variable number both on the dorsal side and the ventral
(aculei extra or dinar ii). Both these kinds of aculei vary
in number, form, and size, are shed periodically or
accidentally broken off, in which case, however, they
always leave, at least for a time, a mark in the skin.
Furthermore both may occur with expanded, flattened
or nail-headed base (aculei clavati), or with expanded,
conical base, deeply grooved on the sides and as it
were radiate (aculei radiati). Besides spinulse and aculei
the Rays have a third kind of spines, but only the
males. These Fries called the cards of the males (car-
mines maris). They are situated on the dorsal side of
each half of the body, partly at the outer margin of
the head, partly and principally on the pectoral fins,
a little way from the tips thereof. They consist of
rather long, simply bent or hooked, and very pungent
spines, set in rather irregular longitudinal rows, and
furnished with a mobile attachment, so that the Ray
can depress or erect them at will. When depressed,
they lie close to the skin, and may easily escape ob-
servation. By the distribution of the aculei on the tail
Fries distinguished between two types among our in-
digenous species, the one with an odd number of lon-
gitudinal rows, the other with an even number. The
latter type — as exemplified in the Shagreen Skate and
the Sandy Ray — seems hardly ever to be impaired by
exceptions or to alter with the age of the individual,
but is always recognisable by the symmetrical rows of
aculei (arranged on each side of the median line). The
former type, on the other hand, is often overlooked
and mistaken owing to the considerable modifications

“ Artedi, Ichthyol., Gen. Pise., p. 70.
b See Vet. Akad. Handl. 1838, p. 126.

“Est vocabulum Plinii.

Derivatio dubia” (Art., Phil., p. 73).

1102

SCANDINAVIAN FISHES.

induced by age. Judging by those species whose de-
velopment Fries had an opportunity of tracing, he
remarked, as a rule probably of general application,
that no species of Ray belonging to the type with an
odd number of rows of caudal aculei, when young or
newly hatched, has more than one such row, which
then invariably occupies the median line of the tail.
The lateral rows do not appear until the fish has
reached a more advanced age. Most of the Scandina-
vian species fall under this type, but. develop in a di-
rection involving something peculiar to and characteristic
of each. In some the lateral rows always project from
the side-margin of the tail, in others between this mar-
gin and the median line. In certain forms the spines
of the lateral rows never attain the same size and
strength as those of the median row, while in others
the relations are reversed. In some species the spines
of the median rows are persistent even at a very ad-
vanced age, in others they are normally lost or shed.

The form of the jaw-teeth, remarked Fries, has
been rejected as a specific distinction by Cuvier and
other writers, and on the whole with good reason, for
greater differences of dentition may often be observed
between the young and the old, between the male and
the female, than can be determined between two nearly
related species. But it is equally certain, he wrote,
that most of the species, not to say all the Scandinavian
ones, have a fixed form of dentition proper to them,
whereby some of them can safely be distinguished, and
some — e. g. the Thornback, Shagreen Skate, and Starry
Ray — are so well marked in this respect that they can
be confounded with no other species. To gain a right
conception of the dentition of each species, the deve-
lopment of the teeth must be carefully followed, and
a good clue is afforded to the investigator by a com-
parison of the teeth nearest the corners of the mouth
with those in the middle of the jaw. The former are
arrested, as it were, at a lower grade of development,
and most nearly resemble the teeth of the young; the
latter exhibit the highest stage in the dental develop-
ment of the species. The several intermediate rows
between the said points show the transitions from the
lowest to the highest development of the teeth. If at-
tention be paid to this, and the gradual detrition to
which the outermost (foremost) transverse rows in the
mouth are subjected be also taken into consideration,

the greater number of the said anomalies in the denti-
tion of a species disappear. As in the preceding fa-
mily, the males are characterized by more pointed jaw-
teeth than those of the females; but only in one of the
species indigenous to Scandinavia (the Thornback) is
the dentition of each sex thoroughly distinct; in the
others the difference is sometimes so slight that it has
even been overlooked.

The form of the snout affords characters of no
little importance and constancy, especially if the alte-
rations of growth be kept in mind ; and the variations
in the form of the snout are attended by differences in
the general configuration of the body, which form is
really determined by that of the snout and the pectoral
fins. To find a safe expression for these variations in
the form of the snout, Fries compared the length of
two lines, one drawn right across the head through the
centre of the pupils, the other at right angles to this
and extending to the tip of the snout. The species in
which the latter line is less than half as long as the
former, he called blunt-snouted; those in which the
length of the snout is more than half or at least half
the breadth of the head at the said point, he ranged
among the sharp-snouted Rays (the Skates). Another
expression of this, which besides affords an often re-
quisite character from the under surface of the head,
may be obtained by comparing the least width between
the inner margins of the nostrils (the base of the nasal
valvule) with the distance between the nostrils and the
tip of the snout, whereby we find that in the blunt-
snouted species the length of the base of the nasal
valvule is more, in the sharp-snouted species less, than
70 % of the distance between the nostrils and the tip
of the snout. The form of the snout also supplies an-
other character, which was indeed remarked by Fries
in his descriptions, but has won greater recognition in
more recent times". In the Rays with a very pointed
snout a line drawn from the tip of the snout to the
anterior margin of the outer tip of the pectoral fin falls
entirely outside the disk, whereas in the blunt-snouted
Rays it at least partly cuts the same.

From the position of the dorsal fins with relation
to each other Nilsson6 deduced a specific character the
value of which was also appreciated by Fries. Some
species, such as the Starry Ray, have the two dorsal
fins sel quite close together on the tail and without

a See E. Mokeau and D6derlein.
b Prodr. Ichthyoi. Scand., p. 119.

KAYS.

1103

aculei between them. In others, as in the Thornback,
the two tins are farther apart, and the median line
between them is armed with one or more spines.

Of the coloration of the body Fries observed that,
in spite of its variability, it is partly characteristic in
some degree of different species — especially that of the
under surface. As a natural result of the life led by
the Rays, which are bottom-tishes, this side of the body
is commonly light and colourless (whitish), and it would
appear as yet to be uncertain whether the dark colour,
entire or distributed in large spots, where it occurs on
the ventral side, affords trustworthy specific characters
or not. But good characters may be drawn from the
sometimes distinctive coloration of the long winding
canals and apertures of the ventral system of the lateral
line, and these characters are of so much greater utility
as being present from earliest youth and persistent in
specimens either stuffed or preserved in spirit. The
spots and ocelli which occur on the dorsal side of certain
species, showing endless variation in size, colour, and
extent, are utterly ..worthless as specific characters.

With regard to the external differences of sex,
which have long been known, Fries stated that they
are, as usual, less marked in young specimens than in
mature individuals, most of the male attributes being
commonly absent or only rudimentary in the former,
and developing in proportion as the age of puberty is
approached. In general the females are armed with
far more numerous and much larger spines than the
males; but this rule is not without exceptions.

Most of the true Rays are good and esteemed food-
fishes. When boiled fresh, their flavour calls to mind
that of the Halibut, or is not unlike that of lobster.
They may also be dried and kept till required for use,
in which case the fish is scored along the body. In this
form they are usually eaten after having been lyed in
the same manner as stock-fish. They are consequently
sought after by the fisherman, and they are in general
well distinguished and familiar to all, their vulgar
names affording valuable aid in the definition of the spe-
cies and their synonymy. Hollberg (1819) knew three
Scandinavian species of the genus, the klorocka ( Raja
radiata), knaggrocka ( Raja clavata, which he called
prickrocka), and slatrocka (Raja batis). In his Prodro-
mus (1832) Nilsson characterized four, the above-men-

tioned species and the blagarnsrocka (R, Untea, which
he called R. fullonica). In 1838 Fries added two, the
gokrocka (R. fullonica) and plogjernsrocka (R. oxy-
rhynchus). To these six species Malm supplemented in
1857 the sandr ocka ( R. falsavela, which he called spdtt-
rocka)- and in 1881 Collett described as Scandinavian
Lilljeborg’s svartbuksrocka (R. nidrosiensis). All these
forms unquestionably belong to the Scandinavian fauna;
but Collett further established in 1878 another species,
Raja hyperborea, to which the said fauna can probably
lay claim. These species may be distinguished as follows:

A: Blunt-snouted. Internasal width more than
70 % of the distance between each nostril
and the tip of the snout: — Raj* clavatse.

a: Root of the aculei nail-headed. Distance
between each nostril and the tip of the
snout 15 % or less of the breadth of
the disk Raja clavata.

b: Root of the aculei grooved. Distance
between each nostril and the tip of the
snout at least 16 % of the breadth of
the disk.

a: Upper median line of the tail armed
with a series of aculei.
aa: Aculei in the median line of the

back and tail at most about 16- . Raja radiata.
bb: Aculei in the median line of the

back and tail at least about 24__ Raja hyperborea.
(3: Upper median line of the tail with-
out aculei' Raja falsavela.

B : Sharp-snouted. Internasal width less than
70 % of the distance between each nostril
and the tip of the snout: — Rajse lteves.

a: Ventral side light, plain, without dark
punctuation.

a: Upper median line of the tail with-
out aculei Raja fullonica.

(3: Upper median line of the tail arm-
ed with aculei _ Raja lintea.

b: Ventral side dark or marked with small
dark dots and streaks.
a: Length of the snout less than 18 %
of that of the body or than 28 %

of the breadth of the disk Raja batis.

(3: Length of the snout more than 18 %
of that of the body o.r than 28 %
of the breadth of the disk.
aa: Aculei (even if small) at the
eyes. Ventral side blackish and

with indistinct punctuation Raja nidrosiensis.

bb: No aculei at the eyes. Ventral
side comparatively light and
with distinct dots and streaks.- Raja oxyrliynchus.

Scandinavian Fishes.

139

1104

SCANDINAVIAN FISHES.

Sectio I: Rajse clavatse, Rays. Snout short, its tip Hunt, the length of the snout from its tip to the
centre of the eyes being less than half its breadth at the latter point.

These species are the smallest, or at least do not attain the same magnitude as the Skates. The females
are larger than the males and more plentifully armed both with spinuhe and aculei. Spines of the latter kind,
known by Fries as extraordinary aculei, occur, sometimes in considerable number, scattered here and there on
the body without definite arrangement and often without symmetry. Iris furnished with lobate lid overlapping
the pupil (Fries).

THE THORNBACK (sw. knaggrockan).

RAJA CL A VAT A.

Plate XL VII, figs. 1 and 2.

Length of the snout ( from the anterior margin of the eyes) about 12 — 13 % of that of the body or 17 — 19 %
of the breadth of the disk. Distance between each nostril and the tip of the snout at most about 15 % of the
said breadth. Dorsal fins well separated, and the median line between them usually spinigerous. Aculei ungrooved,
their number in the median line of the back and tail in front of the first dorsal fin about 26 — 30 %a.

Syn. Raia clavata, Rondel., De Pise., lib. XII, cap. XIII, p. 353;

Schonev., Ichthyol. Slesv. Hols., p. 58; Willughb., Hist. Pise.,
p. 74. Raja aculeata, dentibus tuberculosis, cartilagine trans-
versa in ventre, Art., Ichthyol., Gen., p. 71; Synon ., p.
99; Spec., p. 103; Lin., Fna Suec., ed. I, p. 100; It.
Westrog., p. 175.

Raja clavata, Lin., Syst. Nat., ed. X, tom. I, p. 232; Bl.,
Fische Deutschl., pt. Ill, p. 65, tab. LXXXIII (9 + Raja
rubus, p. 67, tab. LXXXIV, <p ); Ketz., Fna Suec. Lin.,
p. 304; Hollb. (p. p.), Gbgs Wett., Witt. Sainh. N. Handl.,
Ill (1819), p. 29 (nec fig.); Fab., Tidskr. Naturv. Kbhvn, vol.
V (1828), p. 246; Yarr., Brit. Fish., ed. I, vol. II, p. 436;
Fr., Vet. Akad. Ilandl. 1838, p. 140; Bonap. ( Dasybatis ),
Iconogr. Fna Ital., Pesci, tab. 149, figg. 3 et 4 (+ D. aspera ,
ibid.); Ekstr., v. Wr. (Raja), SJcand. Fisk., ed. I, p. 154,
tab. 35; Mull., Hle, Syst. Beschr. Plagiost., p. 135; Kr.,
Damn. Fiske , vol. Ill, p. 962; Nilss., Skand. Fna, Fisk.,
p. 735; Couch, Fish. Brit. Isl., vol. I, p. 99, tab. XXII;
Gtiir, Cat. Brit. Mus., Fish., vol. VIII, p. 456; Coll.,
Forh. Vid. Selsk. Chrnia 1874, Tilliegsh., p. 214; N. Mag.
Naturv., Bd 29 (1884), p. 118; Malm, Gbgs, Boh. Fna,
p. 606; Winth., Naturh. Tidskr. Kbhvn, ser. 3, vol. XII,
p. 60; Mor., Hist. Nat. Poiss. Fr., tom. I, p. 391; Mob.,
Hcke, Fisch. Osts., p. 154; Storm, N. Vid. Selsk. Skr.,
Trondhj. 1883, p. 45; Day, Fish. Gt. Brit., Irel., vol. II,
p. 343, tab. CLXXI; Doderl., Man. Ittiol. Medit., fasc.
Ill, p. 176; Lillj., Sv., Norg. Fisk., vol. Ill, p. 555;
Carus, Prodr. Fnae Medit., vol. II, p. 521.

Raja scabra, Lin., Mus. Ad. Frid., tomi II:di prodr., p. 52
(sec. specimen ex Musseo Drottningholmensi).

Raja Dasybatus punctatus, IIollb., Gbgs Wett., Witt. Samh.
N. Handl., pt. IV (1821), p. 25 cum tab.

Raja pontica, Pall., Zoogr. Ross. Asiat., tom. Ill, p. 58; Rathke,
Mem. div. sav. etrang. Acad. Sc. Petersb., tom. 111(1837), p. 309.

Raja Rubus, Nilss. (ex Bloch), Prodr. Ichthyol. Scand., p. 118.

The males attain a length of about 7 dm. from the
tip of the snout to the end of the tail; the females
sometimes measure 9 or 10 dm. Until they have reach-
ed or nearly reached this size, they are apparently
incapable of propagating their species; Fries at all events
never found an individual under 6 dm. in length with
the external or internal genital organs developed.

The disk is rhombic, with somewhat pointed pec-
toral and ventral fins. The anterior side-margins of the
body form two bends, the anterior situated just behind
the tip of the snout, where a small, blunt muzzle is
thus produced, the posterior beginning a little in front
of a transverse line drawn through the centres of the
eyes. The posterior side-margins are more convex, but
in their outer parts curve slightly inwards. The great-
est breadth of the disk is on an average'' 69 % of the
length of the body. The distance from the tip of the
snout to the extreme end of the hind margin of the
pectoral fin (the length of the disk) is on an average"
75 %, to the anterior angle of the cloaca 64 %d, and
to the tip of the posterior ventral lobe 84 %e , of the
breadth of the disk.

a Provided that the series be complete, no aculei having been lost.

b In 1 1 specimens 36 — 92 cm. long. The variations run between 65 '/2 % and 721/., /•
c The variations run between 71 % and 79 %.
d ,, ,, ,, ,, 6lV2 % and 68 %.

e ,, ,, ,, ,, 79 % and 91 "/.

THORNBACK.

1105

The length of the head from the tip of the snout
to the occiput is about V5 of the entire length of the
body. The length of the snout varies between 12 and
13 % (exceptionally 14 %) of that of the body, or be-
tween 17 and 1 9 1/2 % of the greatest breadth of the
disk. The least interorbital width of the concave fore-
head (exclusive of the dermal fold above the eyes) va-
ries between 30 and 37 % (exceptionally 39 %) of the
length of the snout. The longitudinal diameter of the
oblong eyes is about 1/3a of the length of the snout,
and the longitudinal diameter of the spiracles, which
lie just behind the eyes, is in young specimens only
about % of that of the latter, in old Thornbacks almost
equal to the same. The internasal width is in young
specimens about 3/4, in old 9/io (exceptionally 95 %), of
the distance between each nostril and the tip of the
snout, or about 18 — 20 % of that between the cloaca
and the last-mentioned point. The lateral flaps of the
nasal valvule are fimbriated behind. The breadth of the
mouth, which is transverse and but slightly curved, is
equal in young specimens to the internasal width, some-
what greater in old. The dentition differs considerably
in the young and old and in the males and females.
All young specimens have small, blunt, granular teeth,
set in rows running obliquely on each side, and without
any distinction between the sexes (fig. 315, A); but as
the males approach maturity, the crowns of the teeth
become more and more flattened, with the hind margin
pointed and prolongated, a transition to the characteristic
dentition of the old males. The teeth of the adult males
are set in fairly regular rows, both longitudinal and
transverse, and those forming the 14 — 22 rows in the
middle of the mouth have a somewhat prolongated crown
with subulate cusp (fig. 315, B), which is directed inwards
towards the pharynx, thus lying like a scale over the
tooth behind it. The rows nearest to the corners of
the mouth have quite small, flat crowns, with the hind
margin slightly prolongated into a flat, sharpened heel,
which is more and more extended and approximated to
the subulate form, the nearer the row to which it be-
longs lies to the first-mentioned median rows. Only
the teeth with subulate cusp have a very narrow rim at
the base, and as these are most exposed to detrition,
the tips of the foremost (oldest) teeth are always worn
and blunt. In old females the teeth have broad, lo-
zenge-shaped, and sloping crowns, set in pentagons

(so-called mosaics, fig. 315, C ). This dentition is pe-
culiar to the Thornback, and cannot be confounded
with that of any other among the Scandinavian Rays.

The whole upper surface of the body is densely
strewn with fine, compressed spinulae, branching at the
root into a star with two — four or even six arms.
Similar small spines, more scattered, however, are set
on the ventral side of the body in full-grown speci-
mens, especially under the snout and tail, but are
wanting on the pectoral and ventral fins, which are
smooth. In young individuals, on the other hand,
especially of the male sex, the whole ventral side seems
smooth to the touch; but on careful examination very
fine spines are detected even there, at least under the
very tip of the snout and at the sides of the abdominal
cavity. All the aculei belong to the type which, when

D E

Fig. 315. Jaw-teeth and aculei of the Thornback ( Raja clavata).
A, teeth of a young male, natural size, and (below) a tooth magnified,
lateral and superior aspects; B , tooth of an old male, magnified,
lateral and superior aspects; C, tooth of an old female, magnified,
aspects as before; D , dorsal aculeus of an old Thornback; E , caudal
aculeus of a younger specimen.

fully developed, has a smooth, expanded base ( aculei
clavati). The ordinary consist in young specimens of

1) a row running along the back, containing some
twenty spines, with base as yet but conically expanded,

2) an aculeus on the scapular cartilage, on each side
of this row, 3) one or two before and two or three
behind the eyes, and 4) a few quite small aculei on
the rostral cartilage, set in two rows. In older speci-
mens we find, besides those enumerated above, a row
on each side of the median series at the root of the
tail, and sometimes, especially in the females, two rows
at the same point, containing aculei of the same size
as or even larger than those of the median series.

In a young specimen 14 cm. long the diameter of the eyes, according to Kkoyee, was 37,/‘> / of the length of the suout.

1106

SCANDINAVIAN FISHES.

These lateral rows do not occupy the extreme side-
margin of the tail, but the space between this and the
median series, except where there are two lateral rows
on each side, in which case the outer row projects from
the said margin. They do not extend throughout the
length of the tail, but only a short distance along it; they
are not symmetrical, and contain an extremely variable
number of aculei. A little behind the termination of
these lateral rows and before the origin of the first
dorsal fin, we usually find in old specimens 1 — 3 true
lateral aculei (set at the extreme side-margin), in
most cases the largest on the tail, seldom of equal
number on both sides. These few lateral aculei at the
said point and those occupying the space between the
two fins are, though not constant, good characters,
where they are present, of the Thornback. The extra-
ordinary aculei are so variable, both in distribution
and number, that no rule can be stated. They seem-
ingly appear a short time before the period of puberty,
for in all young specimens there is not even a sign of
their presence. They are most numerous in middle-
aged females, but in very old ones most of them have
again disappeared". They are extremely seldom sym-
metrical, sometimes preponderating in number on the
dorsal side, sometimes on the ventral; sometimes scat-
tered, sometimes collected in groups. Their basal scutes
are often of considerable extent. Among the Scandi-
navian species of the family the Thornback is the only
one in which aculei appear on the ventral side.

The coloration of the dorsal side is yellowish gray,
with a number of large, round, whitish yellow spots of
varying size and without symmetry. Between these
appear smaller, black spots, like round dots, which
gather round the former and compose black frames,
sharply defined especially in young specimens. Some-
times there also occur in old individuals large ocelli, cir-
cular black spots with light margin, in one or two pairs
on the dorsal side. The ventral side is white, with a
dash of violet round the margins of the fins. On the
tail the dark colour now and then, especially in young
specimens, forms several (up to 6 — 8) broad transverse
bands, most distinct on the under surface thereof.

The sex is manifested externally, in part and with
most clearness, by the large pterygopodia of the male
and the cards of spines peculiar to him on the upper

surface of the pectoral fins — two patches of erectile
spines on the sides of the orbital region and another
pair of larger patches within the outer angles of the
fins — in part by several other differences already no-
ticed. However distinct all these characteristics may
be at an advanced age, during youth they are wanting
or only subindicated, and at first the sexes are exter-
nally indistinguishable. As the time of puberty ap-
proaches, they all appear almost simultaneously, and
afterwards the difference between the sexes grows more
marked year by year. In a male 216 mm. long (pre-
sumably a year old) taken in August, Malm estimated
the length of the pterygopodia at only 3 mm. In a
male about 42 cm. long Fries found the pterygopodia
still quite small and short, their tips extending only a
little more than half-way along the ventral fins; there
were no signs of cards; the teeth were still of a gra-
nular, lozenged and blunt form; the ventral side of
the body was quite smooth, only a few spots (the tip
of the snout and the sides of the belly) being rough
to the touch ; there were no extraordinary aculei, and
of the ordinary caudal aculei only the median row was
present. In a specimen 52 cm. long the circumstances
were essentially the same, only that the pterygopodia
were somewhat longer in relation to the ventral fins,
and the ventral side of the body was rougher, especially
under the tail. But in an individual rather more than
60 cm. long the pterygopodia are so developed that
their tips extend more than half-way along the tail,
and all the other characteristics of the male have ap-
peared, though the cards are still small, containing
few spines, and these set in 2 or 3 rows, whereas in
larger and older specimens as many as 6 — 8 rows
may be counted, each containing some twenty spines
or more.

The Thornback is the commonest of all the Swe-
dish Rays. In the Sound it occurs southwards to the
shallows off Saltholm (Winther). It has occasionally
been met with in the south-west of the Baltic, on the
east coast of Schleswig-Holstein (Schonevelde and
Mobius and Heincke) and the north coast of Mecklen-
burg (Boll*). It is most plentiful in the Cattegat
and the North Sea. To the north, according to Storm,
its range probably does not extend beyond the neigh-
bourhood of Trondhjem. To the south it penetrates into

° Fries met with very old males as well as females destitute of a single extraordinary aculens.
6 Die Ostsee, Arch. Ver. Fr. Naturg. Meckl., Heft. I, p. 89 (sep.).

THORNBACK.

1107

the Mediterranean and the Black Sea (Pallas), and
it has been found off Madeira (Gunther).

In the island-belt, of Bohusl&n Thornbacks both
young and old are taken all the year round. During
winter, however, they are less common, and the fisher-
men state that at this season the Thornback repairs to
deep water. In spring and summer it is very plentiful
off the coast, where it seems to prefer water of a mo-
derate depth a and with a sandy bottom. It lives almost
invariably at the bottom and extremely seldom ascends
to a higher level. Its food consists of small fishes,
crustaceans — crabs, lobsters, shrimps, etc. — mollusks,
Annelids, Echinoderms, and Actinice \

Throughout the summer the females contain deve-
loped ova; but according to Fries only one egg is laid
at a time, and the period of oviposition for each female
must consequently be of long duration. Fries observed
that, on opening old females in summer-time, the egg-
clusters in the ovaries are found to be considerably
tumid, the several eggs showing different grades of
development, from the size of a pea to that of a plum.
In the two oviducts however — each of them dilated
below into a uterine organ — he never found more than
a single egg, and this already invested with the sin-
gular corneous shell. When one of the uteri contained
an egg, the other was empty. The ovum is of the form
normal among the Rays, rectangular, with the four
corners prolongated in its longitudinal direction, and
according to Malm0 is 55 — 57 mm. long and 40 mm.
broad, but including the filaments at the corners 110
— 115 mm. long. Fries once found a female Thorn-
back with a half expelled egg in the cloaca. On ex-
amination the protruding end of the egg proved to be

open, the larva having already left the shell. He dis-
sected the specimen, and saw that the left uterus was
empty, but that in the right there lay a recently de-
scended ovum, the shell of which was still very soft.
In this case it was consequently evident that the em-
bryo had freed itself from the egg immediately on the
expulsion of the latter. Malm, on the other hand,
never found a viable foetus, scarcely a distinct embryo,
in newly deposited ova, whence he inferred that the
development of the embryo does not commence until
the egg is laidfi The process of development may thus
vary in its acceleration; but by far the greater number
of the eggs found after liberation are empty. During
their early life, according to Kroyer, the fry keep to
the shallows of shelving coasts.

The Thornback is taken on long-lines and in Floun-
der-nets, less frequently in the seine. As an article
of food it enjoys a good reputation in Great Britain
and Ireland, especially in the latter country0; but
in the island-belt of Bohusl&n, according to Ekstrom,
it is regarded as one of the poorest fishes. It is hardly
ever eaten fresh there, at all events by the islanders,
who instead dry the flesh in the open air on flakes
( gdllar ) or platforms, constructed in elevated situations.
After drying it is sold to the peasants of the agricul-
tural districts, who use it as a substitute for lutfiskf at
their Christmas festivities, and consider it good eating.

Besides knaggrocka (Spiny Ray) this species bears
in the island-belt of Bohuslan the names of piggrocka,
as adult, and rockhok and peruk , while young. On the
coast of Scania it shares with the following species the
name of torr-borr, according to Lilljeborg a corruption
of the Danish Tcerbe. (Fries, Ekstrom, Smitt.)

a According to Malm the full-grown Thornback is only seldom found in less than. 14 fathoms of water.

6 See Olsson, Fiskarnes foda, Lunds Univ. Arsskr., tom. VIII, 1871.
c Ofvers. Vet. Akad. Forh. 1876, No. 3, pp. 94, 95.
d Gbgs, Boh. Fauna , p. 607.
e See Day, 1. c., p. 345.

f Dried Cod etc. soaked in lye before cooking. This is still a staple course in Sweden at Christmas and on other feast-days. Tr.

1108

SCANDINAVIAN FISHES.

THE STARRY RAY (sw. klorockan).

RAJA RADI AT A.

Plate XLVII, fig. 3.

Length of the snout ( from the anterior margin of the eyes) about 13 or 14 % ( in one year old specimens about
12 %) of the length of the body or about 19 — 21 % of the breadth of the disk. Distance between each nostril
and the tip of the snout about 17 or 18 %a of the said breadth. Dorsal fins set close together , and the median
line between them usually without aculeus. Aculei grooved in a stellate form at the base , their number in the

median line of the back and tail at most about 16.

Syn. Raja clavata, Olafs., Reise Isl., pp. 359 et 987, tabb. XLIX
et L; Hollb. (p. p.), Gbgs Wett., Witt. Sarah. N. Haudl.,
pt. Ill (1819), p. 29 cum tabb.; Pall., Zoogr. Ross.
Asiat., tom. Ill, p. 58; Nilss., Prodr. Ichthyol. Scand.,

p. 119.

Raia fullonica, Fabr., Fna Groenl., p. 125; Faber, Tidskr.
Naturv. Kbhvn, vol. V (1828), p. 246; Id. (p. p.), Fisch.
Isl., p. 38.

Raja radiata, Donov., Brit. Fish., tab. CXIV; Yarr., Brit. Fish.,
ed. I, vol. II, p. 439; Fr., Vet. Akad. Ilandl. 1838, p.

146; MOll., Hle, Plagiost., p. 137; Sundev., v. We., Skand.

Fisk., ed. I, p. 178, tab. 43; Kr., Damn. Fisk., vol. Ill,
p. 939; Nilss., Skand. Fna, Fisk., p. 736; Mgrn, Finl.
Fiskfna, p. 72; Gthr, Cat. Brit. Mus., Fish., vol. VIII,

p. 460; Garm., Proc. Dost. Soc. Nat. Hist., vol. XVII

(1874), p. 177; Coll., Forh. Vid. Selsk. Chrnia 1874,
Tillfegsb., p. 214; 1879, No. 1, p. 105; Norsk. Nordh.
Exped., Zool., Fisk., p. 14; N. Mag. Naturv. Chrnia, vol.
29 (1884), p. 118; Cederstr., Ofvers. Vet. Akad. Forh.
1876, No. 4, p. 67; Malm ( Amhlyraja ), Gbgs, Boh. Fna,
p. 607; Winth. (Raja), Naturh. Tidskr. Kbhvn, ser. 3,
vol. XII (1879), p. 60; Mor., Hist. Nat. Poiss. Fr., tom.
I, p. 394; Mela, Vert. Fenn., p. 368, tab. X; Mob.,
Hcke, Fisch. Osts., p. 153; Day, Fish. Gt. Brit., Irel .,
vol. II, p. 347, tab. CLXXIII; Lillj., Sv., Norg. Fisk.,
vol. Ill, p. 547.

The Starry Ray is the smallest Scandinavian spe-
cies of the genus. At a length of 472 dm. it has
attained maturity, and 6 dm. would seem to be its
maximum length, v. Weight’s figure represents a fully
developed female in which the length of the disk was
253 mm., that of the tail 192 mm., the greatest breadth
309 mm., and the distance between the mouth and the
tip of the snout 67 mm. In this species too the males
are commonly smaller than the females.

The disk is almost square, but has strongly rounded
angles, especially the posterior, which is formed by the
very long, broad, and uniformly rounded posterior lobes
of the ventral fins. The anterior side-margins of the

disk show a slight, A-shaped curvature, with broader
and more rounded snout than that of the preceding
species, and from the tip of the latter projects a very
small, blunt muzzle. The posterior side-margins may
almost be described as rounded. The greatest breadth
of the disk is on an average rather more than 3/5 (58
— 64 %), and its length somewhat more than 1/2 (50
— 55 %), of the length of the body, the latter thus
measuring 86 — 88 % of the former. The average dis-
tance from the tip of the snout to the cloaca is rather
more than 7/10 (70 — 75 %) of the greatest breadth of
the body, and the length to the hind margin of the
I ventral fins is almost equal to the said breadth.

The length of the head is here too about 1/5 (18
—21 %) of that of the body, and that of the snout
varies, as in the preceding species, between about 12
and 14 % of the latter, but here between about 1 9 x/2 and
21 % (exceptionally 23 %) of the greatest breadth of the
disk. The interorbital breadth of the forehead and the
dimensions of the eyes and spiracles are about the same
as in the preceding species. In the position of the
nostrils too there is little difference from the Thorn-
back, the distance between them being about 8/i0 — Vio
of that between each of them and the tip of the snout
or 21 — 17 % of that between the cloaca and the last-
mentioned point. The breadth of the mouth is here too
in the young somewhat less than or equal to the inter-
nasal width, in the old somewhat greater. The denti-
tion is very nearly alike in both sexes. The teeth (fig.
316) are small and pointed, set in about 36 rows along
both jaws, the largest in the middle, both the size and
number showing diminution towards the corners of the
mouth. Each tooth consists of a rounded basal disk,

Sometimes, according to Kroyer, 15’/2 %.

STARRY RAY.

1109

somewhat concave in the middle, from the hind margin
of which a line, subulate cusp rises obliquely upwards
and backwards.

Only the upper surface of the body is armed with
spines, the under being quite smooth and soft. The
spinulae are of the same form as the aculei scattered
over the whole surface, having a conically expanded,
stellate base and an extremely tine, pungent tip. The
only difference between them is in the size, though no
such definite line as in the Thornback can here be
drawn between the spinulae and aculei, large and small
of all sizes being indiscriminately interspersed with one
another, and their distribution being more scattered and
irregular, leaving more or less extensive patches of
smooth skin between them. The ordinary aculei occupy
in young specimens the same positions as in young
Thornbacks, and stand in nearly the same relation to
one another; but their distribution is characteristic in
three respects: there is no aculeus between the two
dorsal fins“, which are set so close together that the
membrane of the first often overlaps the anterior mar-
gin of the second; the aculei along the spinal column
are only 12 — 16 in number; and the scapular cartilage
is always furnished with two aculei, one behind the
other, and sometimes with a third, between this pair
and the median series, all with deeply grooved, stellate
base. In old specimens there further appears on each
side of the median series a lateral row, commencing
high up on the dorsal surface not far behind the sca-
pular cartilage, and extending back to the first dorsal
fin. On the tail these lateral rows lie about half-way
between the median series and the side-margin of the
body, and here the aculei never attain the same size as
those of the median series. The lateral margins of the
tail are never spinigerous. The extraordinary aculei
appear, as we have mentioned, only on the dorsal side.
They gradually increase in size, but never attain the
same dimensions as the ordinary aculei on the spinal
column. They are scattered with some degree of sym-
metry, and are never wanting in adult individuals.

Characteristic as the radiate base of the aculei
generally appears in this species, it should be remarked,
however, that during earliest youth, in individuals 9 —
12 cm. long, the base is quite smooth, and the anterior
aculei rise to a height of 5 mm. in a subulate form.
Even at the former length of body all the ordinary

a In adult Starry Rays from the Arctic regions, however,
specimens from the White Sea there is no aculeus between these fins

aculei are already present, and their length as just-
stated is very considerable in proportion to the size of
the body, as compared with their dimensions in older
specimens, this being a sufficient character by which
small young individuals of the two species may im-
mediately be distinguished.

The coloration is above of a plain liver-brown,
faintly marbled with yellowish white, and thinly strewn
with irregular, very indistinct, blackish spots of small
size; but the large, round, whitish spots that appear
in the Thornback are entirely absent. The spinulae
and aculei are of a paler, yellowish white colour. The
whole under surface is white, somewhat pinkish at the
margins of the fins, the canals and pores of the lateral
line being colourless. Kroyer remarks, however, that
black spots, varying in number and size, frequently
occur on the ventral side, especially on the under sur-
face of the tail. Transverse bands may also be observed,
but are less numerous than in the preceding species.

A B

Fig. 316. Jaw-teeth and aculei of a Starry Ray ( Raja radiata ), na-
tural size. A: teeth of upper and lower jaws; a, tooth, magnified,
lateral and superior aspects; B: dorsal aculeus, lateral and superior
aspects, natural size; a , aculeus from the tail of a young specimen,
lateral and superior aspects, magnified.

The external difference of sex is less marked in
young individuals of this species than in young Thorn-
backs. In adult Starry Rays it is distinctly shown by
the long pterygopodia of the male and by the posterior
cards at the outer angle of the pectoral fins. The an-
terior cards, on the other hand, at the side-margins of
the head, may be regarded as wanting, for the female,
which generally has more numerous spines, is, if any-
thing, better armed at these spots than the male.

The Starry Ray belongs to northern latitudes, and
occurs far up in the Arctic Ocean. From Greenland
it was described by Fabricius. The Norwegian North
Atlantic Expedition of 1876 — 78 found it off Bear Is-
land and the north-west corner of Spitsbergen. In the
White Sea it was known even to Pallas. In the Bay
of Biscay, according to Moreau, lies the southern limit

Collett frequently found one aculeus between the dorsal fins. In our

1110

SCANDINAVIAN FISHES.

of its geographical range, but its occurrence is rare
even off the north coast of France. It penetrates into
the Sound and, through the Belts, into the western-
most part of the Baltic in the same manner as the

After Collett.

Fig. 317. Raja hyperborea, <y, 1/3 nat. size.

Thornback, but is somewhat commoner there, though
less numerous on the coast of Bohuslan than the latter.
Equally extensive is its bathymetric range. In Scan-
dinavian waters it generally lives at a less depth than
the Thornback, but appears also to descend to far lower
levels. Storm obtained a specimen from Trondhjem
Fjord that had been taken in 250 fathoms of water,
and the above-mentioned Norwegian Expedition secured
their two young specimens from Spitzbergen at a depth
of 459 fathoms.

In its manner of life the Starry Ray seems else
to resemble the preceding species; but it is smaller,
and is therefore restricted to smaller prey, preferably
small fishes, comparatively minute crustaceans (Amphi-
pods), and worms. “Early in spring (in the month of

March)”, says Kroyer, “it ascends towards the shal-
lows; it appears to lead a secluded life during a great
part of the summer; and at the approach of winter it
returns to the depths on the completion of its spawn-
ing operations.” Its egg-shells, he says, are about 61
mm. long, exclusive of the prolongations at the four
corners, and 39 mm. broad. The fry seem to remain
not far from land during winter, for the small young
specimens mentioned above, whose peculiarities were
remarked by Fries, were caught at this time of year
in Herring-seines.

In Scandinavia the Starry Ray is of less im-
portance than the Thornback, partly Giving to its smaller

DEEP-SEA RAY.

1111

size, partly because it is less common. It is fre-
quently taken, however, on Haddock- lines and in Floun-
der-nets, occasionally too in seines drawn for Herrings.

(Fries, Smitt.)

Fig. 318. Raja hyperborea, cA nat.

size. After Gunther.

THE DEEP-SEA RAY.

RAJA HYPERBOREA.
Figs. 317 and 318.

Length of the snout ( according to Gunther’s figures ) about 13 — 14 % ( according to Collett 17 %) of that of
the body, or 17 — 18 % ( according to Collett 22 %) of the breadth of the disk. Distance between each nostril
and the tip of the snout about 14 — 16 % of the said breadth. Dorsal fins set close together, with (in the young)
or without (in the old) interjacent aculeus. Aculei grooved , their number in the median line of the back and

tail at least about 24a.

Syn. Raja hyperborea , Coll., Forh. Vid. Selsk. Chrnia 1878, No. 14,
p. 7 ; Norsk. Nordh. Exped., Zool., Fisk., p. 9, tab. I,
figg. 1 et 2 ; Gthr, Deep Sea Fish., Challeug. Exped., Zoo].,
vol. XXII, p. 8, tab. IV; Lillj., Sv., Norg. Fisk., vol. Ill,
p. G04.

“ One of Gunther’s young specimens, however, had only 16.

As may be inferred from the diagnosis, as well as
from the figures given above, it is as yet uncertain
whether this Ray should be regarded as a distinct spe-
cies or merely as a variety (perhaps two different va-

Scandinavian Fishes.

140

1112

SCANDINAVIAN FISHES.

rieties) of the common Starry Ray, to which it is also
similar in size. Only six specimens are known, five
of them — including a male 62 cm. long — taken by
the Knight Errant Expedition at a depth of 400 — 600
fathoms between Scotland and the Faroe Islands and
now in the British Museum, the sixth a male 52 cm.
long taken by the Norwegian North Atlantic Expedition
in 459 fathoms of water, off the north-west corner of
Spitsbergen, and now at Christiania.

From the common Starry Ray it differs principally
by the greater number of aculei in the median line of
the back and tail — the ventral side is smooth and
quite free from aculei — and by the maculation of the

THE SANDY RAY (sw.

RAJA FA
Fig.

Length of the snout about 10 — f/1/ 2 % of that of the
between each nostril and the tip of the snout about 16-

No row of aculei in th

Fig. 319. Young female of the Sandy Ray ( Raj

Syn. Raja rulms , Lacep. (p. p.; nec Bl., Gmel.), Hist. Nat. Poiss.,
tom. I, p. 79, tab. 5 (fig. .vera, descr. p. p.).

Sandy Ray , Couch, Mag. Nat. Hist. (Charleswouth), vol. II
(1838), p. 71 (sine nom. system.).

Raja radula, Yarr., Brit. Fish., ed. I, vol. II, Suppl., p. 69
(nec De La Roghe).

Raja falsavela, Bnp., Iconogr. Fna Ital., Pesci , tab. 148, fig.
1; Canestr., Fna Ital., Pesci, p. 56; Gigl., Esp. Intern.

ventral side. This surface of the body is marked on
its white ground with large, almost symmetrical spots
and patches of the same dark grayish brown colour as
appears on the dorsal side. The species is stated to
be further distinguished by more scattered and more
pointed jaw-teeth. The form of the body differs so
widely in Collett’s and Gunther’s figures that in this
respect the Deep-sea Ray would seem to be a highly
variable intermediate form between the Starry Ray and
the sharp-nosed species of the genus. The species- — which
is based principally on a chromatic character — evidently
requires revision, a task which, having no specimen with-
in our reach, we cannot undertake in the present work.

SANDROCKAN OR SPATTIiOCKAN).

LSAA7ELA.

319.

body or 19 — 21 % of the breadth of the disk. Distance
17 % of the said breadth. Dorsal fins set close together.
' median line of the tail.

Pesca Berl. 1880, Sez. Ital., p. 114; D6derl., Man. Ittiol.
Medit., fasc. Ill, p. 202.

Raja ncevvs, Mull., Hle, Plagiost., p. 138 (+ R. radula ,
p. p., p. 133).

Raja circulars, Couch in Yarr., Brit. Fish., ed. 2, vol. II,
p. 574; Malm, Ofvers. Vet. Akad. Forh. 1857, p. 187;
Couch, Fish. Brit. Isl., vol. I, p. 115 (+ R. miraletus,
p. 112); Gthr, Cat. Brit. Mus., Fish., vol. VIII, p. 462;

SANDY RAY.

1113

Coll., Forh. Vid. Selsk. Clirnia 1874, Tillasgsh., p. 214;
1879, No. 1, p. 105; N. Mag. Naturv. Clirnia, vol. 29 (1884),
p. 119; Malm, Gbgs , Boh. Fna, p. 608; Mor., Hist.
Nat. Poiss. Fr., tom. I, p. 397 ; Day, Fish. Gt. Brit., Irel.,
vol. II, p. 348, tab. CLXXIV ; Lillj., Sv., Norg. Fislc.,
vol. Ill, p. 564; Carus, Prodr. Face Med.it., vol. II,
p. 521.

The Sandy Ray attains the greatest size among
our blunt-nosed species, the females growing to a
length of at least 12 dm. It also undergoes the greatest
alterations in its proportions, the young having com-
paratively a much longer tail than the old.

The form of the body is essentially the same as
in the case of the Starry Kay; but the disk is still
more rounded, and the root of the tail broader (flatter).
The breadth of the disk is in young specimens (less
than 4 dm. long) about 48 — 53 %, in old as much as
63 or 64 %, and its length in the former about 44 —
51 %, in the latter 52 — 54 %, of the length of the body,
the length of the disk thus varying between about 85
and 95 %a of its breadth. The distance from the tip
of the snout to the cloaca is about 3/4 of the greatest
breadth of the body, and that to the hind extremity
of the ventral tins about equal to the same.

The length of the head, which varies, according to
Doderlein’s measurements, between 25 and 38 % of
the breadth of the disk, is during youth, in consequence
of the great length of the tail, only about 15 % of that
of the body; subsequently, according to Doderlein, this
percentage increases to 20, while the relative length of
the tail is undergoing reduction, but again diminishes in
the old to 16 % of the length of the body. The length
of the snout is less in proportion to that of the body
than in any among the preceding species of the genus,
but in proportion to the breadth of the disk similar to
the same measurement in the Starry Ray6. The least
interorbital width of the forehead in this species too
is somewhat more than x/3 (about 36 °/o) of the length
of the snout, and the longitudinal diameter of the eyes
varies between about 3/ 4 and 2/3 of this breadth. The
width of the spiracles, which are directed rather straight
outwards, is in young specimens less, in old, according
to Doderlein, i/i greater, than the longitudinal dia-
meter of the eyes. The mouth and its dentition are
subject to considerable alterations of growth. In the
young the cleft of the mouth is much smaller than in

Starry Rays of the same size, but the teeth are similar
to those of the latter; in old specimens the mouth is
broad, and the teeth are sharp and unguiform, both in
females and males, while their number too shows con-
siderable increase with age. In a young specimen about
45 cm. long we find only 34 longitudinal rows of teeth
in the upper jaw, where Malm in older specimens
counted 78. The internasal width is in our young
specimens 71 — 72 % of the distance between either of
the nostrils and the tip of the snout, but only 15 —
16 % of that between the mouth of the cloaca and the
latter point.

Young specimens of the Sandy Ray too have only
the dorsal side armed with spines, the ventral being
perfectly smooth. In older specimens both the ventral
side and the dorsal are more or less densely coated
with spinulte throughout the greater part of the surface;
but certain patches, as, on the dorsal side, the hind
margin of the pectoral fins, the anterior lobe and the
base of the ventral fins together with an oblong patch
further in, and the greater portion of the median line
of the back and tail, and, on the ventral side, the outer
part of the pectoral fins, the whole of the ventral fins,
and the region between the latter pair, are smooth.
Along the anterior margin of the pectoral fins — in old
specimens both on the dorsal side and the ventral — as
well as on the sides of the tail and at its extreme lateral
margin — in young specimens, however, only to aline
with the beginning of the first dorsal fin, where the
lateral margins of the tail develop a broad dermal
fold — are set rows of larger spinulae, intermediate be-
tween spinulae and aculei. The true aculei, which here
too are grooved, though not so distinctly as in the
Starry Ray, have a broad conical base and a compress-
ed, recurved spine. They are set on the front of the
snout, in a curved row on the supraorbital margin, in
a triangular patch between the head and the scapular
cartilage, one or two at each end of the latter carti-
lage, in a row on each side of the median dorsal line
behind the scapular cartilage — sometimes, especially
in old specimens, this row commences further back or
even not before the tail — and in two to four more or
less irregular series on the sides of the tail. The me-
dian line itself is smooth, with the exception that,
owing to the irregularity prevalent in the distribution

° According to DSderlein’s measurements the variations run between 83 and 97 %.

6 In the Mediterranean falsavela, however, it is sometimes, according to DSderlein’s measurements, only about 16 — 17 % of the
breadth of the disk.

1114

SCANDINAVIAN FISHES.

of the spines, an aculeus or two may sometimes be
forced in even to the middle of the upper caudal
surface.

By the coloration Moreau distinguished between
three varieties: falsavela, with plain, brownish or red-
dish gray (clay-coloured) dorsal side; ncevus, with the
base of each pectoral fin spotted with a large ocellus,
the middle of which is black but crossed by rather
broad, yellowish white streaks; circularise with nume-
rous, but smaller, symmetrically arranged, simple spots,
light (cream- white) or dark, scattered on the dorsal
side of the disk. The ventral side is white.

The Sandy Ray has an extensive geographical
range, more southern and to the north more pelagic
than that of the Starry Ray. Its true home in the
Old World lies in the Mediterranean and the adjoining
parts of the Atlantic, but it is frequently found even
so far north as on the fishing-banks off the south-west
of Norway, where Malm was the first to observe it among
the take of fish at Jaderen. Even in the Skager Rack
it has been met with, but only once, so far as is known.
On the 12th of July, 1879, Theel and Forsstrand
took two specimens in a deep-sea trawl at a depth of
370 fathoms, off Arendal, on a bottom of fine, brown
clay. One specimen was a male 33 cm. long and not

yet arrived at maturity, the other a female 45 cm. in
length. Round the coasts of Great Britain and Ireland
the Sandy Ray would appear to be commoner (Day),
but in France it is stated to be rare (Moreau). Yet,
like the Starry Ray, it is apparently an inhabitant of
the West Atlantic too, where MitchiklV and StorerV
Baja ocellata and B. diaplianes from the east coast of
the United States can hardly be distinct in species from
our Sandy Ray. As Moreau has remarked, the Sandy
Ray was certainly known to Lacepede; but the name
he conferred upon it was incorrect and has been so
misused that it hardly deserves recognition. When the
synonymy of the species has been fully elucidated, the
point will be decided whether the systematic name em-
ployed in America has not most right to employment.

Where Malm first found the Sandy Ray, it had
been taken on long-lines in company with all the
other common Rays of Scandinavian waters, except
the Starry Ray; and its manner of life is probably
much the same as that of its congeners. As food, ac-
cording to Couch, it is little esteemed in England,
where it is mostly used to bait lobster-pots and crab-
pots. From spring till the end of autumn it may be
found on the English coast- in water of a moderate
depth; during winter it lives farther from land.

Sectio II. Rajae laeves, Skates. Snout long , tip of the snout acute-angled: length of the snout from the
tip to the transverse line through the body at the centre of the eyes more than half as long as this trans-
verse line.

These Rays possess so-called ordinary aculei only at the eyes and along the spinal column and tail, no
further ordinary aculei being scattered on the surface of the body. All have pointed teeth.

A: Ventral side of the body white, without spots. — White Skates.

These Skates have the two dorsal fins set close together, never with interjacent aculeus. Their under sur-
face is entirely free from spots, they have ordinary aculei both on the back and the tail, and the lobate pupil-
lid originating from the iris is absent. In size they are intermediate between the Baja clavatce and the true
Baja laves. They live at a great depth and seldom approach the coasts of Scandinavia. (Fries.)

a Trans. Lit., Phil. Soc. N. Y., vol. I, p. 477.

6 Mem. Amer. Acad. Arts, Sc., Boston, N. ser., vol. IX, ,p. 240, tab. XXXIX, fig. 1.

SKATES.

1115

THE SHAGREEN SKATE (sw. goicrocican).
RAJA FULLONICA.

Fig. 320 and Plate L, fig. 1.

Length of the snout about 12 — 14 % of that of the body or 20 — 24 % of the breadth of the disk. Distance
between each nostril and the tip of the snout about 17 % of the said breadth or nearly tivice ( 175 — 200 % of)
the internasal width. Aculei grooved. No aculei in the median line of the back and tail.

Fig. 320.

Shagreen Skate ( Raja fullonica ), 9? Vs na^- size- The Cattegat, July, 1835. B. Fries, a, a tooth, magnified, lateral and

outer aspects, b , a dermal aculeus.

Syn. Raja aspera nostras, the White Horse dicta, Willughb., Hist.

Pise., p. 78; Ray, Synops. Meth. Pise., p. 26. Raja toto
dorso aculeato, duplici ordine aculeorum in cauda, simpli-
cique ad oculos, Art., Ichtli., Gen. Pise., p. 72; Syn. Pise.,
p. 101. Shagreen Ray, Penn., Brit. Zool. (ed. 1776), vol.
Ill, p. 77.

Raja Fullonica , Lin., Syst. Nat., ed. X, tom. I, p. 231 ;
Rathke in Ascan., Icon. Rer. Nat., ad. tab. XLIII; Fr., Vet.
Akad. Handl. 1838, p. 150, tab. II, fig. 2, tab. Ill, figg. 5
et 9; Mull., Hle, Plagiost., p. 145; Kr., Damn. Fisk.,
vol. Ill, p. 996; Nilss., Skancl. Fna, Fisk., p. 737; Yarr.,
Brit. Fish., ed. 2, vol. II, p. 578; Gthr, Cat. Brit. Mus,,
Fish., vol. VIII, p. 467; Coll., Forli. Vid. Selsk. Chrnia
1874, Tillaegsli., p. 217; 1879, No. 1, p. 106; N. Mag.
Naturv. Chrnia, Bd 29 (1884), p. 119; Malm ( Leucoraja ),
Gbgs, Boh. Fna, p. 609; Winth. {Raja), Naturh. Tidskr.
Kbhvn, ser. 3, vol. XII, p. 61; Day, Fish. Gt. Brit., Irel.,
vol. II, p. 342, tab. CLXX; Lillj., Sv., Norg. Fisk., vol. Ill,
p. 569.

Raja chagrinea, Mont., Mem. Wern. Nat. Hist. Soc., vol. II,
p. 420, tab. XXI; Parn., ibid., vol. VII, p. 431, tab. XLI;
Couch, Fish. Brit. Isl., vol. I, p. 117, tab. XXIX; Mor.,
Hist. Nat. Poiss. Fr., tom. I, p. 401 ; D6derl., Man. Ittiol.

Medit., fasc. Ill, p. 172; Carus, Prodr. Fnce Medit., vol. II,
p. 521.

Raja aspera, Flmng, Brit. Anim., p. 172.

The ordinary dimensions of the Shagreen Skate,
as it is brought home by the deep-sea fishermen of
Bohuslan from the fishing-banks off Jutland and Nor-
way, are a length of about 1 m. and a breadth of about
6 dm. At this size it has reached full sexual maturity.
According to Moreau its length varies between 8 and
12 dm.

The form of the disk is rhombic. The distance
from the tip of the snout to the hind margin of the
ventral fins is about equal to the greatest breadth of
the body. The anterior side-margins of the disk are
undulate, somewhat concave, so that the most promi-
nent margin of the head falls short of the line between
the tip of the snout and that of either pectoral fin.

1116

SCANDINAVIAN FISHES.

The snout is prolongated in a securiform shape. The
posterior side-margins of the disk are arcuate, with
a slight inward curve near the tip of each pectoral fin.
The ventral fins are conically pointed and rounded,
rather long, their length, measured from the sacral
prominence, being not much less than 1/3 of the entire
length of the tail, measured from the same point. The
body as a whole is flat and not fleshy, its depth at
the shoulder-girdle being only about 9 % of the greatest
breadth of the disk. The tail is furnished, as in the pre-
ceding Rays except the Thornback, with two juxtaposited
dorsal tins, often united at the base. The hind margin of
the posterior lies half a fin-length from the tip of the tail,
which bears above a low and almost semicircular caudal fin.

The length of the head to the occiput measures in
specimens 1 m. long about 18 % of the length of the body
or 30 % of the breadth of the disk. The length of the
snout as stated above distinguishes the Shagreen Skate,
as well as the following species, from the preceding
forms of the genus, but the distinction is least- marked
in the Shagreen Skate, as appears from the relation
between this length and the least interorbital width of
the forehead. The latter is at least about 30 — 35 %
of the length of the snout from the anterior margin
of the eyes. The longitudinal diameter of the eyes is
about half the interorbital width. The spiracles, which
are almost transversely set, are of about the same size
as the eyes. The mouth is broad and curved, power-
fully armed with teeth. The form of the teeth is also
highly characteristic, and among the other Scandinavian
Rays there is none with whose dentition it can be con-
fused. Each tooth (fig. 320, a) lias a fairly long, un-
guiform crown, subulate at the tip, with somewhat
compressed sides and without other indication of the
ordinary basal rim than a very small intumescence on
each side. The largest teeth are set in the middle of
the jaws, and their size decreases towards the corners
of the mouth. All these teeth are arranged in about
60 regular, longitudinal rows, which, in consequence
of the length of the pointed crowns, the superimposed
(imbricated) position thereof, and the fixed intervals
between the rows, have a resemblance to thin ridges,
fitting in between each other when the jaws are closed.
The internasal width is about V5 — 1/3 greater than the
least interorbita-1 width (120 — 133 %), somewhat more
than half (55 — 60 %) of the distance between each
nostril and the tip of the snout, and about 1 2 % of that
between the tip of the snout and the anus.

Among all the Scandinavian Rays the present spe-
cies has the most shagreened skin. The whole upper
surface of the body is covered everywhere with dense,
but very short spinuke, partly pointed with stellate
base, partly granular and hardly distinguishable to the
naked eye. On the snout, both above and below,
throughout the anterior side-margins of the disk,
and throughout the upper and under surfaces of the
tail, these spinulae show a somewhat greater develop-
ment, and with their pungent tip, recurved in a hamate
form, they greatly resemble, on a small scale, the aculei
proper. The ventral side too is strewn with spinuke,
but more sparsely, large patches of smooth and naked
skin being left on the pectoral and ventral tins, in the
anal region, and at other points. The ordinary aculei
have an expanded, stellate or grooved, laterally com-
pressed, and flat- base and a hamately recurved tip.
Two short rows of small aculei, varying in number
and size, are set on the rostral cartilage. In a semi-
circle around the inner margin of each orbit runs a
row of larger, but unequal aculei, the middle ones,
however, often absent or lost. The spinal column be-
tween the head and the scapular cartilage is followed by
a series of aculei, 8 — 10 in number and not very large,
and sometimes a row of still smaller ones skirts this
series on each side; but- often only vestiges of these last
aculei remain in the form of small, worn protuberances
occupying the said places. Two rows on the back begin
just behind the scapular cartilage, the aculei being at
first quite low, but increasing in size a-s they approach
the tail, along which the rows proceed parallel to each
other, on each side of the median line and nearer to
this than to the side-margin of the tail. In these
rows the aculei are very densely packed, sometimes,
as it were, extruding one another. The largest aculei
lie in front of the middle of the tail; further back
their size again gradually decreases. On the spinal
column itself, behind the scapular cartilage and in the
caudal region, there is no row of aculei.

The dorsal side is of a plain yellowish gray, the
ventral side of a translucent milky white, the muscles
of the pectoral fins being visible, and the tips of the
fins showing a bluish tinge. According to Montagu the
back is of a plain grayish brown; but in one specimen
he found a few black spots on this ground-colour.

The external differences of sex are confined, ac-
cording to Fries and Malm, to the ordinary cards of
spines on the sides of the head and the outer parts

SHARP-NOSED SKATE.

1117

of the pectoral fins in the male and to his pterygo-
podia.

The Shagreen Skate is stated to have been taken
once on the coast of Sweden. Baron Gyllenstjerna
received it, according to Nilsson, in October, 1849,
from the fishing-village of Arild (Mount Kullen). But
the specimen has disappeared, and with it the certainty
that no confusion of the species has taken place. Ac-
cording to Collett a female 9x/2 dm. long was caught
in January, 1880, off the Onso coast, at the entrance
of Christiania Fjord and consequently on the confines
of Sweden. Its favourite haunts in Scandinavian wa-
ters, however, are the rich fishing-banks off Southern
Norway and the west of Jutland, where the water is
from 50 to 120 fathoms deep. Iyroyer and Collett
assign Trondhjem Fjord as the northern limit of its
range", and to the south it occurs, according to Gun-
ther, off Madeira and, according to Moreau, in the
Mediterranean, where it is described, however, as rare.
Like the other Rays it ascends nearer land in summer;
in winter it is not taken, except where the fishermen

follow it to the deep sea. It feeds on all kinds of
fishes, such as Sharks and Codfishes, as well as on
crustaceans and mussels6. The strong, pointed teeth
in its jaws are suggestive of considerable voracity. As
food it is little esteemed both in England (Day) and
France (Blanciierec).

The systematic name of the Shagreen Skate ( fullo -
nica, Lat. fullo, a fuller) has reference to the similarity
between the spiny . skin and the implements used in
cleaning cloth and the like. It was adopted by Lin-
naeus from Rondele'G, but was not applied originally
to this species. It is also probable that the Swedish
name ( gokrocka = Cuckoo Ray) was borrowed from
England, though the latter name is ascribed by Couch
to the preceding species. In Norway, according to
Ratfike, the Shagreen Skate is called Ncebskate (Beaked
Skate), a name which, as Iaroyer has remarked, is
more appropriate to one or other of the following spe-
cies. In Bo hu si an Fries also heard the name of Gcidsk-
rocka (Jutland Ray).

(Fries, Smitt.)

THE SHARP-NOSED SKATE (sw. hvitrockan or blagarnsrockan).

RAJA LINTEA.

Fig. 321.

Length of the snout from the anterior margin of the eyes about 14 — 18 % of the length of the body or 24
of the greatest breadth of the disk. Distance between each nostril and the tip of the snout about 21 — 25
said breadth or more than twice ( 207 — 243 % of) the internasal width,
median line of the tail and its lateral margins armed with rows of aculei.

the tip of the tail. Caudal fin rudimentary.

Syn. Raja Icevis , major (p. p.), Duh., Tr. Pitches, part. II, sect. IX,
p. 285.

Raja oxyrinclius ( Sharp-nosed, Ray), Penn., Brit. Zool. (ed.
1776), vol. Ill, p. 73; Mont., Mem. Wern. Nat. Hist. Soc.,
vol. II, p. 423; Yarr., Brit. Fish., ed. 1, vol. II, p. 424;
ed. 2, vol. II, p. 556; Couch ( Barton Skate), Fish. Brit.
1st., vol. I, p. 97, tab. XXI.

? Raja alba , Lacep., Hist. Nat. Poiss., tom. Y, p. 663, tab.
20, fig. 1 (+ Raja marginata, ibid.); Mor., Hist. Nat. Poiss.
Fr., tom. I, p. 412; Day, Fish. Gt. Brit., Irel., vol. II,
p. 339, tab. CLXVIII; ? = Raja ( Lceviraja ) bramante,
Sassi, vide Canestr., Mem. Accad. Sc. Torino, ser. 2, tom.
XXI (1865), p. 361, tab. I, figg. 2—5.

— 29 %
% of the

Aculei faintly grooved. Both the upper
Second dorsal fin extended nearly to
Ventral side white.

Raja Fullonica, Nilss., Prodr. Ichth. Scand., p. 119; Storm,
Norsk. Vid. Selsk. Skr., Trondhj., 1883, p. 47.

Raja lintea, Fr., Vet. Akad. Handl. 1838, p. 154; Mull.,
Hi.e, Plagiost., p. 147; Kr., Damn. Fisk., vol. Ill, p.
1005; Nilss., Skand. Fna , Fisk., p. 738; Malm, Ofvers.
Vet. Akad. Fork. 1857, p. 193; Gthr, Cat. Brit. Mus.,
Fish., vol. VIII, p. 466 (? + Raja marginata, p. 465);
Coll., Fork. Vid. Selsk. Chrnia 1874, Tilleegsh., p. 217;
1882, No. 29, p. 4; Malm ( Leucoraja ), Gbgs, Boh. Fna,
p. 611; Lillj. (Raja), Sc.. Norg. Fisk., vol. Ill, p. 580.

Obs. That the blagarnsrocka of Bohuslan (the Norwegian hvid-
skade) is identical in species with the English White Skate (the French

“ Storm, who states (Norsk. Vid. Selsk. Skr. Trondhj. 1883, p. 47) that Raja fullonica is very commonly taken in Trondhjem Fjord,
evidently referred here to the following species, for he says that “the middle row of caudal spines extends far forward on the back”.

5 Cf. Olsson, Om fiskarnes foda, Lunds Univ. Arsskr., tom. VIII (1871).
c La Peche et les Poissons , Diet. Gen., p. 666.
d De Pise., lib. XII, p. 356.

1118

SCANDINAVIAN FISHES.

raie blanche ), is extremely probable, though so large specimens have
never been found in Scandinavia as further south, nor has the juve-
nile form ( Raja marginata ) referred by Moreau to the said species
been met with in Scandinavian waters. Fries too was of this opinion,
for he combined Montagu’s Raja oxyrhynchus and his own Raja
Untea. In recent times (Dumeril®, Gunther, and Day) it has been
proposed again to separate them; but the best descriptions (Moreau,
1. c. and DSderlein6) seem to favour the opinion that they are iden-
tical, if we bear in mind the significance within the genus of the
alterations of growth and the external differences of sex.

Among the Sharp-nosed Skate, which were certainly
not few in number, brought home by the fishermen of
Bohusliin from the deep-sea fisher)’ in the North Sea,

Fig. 321. Sharp-nosed Skate ( Raja lintea ), Q, Ys nat- size- The
Cattegat, July, 1837. B. Fries. From a stuffed specimen, a, an
aculeus, after W. von Wright.

Fries never came across specimens more than 9 or 10
dm. long; but none of the females he opened had fully
developed ovaries, nor had the males full-sized ptery-
gopodiaY “It is thus probable,” wrote Fries, “that
these Skate attain a greater length before they are ca-

pable of reproduction, which is borne out by Pennant’s
statement that a specimen found by him measured 7
feet (21 V3 dm.).”

In form of body the Sharp-nosed Skate most nearly
resembles the preceding species. The anterior lateral
margins of the disk, however, show hardly so deep un-
dulation or concavity^, the most prominent side-margin
of the head approaching close to the line from the tip
of the snout to that of either pectoral fin. The ventral
tins too are shorter, more truncate at the tip, rectilinear;
their length from the sacral prominence being contained
3V2 times in the entire length of the tail. The two
dorsal fins on the tail are closely juxtaposited, though
sometimes without being confluent at the base, and the
posterior is set so far back that the upper fin-margin
projects beyond the tip of the tail, which is finless,
save for a lobe formed by an incision in the hind mar-
gin of the second dorsal fin.

The length of the head in a female 9 dm. long

O C>

measures nearly 1/4 of that of the body or 9/ 5 of the
greatest breadth of the disk. The interorbital width
(the least breadth of the cranial forehead) is V4 of the
length of the snout to the anterior margin of the eyes.
The diameter of the transversely set spiracles is much
less (V5, according to Krgyer) than the longitudinal
diameter of the eyes, which measures, according to Krd-
yer, % °f Fie interorbital width. The mouth is not so
broad as in the Shagreen Skate, and the dentition too
is feebler. The form of the teeth is not very charac-
teristic. It almost exactly resembles that of the common
Skate, with the exception that every tooth is compara-
tively smaller both in the area of the basal disk and the
length of the cusp, and that the number of longitudinal
rows is somewhat less, in the above-mentioned female
45 Y The internasal width is much greater than the
least interorbital width (150 — 175 % thereof), but at
most somewhat less than half the distance between each
nostril and the tip of the snout.

The Sharp-nosed Skate has the smoothest skin of
all the Scandinavian speciesY The ventral side is per-

a Hist. Nat. Poiss. (su. a Buff.), tom. I, p. 564, note.
b Man. Ittiol , Medit., fasc. Ill, p. 165.

c In the male 1,125 nun. long and with pterygopodia 190 mm. in length described by Malm ( Gbgs , Boh. Fna ), these organs had not
attained their full development, though nearly so. The largest female mentioned by Malm was nearly 12 dm. long. According to Doderlein
the Mediterranean Raja bramante grows to a length of more than 2 m. According to Day the White Skate of English waters sometimes
weighs nearly 500 lbs.

d In this respect the descriptions and figures of the present species vary considerably. According to KrOyer the tip of the snout in
a male is “strongly prolongated from the disk and of a considerable length.”

' In a male Kr0yer counted 48.

f Young specimens of the common Skate, however, are sometimes equally smooth.

SKATES.

1119

fectly soft and smooth", without a sign of spinulae, in
ordinary cases even under the snout and tail, except at
the extreme tip. The dorsal side is for the most part
naked, if we except the edge of the anterior lateral
margins of the disk, which is fringed with coarse, scat-
tered spinulae with stellate base, the rostral cartilage,
where similar, but still more scattered spinulae appear,
the dorsal fins, and the tail. The latter is somewhat
roughened with fine, very dense spines. These spines
present the peculiarity of being distributed in two
longitudinal bands, broad in front, narrowing behind,
which extend between the aculei of the median and
lateral rows, back to the sides of the base of the second
dorsal fin, the outer margin of each band touching the
lateral row of aculei, but the inner margin falling a
little short of the median row, so that the spinal column
itself, on which the aculei are situated, is covered between
them with quite smooth skin. The ordinary aculei are the
following: a) a few before and behind the eyes, b) a row
along the back and the middle of the tail, though it should
be observed that the aculei in the anterior part of this
row are often worn, short and tuberous, here and there,
or wanting at certain spots, c) 2 to 4 on each side of
the spinal column above the scapular cartilage, and d)
a row of smaller aculei on each side of the tail, oc-
cupying the extreme lateral margin, just above the
membrane with which this is edged. The aculei have
a singular and rather easily recognisable form. The
base is a triangular cone, with compressed sides, sharp
in front, truncate behind, and with a hollow and grooved

or somewhat radiate surface, and from this rather elevated
base there springs abruptly a very short cusp, usually
directed obliquely backwards, but on one or two of the
aculei at the side-margins of the tail antrorse. No
aculeus is ever present between the two dorsal fins.

The coloration is above of a plain bluish gray,
underneath of a pure milky white, with a grayish band
along the middle of the tail and a spot of the same
tint on each side of the anus. Sometimes a darker
tinge appears on the margins of the disk, both above
and below, being apparently a remnant of the juvenile
dress which has given rise to the name of marginata.

The external differences of sex are the ordinary

ones.

The haunts of the Sharp-nosed Skate are the same
as those of the Shagreen Skate, but it is more frequently
caught by the fisherman. It lives at the same consider-
able depth. Middle-aged individuals preponderate among
the Sharp-nosed Skate taken on the Norwegian banks.
In Bohuslan this species bears the names of Spanjor
(Spaniard) and Blagarnsrocka (Canvas Skate), and is well
known and distinguished from the others by every ex-
perienced fisherman. As we have already mentioned,
our Sharp-nosed Skate is probably the same species as
the French Bale lisse et blanche , which has been an
article of trade between French and English fishermen
for centuries, the former visiting the English coast to
purchase this fish for transportation to their own country,
where on account of its fleshiness it is more in demand
than other Rays. (Fries, Smitt.)

B: Ventral side of the body gray or grayish black, punctated with black dots and lines. — True Skates.

The two dorsal fins are separated, sometimes with, sometimes without interjacent aculei. The ventral side
is of a dirty grayish or blackish colour, with numerous black pores, opening into curvilinear muciferous canals
(Lorenzinian ampullae and divisions of the lateral line). No ordinary aculei on the back, but only on the tail
and sometimes at the eyes. These species attain a considerable size, live in very deep water, and are highly
valued for their flesh. (Fkies.)

The relations between the three following species
are very highly suggestive of the same close kinship as
we have considered at length above, especially in the
families of the Flounders and Salmons. These Skates
compose a form-series in which the specific distinctions

are more or less obvious expressions of the differences
of age and sex. The most prominent specific charac-
ters depend on the prolongation and contraction of the
anterior part of the body, especially the rostral region.
In this respect one of the Scandinavian forms, the com-

° Sometimes, however, spines occur on the \mder surface of the snout; and in Raja alba ( bramante ) the anterior margins of the
pectoral fins are also sometimes spinigerous underneath.

Scandinavian Fishes.

141

1120

SCANDINAVIAN FISHES.

mon Skate, stands side by side with the most advanced
stages of development in the species immediately pre-
ceding it. The course followed by its alterations of
growth is such that, during the growth of the body
from a length of 3 dm. to one of 15 dm., the length
of the snout increases from about 16 to 18 % of that
of the body, in exceptional cases exceeding the latter
percentage, or from 21 to 26 % (sometimes 27 %) of
the greatest breadth of the disk. In another of our
species, the so-called plogjernsrocka ( Baja vomer) of
Fries, where the alterations of development, however,
are scarcely known, these percentages for the length of
the snout vary between about 20 and 24 in relation to
the length of the body and between about 31 and 35
in relation to the breadth thereof. The first-mentioned
alteration during growth of the proportion to the length
of the body indeed depends in great part on another,
which we have also observed above, namely the rela-
tive abbreviation of the tail with increasing age; but
herewith is associated in the common Skate a difference
of sex, which is expressed by the comparatively greater
length of the abdominal region in the females than in
the males. This is distinctly shown by the relations
between the distances from the tip of the snout on the
one hand of the mouth or nostrils, on the other of the
cloacal aperture. The distances of the mouth and ej’es
from the tip of the snout are about equal in these Rays,
and decrease in male specimens of the common Skate
from about 53 to 38 %. of the distance between the

mouth and the cloacal aperture, whereas in the females
these percentages diminish from about 51 to 27. The
Long-nosed Skate represents in this respect the male
characters of the common Skate; but in the former
species, owing to the different prolongation of the snout,
which is longest in the females, the sexual distinction has
been reversed, the distance between the mouth and the
tip of the snout being in adult males about 58 — 60 %
of that between the former and the cloacal aperture,
in the females about 75 — 80 % of the same. The males
of the two species may consequently approximate so
closely to each other in form of body that the limit
between the species is difficult to fix. Furthermore
Parnell in Scotland, Bonaparte in the Mediterranean,
and Collett in Trondhjem Fjord have each distinguished
an intermediate form to fill the gap between the lines
of demarcation. This intermediate form has retained
one of the juvenile characters of the common Skate,
namely the aculei arming the supraorbital margin,
which disappear in old specimens of the common Skate,
and are wanting in the Long-nosed Skate. In Trondhjem
Fjord this intermediate form has acquired a still darker
(brownish black) ventral side; but even this peculiarity
seems primordially to have been a sexual character,
the males, according to Storm, being darker than the
females. The more northern common Skate, with its
more strongly marked female characters, thus repre-
sents an original form from which the two remaining
species are descended.

THE COMMON SKATE (sw. slatrockam).
RAJA BATIS.

Plate XLVI1I.

Length of the snout from the anterior margin of the eyes about 14 — 18 % of the length of the body or 21 — 26%
of the greatest breadth of the disk. Distance betiveen each nostril and the tip of the snout about 21 or 22 % of
the said breadth and less than twice ( 180 — 140 % of) the internasal width. Least interorbital width more than
V4 ( about 30 — 42 %) of the length of the snout. Aculei without grooves {or with extremely faint ones). A row
of aculei either in the upper median line of the tail or on each of its lateral margins , seldom simultaneously
present on the former and the latter. The second dorsal fin ends at a distance from, the tip of the tail measuring
more than half its oivn base , and the tip of the tail is furnished above with a distinct caudal fin. Ventral side

grayish or darker , dotted and streaked with black.

Syn. Raja undulata sive cinerea , Rondel., De Pise., p. 346. Raja
Icevis (Tepel), Schonev., Ichth. Slesv. IIols., p. 58. Raja

Icevis undulata seu cinerea Rondeletii, Willtjghb., Hist.
Pise., p. 69, tab. C, 5. Raja varia, dorso medio glabro, unico
aculeorum ordine in cauda, Art., Ichtli., Gen. Pise., p. 73;
Syn. Pise., p. 102. Skat a,, Raja (major et vulgaris) dorso

non aculeato, Olafs., Reise Isl., pp. 359 et 987. Skate
1. Roklce, Raja clavata Auctt., Strom, Sondm. Beskr., p.
309; efr Trondhj. Selsk. Skr., vol. I, p. 148.

Raja batis, Lin., Syst. Nat., ed. X, tom. I, p. 231; Penn.
(Skate), Brit. Zool. (ed. 1776), vol. Ill, p. 72, tab. IX;
Rl., Fiscli. Deutschl., part. Ill, p. 54, tab. LXXIX; Hollb.

COMMON SKATE.

1121

( Dasybatus ex Blainv.), Bohusl. Fisk., G-bgs Welt., Witt.
Samh. N. Handl., part. Ill (1819), p. 21 cum 2 tabb.;
Fab. (Baja), Fiscli. Isl., p. 33; Nilss., Prodr. Iclithyol.
Scand., p. 120; Yarr., Brit. Fish., ed. I, vol. II, p. 421;
Fr., Vet. Akad. Handl. 1838, p. 158; Mull., Hle, Plagiost.,
p. 146; Kr., Damn. Fisk., vol. Ill, p. 978; Nilss.,
Slcand. Fan, Fisk., p. 739; Malm, Olivers. Vet. Akad.
Forh. 1857, p. 193; Gthr, Cat. Brit. Mas., Fish., vol. VIII,
p. 463; Ltkn, Vid. Meddel. Naturh. For. Kb*hvn 1873,
p. 36; Coll., Forlr. Vid. Selsk. Chrnia 1874, Tillsegsh.,
p. 216; 1879, No. 1, p. 106; N. Mag. Naturv. Chrnia,
Bd 29, p. 119; Malm ( Lceviraja ex Salv. et Bonap.),
Gbgs, Boh. Fna, p. 615; Winth. {Raja), Naturh. Tidskr.
Kbhvn, ser. 3, vol. XII, p. 60; Mor., Hist. Nat. Poiss.
Fr., tom. I, p. 409; Mob., Hcke, Fiscli. Osts., p. 156;
Day, Fish. Gt. - Brit. Irel. , vol. II, p. 336, tab. CLXVI;
Lillj., Sv., Norg. Fna, Fisk., vol. Ill, p. 584.

Ruia gaimardi, Robert in Gaim., Voy. Isl., Groenl., Zool.,
Poiss., tabb. 2 et 3 ; Dum., Hist. Nat. Poiss. (n. su. a Buffi.),
tom. I, p. 565.

Batis vulgaris, Coitch, Fish. Brit. Isl., vol. I. p. 87, tab.
XVIII.

Most of the common Skate taken in Bohuslhn
measure between 12 and 15 dm. in length, but at this
size are hardly mature. On the south and west coasts
of Norway Krgyer met with specimens 22 — 25 dm.
long and rising 200 Norwegian pounds (100 kilo.) in
weight.

The form of the body is rhombic, with the anterior
side-margins somewhat undulate and more or less in-
curved, the posterior, on the other hand, very faintly
rounded (convex), with a slight incurvature just behind
the tip of each pectoral fin. The distance from the tip
of the snout to the hindmost part of the posterior
margin of the pectoral tins is about 74 — 84 % (as a
rule about 77 %), and to the posterior margin of the
ventral tins about 80 — 94 %, of the greatest breadth of
the disk. The greatest thickness (the depth at the
shoulder-girdle) is about 7 or 8 % of the greatest breadth.

The form and length of the head vary consider-
ably, in proportion to the greater or less prolongation
of the snout as mentioned above. The longitudinal
diameter of the eyes is about 1j5- — 1/6 of the length of
the snout. The maximum diameter of the spiracles is
about 3/5 — 4/5 of that of the eyes. The internasal width
measures about 57 (in young specimens, according to
Krgyer, sometimes 55) — 68 % of the distance between
each nostril and the tip of the snout, which distance
is about 88 (in young specimens, according to Kroyer,
sometimes 91) — 83 % of the length of the snout. The
breadth of the mouth is about equal in young speci-
mens to the internasal width, in older ones perceptibly

greater, at least 112 — 114 % thereof. The jaw-teeth
are comparatively small (tig. 322). On a neck-like,
but low base is extended an almost nail-headed disk,
the posterior part of which rises in a somewhat conical,
retral tip, short in the females (fig. 323), longer in the
males. Teeth of this form are distributed in fairly
regular rows, denser in young individuals than in old,
both along and across the jaws. The largest teeth, with
the longest cusps, are set in the middle of the mouth,

Fig. 322. Half of a jaw in a male {A) and a female (B) of the
common Skate {Raja hatis). Natural size. After Lutken.

the smallest, with hardly a sign of cusp, innermost at its
corners. As a rule the longitudinal rows number some
40 — 50, the transverse 6 — 10. The various ages ex-
hibit less difference in the number of the rows than
in the form of the teeth, for in very small Skate there
is scarcely an indication of the projecting cusps which
are so prominent in older specimens, especially in the

Fig. 323. A tooth of Raja batis, magnified.

males; and the whole tooth thus acquires in the young
a plane, lozenge-shaped appearance.

The spiny armature of the body consists in very
young specimens merely of a restricted number of ordi-
nary aculei, namely two before and one behind each
eye and a row along the upper median line of the tail
with one or two spines between the two dorsal fins.
Of spinulm there is not a sign in these young speci-
mens, either on the dorsal side or the ventral, the
body being smooth on both sides. In middle-aged in-

1122

SCANDINAVIAN FISHES.

dividuals both the upper and under surfaces of the
body are roughened with spinulae, which are, however,
somewhat unevenly distributed, large and small patches
of perfectly smooth skin occurring here and there. The
spinulae are also rather scattered, except on the under
surface of the snout and around the inner orbital mar-
gins, where they are card-like. They do not extend
to the hind margin of the pectoral fins, which is per-
sistently smooth. In form these spinulae are very
pointed and fine, with a somewhat, expanded, stellate
base, the diameter of which is less, however, than the
entire length of the spine, and which is scarcely dis-
tinguishable to the naked eye. Here and there they
are worn and more granular, and their size is highly
variable. In middle-aged specimens the orbital aculei
are as a rule already worn down or missing. This is
partly the case too with the median caudal row, of
which there usually remain only one or two aculei and
the marks of the rest. Each lateral margin of the tail,
on the other hand, has developed by this age a row
of aculei, which is also very variable, seldom symmet-
rical, and displays the peculiarity that most of its
aculei are oftenest directed forwards. Full-grown fe-
males are rather densely covered with spinulae through-
out the dorsal side, except on the outermost parts of
the hind margin of the pectoral fins, and on the pos-
terior fold and anterior lobe of the ventral fins. The
ventral surface of the snout is almost as densely armed
with spin uhe as the dorsal side. At the middle of the
body and on the tail, as well as on the inner parts of the
hind lobe of the ventral fins, the ventral side is also spini-
gerous, but the spinulae are more scattered and grow
sparser and sparser outwards, until they disappear on
the outer parts of the pectoral and ventral fins, the an-
terior lobe of the ventral fins being also naked. In adult
males the spiny armature is as usual less developed.

The coloration of the dorsal side is yellowish gray
or brown, with irregularly distributed and highly vari-
able spots of lighter and darker tint. Young and
middle-aged specimens frequently have one or two pairs
of ocelli, varying in size, on the posterior and inner parts
of the pectoral fins. One of our figures (PI. XLVIII,
fig. 2) represents a specimen from Gull mar Fjord with
the dorsal side strewn everywhere with light spots (van.

a Prodr. Face Medit ., vol. II, p. 523.

b Expos, intern, di Pesca, Berlino 1880, Sez. Ital. Catal., p.
c Mannuale Ittiol. Medit ., 1. c.

guttata). The ventral side is of a dirty grayish white
or darker, chocolate-coloured, shading towards the hind
margins of the disk into a grayish violet. It is also
dotted and streaked, as we have mentioned above, with
black, the apertures and opening ducts of the system of
the lateral line being thus indicated. This black punc-
tuation extends 'sometimes even to the anterior part of the
back. Sometimes, however, the ventral side is so white
— Baron Cederstrom has presented to the Royal Museum
a specimen of this description, a male 1 m. long and not
yet mature, from Stromstad — and the black punctua-
tion so sparse and faint that the difference from the
Sharp-nosed Skate in this respect is inconsiderable.

The common Skate is strictly a North Atlantic fish,
its range extending from Norwegian Finmark and Ice-
land south to the Bay of Biscay. According to Moreau
and Carets'1 it also occurs in the Mediterranean, being
fairly common, especially off Cette and Riou; but neither
GigliolP nor Doderlein0 includes it among the list of
Mediterranean fishes. The difficulty of fixing a specific
limit between this species and the so-called Black-bellied
Skate, which at least has a representative form in the
Mediterranean, renders it impracticable to pass an opi-
nion on this head without access to specimens. Our
knowledge of the North American Rays is also too un-
certain to decide the question whether the common
Skate does not belong as well to the western parts of
the North Atlantic. The waters round the British Isles
and the fishing-banks off' the south and middle of Nor-
way are undoubtedly the most frequented haunts of
the common Skate. It also enters in numbers the deep
fjords and the island-belts of the west coast of Scandi-
navia. In Bohuslan it is commonly taken. In the
Sound it penetrates at least to the neighbourhood of
Landskrona, and through the Belts it makes its way,
though rarely, at least to Travemiinde Bay (Mobius
and Heincke). Its bathymetric range is also rather
extensive. The older individuals are taken in Norway,
according to Lilljeborg, at depths sometimes amounting
to 200 fathoms. But, like the other Rays, it ascends
to higher levels during summer; and Kroyer once saw
4 specimens, 12 to 15 dm. long, taken at the end of
September in a stake-net off Gilleleje (the north coast
of Zealand), in water probably not more than 4 fa-

114.

BLACK-BELLIED SKATE.

1123

thorns deep, this being the maximum depth of an or-
dinary stake-net.

The common Skate seeks its food, like other Rays,
among all kinds of marine animals, its size rendering
it a still more dangerous enemy of the larger among
them, such as Flatfishes, Codfishes, and large crusta-
ceans. Even its congeners do not escape.

The breeding season of the common Skate occurs
in spring. During summer, from May till September,
it deposits its eggs, the shell of which, according to
Kroyer, is of a handsome sea-green colour when fresh.
While dredging in very deep water off the outermost
islands in Bohuslan, Fries secured the empty egg-shell
of a Ray. It measured 32 cm. in length and rather
more than 1 3 1/2 cm. in breadth0, and in all probability
belonged to the common Skate.

The fairly plentiful occurrence of this species ren-
ders it of no little economical importance to man. The
largest Skate are less esteemed, however, than those of
medium size. When boiled fresh, Skate finds no great
favour with the fishing population, who prefer either
to keep it a day or two before eating or to score it
with deep long slits, after which it is either salted or
dried. Under the last-mentioned form it is often sub-
stituted for stockfish in their Christmas fare6.

No special fishery can be said to be practised for
the common Skate; but on his long-lines the fisherman
usually leaves every fifth snood, the so-called Ray-snood,
without float, so that it may sink to the bottom, where
the bait, generally the head of a Herring, is more easily
reached by the Skate.

(Fries, Smitt.)

THE BLACK-BELLIED SKATE (sw. SVARTBUKSROCKAN c).

RAJA NIDROS1ENSIS.

Figs. 293, A and 324.

Length of the snout from the anterior margin of the eyes (in adult specimens ) about 19 — 22 % of the length of
the body or 28 % (27 — 30 %) of the greatest breadth of the disk. Distance between each nostril and the tip of
the snout about V4 of the said breadth and more than twice the internasal width. Least interorbital width less
than V4 (24f2 — 21 %) of the length of the snout. Females'3 with three roivs of aculei on the tail, males with
only one (that of the upper median line). Otherwise: aculei , dorsal fins, and caudal fin as in the common Skate.

Ventral side so dark (blackish) that the

Syn. ? Raja Macrorynchus, Rafin., Caratt. ale. n. gen., p. 15;

Bonap. ( Lceviraja ), Iconogr. Fna Ital., Pesci, tab. 151, fig.

2; Gthr (Raja), Cat. Brit. Mus. Fish., vol. VIII, p. 468;
Mor., Hist. Nat. Roiss. Fr., tom. I, p. 405; Day, Fish.

Gt. Brit., Irel., vol. II, p. 338, tab. CLXVII.

? Raia intermedia, Parn., Mem. Wern. Nat. Hist. Soc., vol. VII,
p. 429, tab. XL; Rich, in Yarr., Brit. Fish., ed. 3, vol.

II, p. 557.

Raja nidrosiensis, Coll, apud Storm, N. Vid. Selsk. Skr. Trondhj.
1880, p. 80; Coll., Forh. Vid. Selsk. Chrnia 1881, No. 7;
Storm, 1. c. 1883, p. 47 ; Coll., N. Mag. Naturv. Chrnia,

Bd 29, p. 121; Lillj., Sv., Norg. Fna, Fisk., vol. Ill,
p. 576.

Of the Black-bellied Skate only large specimens,
about 14 — 20 dm. long and 10 — 14 dm. broad, have
hitherto been found. In form of body it is so similar
to the common Skate that the longer snout can merely
be adduced as a character which is generally distinctive

“ In its present dried condition it measures, including the short
b See p. 1107, note /.
c Lilljeborg, 1. c.
d According to Collett.

black dots and streaks are indistinct.

of the Black-bellied Skate, but which — to judge by
the course of development known to be followed by
the other Rays — will probably lose its significance
when information has been obtained of the earlier staves
in the growth of this fish. The skeletons of the two
species are also alike, apart from the length of the
rostral region. The only difference worthy of remark
in skeletons of about equal size seems to be that in the
pectoral fins of the Black-bellied Skate the mesoptery-
gium is bifid; but the number of the radialia applied
to the two parts is the same (13) as that possessed by
the undivided mesopterygium of the common Skate.
The outer (anterior) lobe of the ventral fins also appears
in the Black-bellied Skate to be longer in proportion to
the posterior lobe; but to judge by Collett’s measure-

horns, 2 4 1/2 cm. in length and 8Y2 cm. in breadth.

1124

SCANDINAVIAN FISHES.

ments of these parts, the variations in this respect are
equally great in the two species and overlap each other.

The spiny armature of the skin is generally weaker
and on the dorsal side sparser than in the common Skate
— sometimes it is entirely wanting — but on the ventral
side it is more uniform and more extensive, except at
the outer margins of the ventral tins and on the tail,
which in some instances, however, has a strip of tine
spinulse along the middle of its under surface. The aculei
of the young are persistent, at least in the males, on

In a male 1 43/4 dm. long the length of the head
to the occiput measured 27'8 %, that of the snout from
the anterior margin of the eyes 21 %, and the distance
between the mouth and the tip of the snout 22 ‘3 %, of
the length of the body. The last-mentioned distance
was 68 % of that between the mouth and the cloaca!
aperture, a percentage which, according to Collett’s
figure, amounts in a female 19 dm. long only to 56.
The breadth of the mouth was 40 % and the internasal
width 45'8 % of the length of the snout". The jaw-

Fig. 324. Black-bellied Skate ( Raja nidrosiensis), ok from Trondhjem Fjord, March 17, 1891. Conservator Storm. 1/1 0 nat. size.

The skeleton of this specimen is figured above, p. 1064.

the orbital margins, and are also retained in both sexes
on the upper median line of the tail, a few usually re-
maining as well between the two dorsal fins; but ac-
cording to Collett only the females are furnished with
a row of aculei on each lateral margin of the tail.

" According to Collett’s figure the last-mentioned percentage
to have been even narrower than that of the specimen described abov
b The distinction adduced by Moreau between the dentition:
of age and sex.

c According 1o Carus Raja macrorhynchus is also dark brown

teeth are similar to those of the common Skate both
in number and form b.

The coloration of the dorsal side, according to Col-
lett, is of a dark grayish brown, and the ventral side
is nearly brownish black, blackest in the middle c. The

amounts in a female 19 dm. long hardly to 37; and the mouth seems
e.

3 of Raja macrorhynchus and R. batis evidently depends on differences
{fused) both above and underneath.

LONG-NOSED SKATE.

1 125

iris is bronze-coloured, the pupil shades into green and
blue-black. The male Black-bellied Skate from Trond-
hjern Fjord presented by Storm to the Royal Museum
in March, 1891, and the original of our figure, showed
on the dorsal side about the same coloration as the
Long-nosed Skate represented in Plate XLIX; the black
skin of the ventral side had been for the most part
chafed away during the railway-journey to Stock-
holm.

The Black-bellied Skate is hitherto known only
from the deepest parts of Trondhjem Fjord, where Storm
first distinguished it in 1880, among the fish taken at

a depth of 150 — 300 fathoms. It is stated to be fairly
common there. As mentioned above, however, there is
scarcely any reason to regard it as distinct in species
from the Scotch Baja intermedia and the Mediterra-
nean B. macrorhynchus. Its manner of life, says
Storm, is about the same as that of the other deep-
sea Skates. It voraciouslv devours both large fishes
and crustaceans. The fishes he most frequently found
in its stomach were Macrurus and Sebastes as well
as Spinax niger and Pristiurus catulus, and the crus-
taceans commonest among the contents were Lithodes
maja, Pasiphae tarda , and Pandalus borealis.

THE LONG-NOSED SKATE (sw. plogjernsrockan).

RAJA OXYRHYNCHUS.

Plate XLIX.

Length of the snout from the anterior margin of the eyes about 20 — 24 % (241fJ %) of the length of the body
or 31 — 35 % (36 %) of the greatest breadth of the disk. Distance between each nostril and the tip of the snout
about 27 — 31 % of the said breadth and more than twice the internasal width. Least interorbital width less
than 1/4 of the length of the snout. Other essential characters as in the common Skate.

Syn. Leviraja mucusa, buvosa, Salv., Aquat. anim. hist., fol. 149,
fig. 52. Raia oxyrliynchos major , Willughb., De Pise.,
p. 71, tab. C, 4 (ex Salv.). Raja varia, tnberculis deeern-
aculeatis in medio dorsi, Art., Ichthyol., Gen. Pise., p. 72;
Syn. Pise., p. 101.

Raja Oxyrinchus , Lin., Syst. Nat., ed. X, tom. I, p. 231;
Bonap. ( Lceviraja ), Iconogr. Fna Ital., tab. 151, fig'. 1; Gthr
(Raja), Cat. Brit. Mus. Fish., vol. VIII, p. 469; Mor.,
Hist. Nat. Poiss. Fr., tom. I, p. 403; Day, Fish. Gt. Brit.,
Irel., vol. II, p. 341, tab. CLXIX; Doderl., Afan. fttiol.
Medit., fasc. Ill, p. 152.

Raja chagrinea, Y-arr., Brit. Fish., ed. 1, vol. II, p. 414.
Raja Vomer, Fr., Vet. Akad. Handl. 1838, p. 161; Mull.,
Hle, Plagiost ., p. 144 (+ R. Salviani, p. 143); Kr., Damn.
Fisk., vol. Ill, p. 1011; Nilss. ( oxyrhinehus ?), Slcand.
Fna, Fisk., p. 740; Malm, Ofvers. Vet. Akad. Fork. 1857,
p. 193; Rich, in Yarr., Brit. Fish., ed. 3, vol. II, p. 548;
Gthr, Cat., 1. c., p. 468; Coll., Forh. Vid. Selsk. Clirnia
1874, Tillsegsli ., p. 217; 1879, No. 1, p. 106; Nyt Mag.
Naturv. Clirnia, Bd 29 (1884), p. 120; Malm, Gbgs, Boh.
Fna, p. 617; Storm, Norsk. Vid. Selsk. Skr. Trondhj. 1880,
p. 81; 1883, p. 46; Lillj., Sv., Norg. Fna, Fislc., vol.
Ill, p. 598.

Raja mucronata, Couch, Corn. Fna, p. 25; Fish. Brit. 1st.,
vol. I, p. 93, tab. XIX; Yarr., Brit. Fish., ed. 2, vol. II,
p. 550.

That this species also ranks among the largest
Scandinavian forms of the genus and perhaps rivals the
common Skate in size, appears from the fact that females
measuring li/3 m. are found that have not yet reached

maturity. As a rule, however, the Long-nosed Skate
is smaller than the common Skate, and the males are
sometimes mature, as our figure shows, at a length of
121 cm. The largest females found by Malm among
the take from the fishing-banks of J itderen were 14
dm. long, and the largest males measured 137 cm.

The form of the body is highly characteristic of
the Long-nosed Skate. Owing to the strong prolongation
of the snout and its pointed or ploughshare-like form,
the disk acquires the appearance of a sector of a circle,
the two radii (the anterior side-margins of the disk)
being undulate and incurved, so that the most pro-
minent lateral margin of the head falls a good way
short of the straight line from the tip of the snout to
that of either pectoral fin. The posterior side-margins
of the disk together form a handsome circular curve,
the centre of which lies in the middle of the mouth
aperture. The distance from the tip of the snout to
the hindmost part of the posterior margin of the pec-
toral fins measured in two females about 83 — 84 % of
the greatest breadth of the disk, in two males about
88 — 90 % of the same. The distance from the tip of
the snout to the posterior margin of the ventral fins
is very nearly the same as the greatest breadth of the
disk (about 92 — 104 % thereof).

1126

SCANDINAVIAN FISHES.

The length of the head to the occiput, is about
28 — 30 % (according to Lilljeborg sometimes 1j 3) of
the length of the body. The longitudinal diameter of
the eyes measures about 7 io' — V12 °f the length of the
snout. The spiracles, the diameter of which is as usual
variable, are of about the same size as the eyes. The
internasal width is about 40 — 45 % of the distance be-
tween either nostril and the tip of the snout, which
distance measures about 90 % of the length of the snout.
The breadth of the- mouth, even in old specimens, is
perceptibly less than the internasal width. The denti-
tion almost exactly resembles that of the common Skate.
The only difference that might be adduced is that, on
comparing specimens of equal size, the teeth of the
Long-nosed Skate prove to be somewhat smaller, and
the retral cusp ascends in a more conical form. The
rows of teeth are nearly the same in number as those
of the common Skate. The divergencies to be observed
in this respect are most naturally regarded as the re-
sults of individual circumstances.

In middle-aged and large specimens the whole skin,
both of the dorsal side and the ventral, is densely and
almost uniformly covered with spinulm of almost equal
size, which have a stel lately expanded base, fairly dis-
tinct to the naked eye, and consisting of 3 — 5 prostrate
rays, each of a length nearly equal to the height of
the spine. In the females no smooth gaps are left,
except the circumanal region and the under side of the
ventral fins and tail, which surface is partly naked. The
rest of the skin is uniformly roughened throughout,
even to the hind margins of the pectoral fins. In the
males the smooth patches are rather more extensive.
Around the eyes the spinulaa are somewhat coarser,
and in jmung specimens, according to Doderlein, a
well-developed aculeus is set at the preorbital margin.

The only other aculei possessed by this species
consist of a row on each lateral margin of the tail.
They are comparatively small, but of unequal size,
pointed and compressed, with the base prolongated and

as it were lobed. Their arrangement is not symmetrical,
and their tips are directed indiscriminately forwards or
backwards.

The coloration of the dorsal side is brownish gray
or reddish brown, with sparse, light brown or even
milk-white spots, scattered in curved rows or irregu-
larly. The anterior side-margins and the spine-cards
of the males are light gray. The under surface is
coloured and marked as in the common Skate, but
seems in general to be lighter.

The Scandinavian distribution of the Long-nosed
Skate is probably the same as that of the common
Skate, only that the former does not enter the Cattegat
proper, though it no doubt approaches the Swedish coast
in the Skager Rack. On the 14th of November, 1889,
Mr. C. A. Hansson secured in Koster Fjord, off Helsft,
the male figured in our plate. The northern limit of
the species apparently lies near Trondhjem, the southern
limit off Madeira. The Long-nosed Skate is besides a
rather common fish in the Mediterranean. Its bathy-
metric range appears to extend in Scandinavia between
depths of about 70 and 150 fathoms.

In its manner of life the Long-nosed Skate re-
sembles the common Skate. The same fishes are found
in the stomach of both, and off our coasts they are
taken on the same tackle (long-lines) and in company.
With regard to the deposition of the ova Grieg noted"
that a female 129 cm. long, which was caught on the
18th of May, 11 miles west of Bergen, laid a fully
ripe egg when drawn ' into the boat.

In Bohusl&n the Long-nosed Skate is well known
among the deep-sea fishermen and to the fish-dealers.
It is there called, according to Malm, varndbb (Big
Beak). It is taken less frequently by the Swedish
fishermen than the common Skate; but according to
Storm, Jensen, and Grieg it is common in Trondhjem
Fjord and off Bergen.

As a food-fish it is considered inferior to the com-
mon Skate. (Fries, Smitt.)

Bergens Museums Aarbog 1892, p. XVIII.

SHARKS.

1127

PLAGIOSTOMI SELACHOIDEI”.

Pectoral fins of ordinary form , situated behind or below the branchial apertures. Tail gradually contracted in
unbroken continuity with the trunk. Anal fin present or wanting. One or two dorsal fins, the anterior generally

belonging to the abdominal portion of the body.

Among the Sharks instances are indeed afforded
of comparatively sluggish bottom-fishes; but this sub-
order consists principally of the most powerful and
enduring swimmers, which chase their prey in speedy
pursuit and seize it with dexterous agility. Their form
of body adapts itself to their manner of life. The quiet

body and more pointed fins. Especially their caudal
fin, the propeller of the body, is in general powerfully
developed and, in contradistinction to that of the Rays,
principally ventral (heterocerca.1).

The sensory organs of the Sharks are also adapted
to the requirements of their predatory existence. The

Fig. 325. Diagram showing the ramification of the cranial nerves and the system of the lateral line in the anterior part of a Greenland

Shark ( Acanthorhinus carcharias). After Ewart.

aq, aquceductus vestibuli; Au , auditory nerve; bu , buccal branch of the facial nerve; cgl, ciliary ganglion; Fa, roots of the facial nerve;

fa, facial nerve proper; Gl , glossopharyngeal nerve; gbu, ganglion of the buccal branch; ggl, ganglion of the glossopharyngeal nerve; g. int,

ganglion of the intestinal branch of the nervus vagus; g. Vag 4, ganglion of the fourth branch of the nervus vagus ; Inn, ganglion of the
hyomandibular branch of the facial nerve; by a, hyoid ampullae; lab, labyrinth; Ira, lateral ampullae of the snout; ma, mandibular ampullae;
md, mandibular branch of the trigeminal nerve; mx, maxillary branch of the trigeminal nerve; N, nasal cavity; rt. int, intestinal branch of
the nervus vagus; n. lat, lateral branch of the nervus vagus; o, mouth; oa, oral ampullae; Oc , eye; ora, oculomotor nerve; pb, last prebranchial
division of the nervus vagus; ph, last pharyngeal branch of the nervus vagus; pi, palatine branch of the facial nerve; po, pit organs; Pr and
pr, nervus ophthalmicus profundus; ptb, last postbranchial division of the nervus vagus', ptbf, postbranchial division of the facial nerve (running
behind the spiracle): Pa, superior rostral ampullae; sof, superficial ophthalmic branch of the facial nerve; sot, superficial ophthalmic branch

of the trigeminal nerve; sp, spiracle; Tv, trigeminal nerve; Vag] — °, first — fifth branches of the nervus vagus; 1 — 27, canals of the system

of the lateral line. The signification of these numerals is explained below fig. 308, see above, p. 1091.

and . indolent Sharks have a more depressed body — an-
teriorly at least — and more rounded fins; the active
and more distinctly raptatorial a more terete, fusiform

system of the lateral line (see the explanation of fig.
325) consists of the same divisions as in the Rays,
only that here the opercular (hyomandibular) canal

a From the Greek oslayog, a cartilaginous fish.

Scandinavian Fishes.

142

1128

SCANDINAVIAN FISHES.

(12 — 13) does not extend beyond the branchial aper-
tures, sometimes not even to them, and that the man-
dibular branch is continuous with this canal.

In Scandinavian waters Sharks are less common
and consequently less known and feared than in more
temperate or tropical seas. About one hundred and
fifty species are recognised, but of these only eleven
can be claimed for the Scandinavian fauna.

In the sketch given by Hasse" of the evolution of
the Elasmobranchs one group, the Palceonotidani, in the
present age most nearly represented by the Notidanidceb,
a family foreign to Scandinavia, rank as the primordial
type both of the Sharks and Rays. Externally this fa-
mily is characterized by the possession of only one
dorsal hn, but of six or seven gill-openings. The proofs
of its primitive nature must be sought, however, in the
structure of the spinal column, with its undifferentiated
vertebrae. From the Palceonotidani, according to ITasse,
three different evolutional series may be traced, 'with a
calcified double-cone, more or less distinct and of varying
development, in the centra of the vertebrae. Its deve-
lopment is most imperfect in the so-called Cyclospon-

dyli, where it appears merely as a ring of calcification.
Externally all these Sharks (the family Spinacidce ), with
their typical form of body, may lie recognised by
their want of anal fin. In the other two evolutional
series the double-cone (composed of two more or less
deeply hollowed cones, confluent at the vertices) is
generally developed to such a degree that the bodies
of the vertebrae are fully amphiccelous; but in one of
them — the so-called Tectospondyli — it consists of
smooth, concentric layers of calcification, in the other
— the so-called Aster ospondyli — calcareous rays issue
radially from the centrum. The Tectospondyli are
without anal fin, and their series comprises both two
Shark families foreign to the Scandinavian fauna, the
Angel-fishes ( Tthinidce ) and the Saw-Sharks ( Pristiopho -
ridce), which are transitional even in their external
form to the Batoidei, and the whole phalanx of the
Rays. All the remaining Sharks — all furnished with
anal fin and two dorsal fins — are Asterospondyli ,
and in this series the Plagiostomous type has attained
its richest development, with the greatest wealth of
families.

ASTEROSPONDYLI.

Sharks with anal Jin and two dorsal Jins.

Among the Scandinavian Sharks the families be-
longing to this series may be distinguished as follows:

A: First dorsal fin situated between the perpendiculars drawn
through the pectoral and ventral fins.

a: Eye with a nictitating membrane Fam. Carcliariidce.

b: Eye without nictitating membrane... „ Lamnidce.

B: First dorsal fin situated above the ventral
fins or behind the perpendicular drawn
through these fins „ Scylliidce.

Fam. CARCHAKIIDjE.

Two dorsal Jins and one anal , the first dorsal opposite to the space between the pectoral and ventral fins. Eye
with a nictitating membrane, which is drawn up from the lower part of the orbit. Spiracles obliterated or minute.
The hindmost or even the penultimate gill-opening situated above the base of the pectoral fin.

According to Gunther’s definition c of this family it
includes among its 60 — 70 species the true Hounds {Mus-
teli) and their nearest relatives • — with small, pointed
or flat, more or less paved jaw-teeth — as well as the
Hammerheads ( Sphyrnce ), with their singular lateral

production of the orbital and nasal regions. These two
subfamilies are indeed strangers to the Scandinavian
fauna, as far as we know at present, but approach very
near to its limits, a species of each having been found
on the Scotch coast, and one of these ( Mustelus vulgaris)

a Naturl. Syst. Elasmobr., Jena 1879, Allgem. Theil, pp. 35, cett.

6 Two, perhaps three species of this family are inhabitants of the Mediterranean and the Atlantic; and one of these species, Noti-
danus ( Hexanclius ) griseus, is not so very rare on the English coast and has been met with so near the limits of the Scandinavian fauna as
off the coast of Scotland.

c Cat., vol. VIII, p. 353.

MAN-EATER SHARKS.

1129

extending its wanderings up to the Shetland Islands.
The third division, the subfamily Car char iince, is re-
cognised by the normal development of its head and
its large, triangular or subulate jaw-teeth, and to it
belong most of the species within the family.

The two genera which occur in Scandinavian wa-
ters may be distinguished as follows:

A: Peduncle of the tail crossed by a trans-
verse groove (a notch) above and below,
just in front of the base of the caudal fin Genus Carcharias.
B : Contours of the tail uninterrupted (with-
out notch) in front of the caudal fin... ,, Galeorliinus.

Obs. From Iceland Faber described" a species of Shark 7 —
9 ft. long, which he called Squalus arcticus. Nilsson assumed that
so large a Shark, with the well-known roving proclivities of these
fishes, could not occur off Iceland without paying at least an occa-
sional visit to Norway; and on this ground Squalus arcticus Avas in-

troduced into the Scandinavian fauna* *. The elder Reinhardt showed,
hoAvever, that, admitting the possibility of one or two slips of the
pen or lapses of memory. Faber’s description is most appropriate to
Isurits ( Lamna ) cornubicus c. More recently Muller and Henle found
in the Museums of Berlin and Leyden a species of Sharkd which they
assumed to be Faber’s Squalus arcticus, and which they placed in
their genus Galeocerdo , most nearly resembling Galeorhinus, but with
a groove in front of the caudal fin. Kr0Yer also states e that he has
examined two mandibles of Galeocerdo arcticus from Iceland. But
Faber’s description by no means coincides with that given by Muller
and Henle of the last-mentioned species. Especial attention is due
to the following points in Faber’s description: “Schnauze verlangert,
zugespitzt . . . Nasenoffnungen dicht vor den Augen . . . Schwanzflosse
fast halbmondformig.’ Nilsson too was probably justified iu his as-
sumption that, if so large a Shark had occurred regularly off the
coast of Iceland, it ought also to have been met with in Norwegian
waters. As this has not happened for more than sixty years since
attention was first drawn to the possible occurrence of the species,
we see no reason for noticing this species among the fishes of Scan-
dinavda at greater length than we have done in the present note.

Genus CARCHARIAS.

Spiracles obliterated. A transverse groove in the superior and inferior margins of the tail just in front of the

caudal fin. Valve of the spiral intestine longitudinally coiled.

The Sharks most commonly known as man-eaters ,
dreaded and detested with equal fervour by the sailor,
belong to this genus. With their elongated form of
body and their well-developed fins — especially the
usually long and scythe-shaped pectorals — they are
confirmed rovers and readily strike the eye, whether
disporting themselves at the surface of the open sea
or swimming in shallower water, sometimes close in
shore. Their insatiate voracity renders them formidable
pirates throughout their range in the tropical and tem-
perate seas. The seaman that falls overboard, in most
cases, no doubt, becomes their prey, and the incautious
bather, even on a shelving coast, may meet with the
same fate. Sailors therefore take a delight in torment-
ing these fishes in every conceivable manner when they
have succeeded in getting them on board.

During his voyage to India Sundevall made the
following notes with respect to the genus Carcharias :
“These species are commonly seen slowly following the
vessel in fine weather and light winds, especially when

“ Fiscli. Isl., p. 17.

* Prodr. Ichth. Scand., p. 115.

c Maanedskr. f. Lit., Kbhvn, Bd 7, p. 212 and Bd 9, p. 263.
d Plarjiost., p. 60, tab. 24.
e Danm. Fisk., Ill, p. 933, note.

the salt meat for daily consumption is towed behind
the ship, as is the general custom, in a sort of keg, to
be soaked. The Shark is then seen cruising about,
with dorsal fin projecting above the surface, and as
soon as anything is thrown into the water, it is at
once pounced upon by the fish. Bits of wood and
other inedible substances are readily seized, but soon
discarded. The Shark is consequently an easy catch,
but the hook should be of a suitable thickness, the line
strong and furnished with some iron links next the hook
to prevent it from being bitten off. These fishes do not
disdain any animal substance or filth. In securing any
floating object, they open the jaws over it quite lei-
surely and raise the whole head above the surface, the
mouth being situated on the ventral side; but upon
objects that sink in the water they dart with great ra-
pidity. In taking a baited hook, the Shark turns upon
its side or completely over, with the belly upwards, as
the snout would else come in contact with the line
and push the bait away. When the wind is at all

1130

SCANDINAVIAN FISHES.

high, these Sharks are never seen, presumably because
they are sluggish swimmers and cannot keep up with
the vessel. In all their actions they greatly resemble
a hungry dog or a wolf prowling in quest of food;
hence the names of chien de mer, grand chien bleu , etc.,
which are conferred upon them in several languages.
Most frequently these Sharks are attended by a little
fish known among seamen as the Pilot ( Naucrates due-
tor, see above p. 82, note d). This fish, which is about
a foot long, presents a handsome appearance, being
silvery blue with broad transverse bands of dark blue.
Seafaring folk in general suppose that it guides the
Shark to the prey in the hope of gaining its share
thereof, and this belief has been supplemented by all
kinds of romantic additions. Cuvier, who discerned
the improbability of these fabulous narratives, was of
the opinion that the companionship of the Shark and
the Pilot-fish is merely fortuitous, each of them fol-
lowing the vessel. But this is not the case. The Pilot-
fish really attends upon the Shark, but only to feed
upon its excrements, as a few writers, ancient and mo-
dern, have correctly stated. I have often had the op-
portunity on tropical seas of carefully watching Sharks
accompanied by one or two Pilot-fish, and have always
made the same observation. The Pilot-fish keeps close

to the body of the Shark, at the dorsal, pectoral, or
ventral fins, but now and then quits his post to taste
some floating object he has espied. Apparently, however,
he seldom finds anything that tickles his palate, and
soon returns to the Shark; but the moment the Shark
passes any excrementitious matter, the Pilot promptly
and eagerly makes off to secure the prize, and then
returns to its former station, from which it refuses to
be enticed for a long while, whereas, when it is hungry,
it readily swims after any small object thrown into the
water. I never succeeded in hooking a specimen. In
the Bay of Bengal I had good opportunity of observing
that large turtles are also attended by the same kind
of Pilot-fish; and Sucking-fishes ( Echeneis , see above,
p. 89, note a) accompany both Sharks and turtles for
the same reason as the Pilot, but always adhere firmly
to the body of their host, till they see any morsel of
food, when they vie with the Pilot-fish in agility, and,
their errand accomplished, immediately return to attach
themselves as before.”

These Sharks are, however, not exclusively salt-
water fishes. Some of them ascend the great rivers of
the tropics, even beyond the tidal portion of their
course. In Scandinavian waters they are strangers;
but one species has strayed to our coasts.

THE BLUE SHARK (sw. blahajen).
G ARCH ARIAS GLAUCUS.

Plate L, fig. 3.

Teeth serrated at the margins. Snout more or less conically prolongated to a length of about half that of the
head. First dorsal fin nearer to the ventral fins than to the pectoral , beginning about half-way between the tip of
the snout and the base of the caudal fina. Coloration above of a blackish or grayish blue , underneath white.

Syn. riav'rog, .ZElian., Anim. Nat., lib. I, cap. XVI, p. 22.

Galeus glaucus , Rond., De Pise., p. 378; Willughb., Hist.
Pise., p. 49. Squalus fossula triangnlari in extremo dorso,
foraminibus nullis ad oculos, Art., Ichth., Gen., p. 69;
Syn., p. 98.

Squalus glaucus , Lin., Syst. Nat., ed. X, tom. I, p. 235;
Bl., Naturg. Fisch. Deutschl., pt. Ill, p. 78, tab. LXXXVI ;
Rf/tz., Fna Suec. Lin., p. 306; Blvlle {Car char inns'),
Bull. Sc. Soc. Philom. Paris, 1816, p. 121 (+ C. cceru-
leus, ibid., efr Fne Franc., I, Poiss., pp. 90 — 92); Guv.
(subg. Oarcharias), R'egn. Anim., ed. I, tom. II, p. 126;

Yarr., Brit. Fish., ed. 1, vol. II, p. 381; ed. 2, vol. II,
p. 498; Bonap., Iconogr. Fna Ital., Pesci, tab. 133, fig. 2;
Mull., Hle ( Oarcharias , subg. Prionodon), Plagiost., p. 36,
tab. 11 ( + C. hirundaceus ex Val., p. 37); Couch, Fish.
Brit. Isl., vol. I, p. 28, tab. VI; Dum., Hist. Nat. Poiss.
(su. a Buff.), tom. I, p. 353; Barb. Boc., Cap., PeLv.
Plagiost., p. 17; Gthr, Cat. Brit. Mus ., Fish., vol. VIII,
p. 364; Mor., Hist. Nat. Poiss., tom. I, p. 329; Mob.,
Hcke, Fisch. Osts., p. 150; Jord., G-ilb. ( Carcharinus ),
Bull. U. S. Nat. Mus., No. 16, p. 22; Day ( Oarcharias ),
Fish. Gt. Brit., Irel., vol. II, p. 289, tab. CLII; Coll.,

a The distance between the beginning of the first dorsal fin and the upper transverse notches on the peduncle of the tail is in our -
young specimen 91 % of that between the same fin and the tip of the snout, which latter distance is equal to that between the anterior
angle of the pectoral fin and the beginning of the anal fin. In older Blue Sharks the last-mentioned distance is even relatively somewhat
greater.

BLUE SHARK.

1131

N. Mag. Naturv., Bd 29 (1884), p. 116; Doderl , Man.
Ittiol. Medit., fasc. II, p. 42; Lill.j., £v Norg. Fna,
Fisk, vol. Ill, p. 609.

Iii Plate L, tig. 3 we give a representation of a
young Blue Shark, 41 cm. long, from the Indian Ocean.
It was cut out of the belly of the mother-fish. The
specimen is sufficiently developed and well preserved
enough to give a better idea than a stuffed example
of the appearance of the species. The Blue Shark,
however, attains a considerable size, in ordinary in-
stances a length of 2 to 3 or 4 m.°, and the form
varies during growth. The pectoral fins become con-
siderably longer and narrower, their length being some-
times thrice their breadth. The relative size of the
eyes diminishes from about '/? to Vio °f the length of
the head. The form of the jaw-teeth (fig. 326) is
altered as shown in the figure; and the number of

specimen 3 1,/2 m. long was taken in Travemtinde Bay
and described by Walbaum6. During the previous
year a specimen is said to have been caught off Kielb
According to Cough the Blue Shark roves round the
English coast during summer, but leaves those waters
at the approach of winter.

Though the Blue Shark attracts most attention as
it swims at the surface, it also descends to a consider-
able depth (at least about 100 m.), and here is perhaps
its most congenial home. Light is apparently obnoxious
to it, for at the surface and in the upper strata of the
water, so long as the light is powerful, and when it is
desirous of fixing a steady gaze on some object, it keeps
drawing the nictitating membrane up and down over
its eyes. In the Mediterranean the young make their
appearance during May and June (Doderlein). Oppian
celebrated in verse/ the care devoted by the parents

A

Fig. 326. Jaw-teeth of the right side in a young (M) and an adult- ( B ) Blue Shark. J/2 nat. size. After Muller and Henle.

their transyerse rows, one behind the other in each
jaw, increases from 2 to 5.

In the tropical seas the Blue Shark is dispersed all
round the globe. In Europe it is common in the Medi-
terranean and at least not rare on the south coasts of
England and Ireland. Even in St. Andrews Bay (Scot-
land), according to McIntosh, it is not uncommon,
and according to Couch it strays up to the Orkney Is-
lands; but further to the north and east it is appa-
rently but an occasional visitor. In November, 1883,
Collett received a specimen 16 dm. long that had
been taken at the very head of Christiania Fjord, and
Blue Sharks have been caught in the westernmost parts
of the Baltic. At the beginning of October, 1753, a

to their offspring. When any danger threatens, the
mother (or, according to others, the father) opens its
mouth for the young to take shelter in its pharynx6 or
even in its stomach. The fry are, however, soon ca-
pable of defending and feeding themselves, for they
come into the world fully developed for leading an
independent existence, and measuring, according to a
note of van BenedknV, at least 60 cm.

The diet of the Blue Shark consists principally of
fish of all kinds. In the stomach of a specimen 6 ft.
long Couch found a large Picked Dog-fish and a Con-
ger, both bitten right in two, and a Gurnard. In an-
other he found four Mackerel, half a Garpike, and so
many Herrings, quite whole, that the fisherman sold

a Lilljeborg states as much as 71/, m.
b Cf. Bloch, 1. c.
c Cf. MSbius and Heincke, 1. c.

d Halieuticorum lib. 1. From the beginning of the third century, A. D.

e Cf. the method employed by the Chromidce of preserving their eggs and fry in the branchial cavity. Agassiz, A Journey in
Brazil , p. 222.

f Poiss. Cotes Bely ., p. 4; Mem. Acad. Roy. Belg., tom. XXXVIII (1871).

1132

SCANDINAVIAN FISHES.

them for eighteenpence ; and yet the Shark had been
hungry enough to gorge the bait. To the fisherman the
Blue Shark causes great annoyance, as it robs the long-
lines of their bait, and cuts the line in two or twines
the latter round its body, by rolling over and over,
in such a complicated manner that the task of disent-

anglement is hopeless; and it plucks the fish out of
the net and tears asunder the meshes.

The flesh of the Blue Shark is hard and has a nau-
seous smell, but it is eaten in Italy by the poor. The
only other value of the fish consists in its oily liver
and finely shagreened skin. •

Genus GALEORHINUS.

Spiracles open , though small. Peduncle of the tail without t r

Both in form of body and manner of life the ge-
nus of the Smooth Sharks closely resembles the pre-
ceding one; but they do not attain the same dimensions
as the man-eater Sharks, and the narratives related ever
since Pliny’s time of combats between divers and
Smooth Sharks are probably based on a confusion with
Blue Sharks, though the Tope is sometimes large enough
to take a substantial mouthful from the body of a

ansverse notch. Valve of the spiral intestine spirally coiled.

swimmer. The ground-colour of the body, though it
does not distinguish them in the least from several
man-eating Sharks, has given rise to the name of
Gray Sharks (Nilsson in Skand. Fauna); but this
name is more commonly applied to another genus, also
occurring in the North Sea, namely Notidanus.

Only two species are known, one from Japan alone,
the other cosmopolitan in the tropical and temperate seas.

THE TOPE (sw. HASTOIiJEN OK BETHAJEN).

GALEORHINUS GALEUS.

Plate L, fig. 2.

Inner margin of the teeth smooth , their outer margin obliquely notched , finely serrated , but with a coarser denti-
culation at the base. Snout in great part translucent and prolongated in a more or less flattened form to a length
measuring about half that of the head. First dorsal fin at least about tivice as large as the second and situated
nearer to the pectoral fins than to the ventral , the distance betiveen it and the tip of the snout being slightly more
than 2/5 of the length of the body to the beginning of the caudal fin. Beginning of the second dorsal fin some-
what further forward than that of the anal. Coloration above of a more or less light bluish gray, underneath white.

Fig. 327. Teeth of the upper and lower jawrs in a Tope ( Galeorhinus galeus ) 13 dm. long, 2/3 nat. size.

TOPE.

1133

Syn. raledg Kviuv, Abistot., De anim. hist., lib. VI, cap. XI. Galeus
cants, Rondel., De Pise., p. 377. Canis galeus, Willughb.,
Hist. Pise., p. 51 (ex Salv.). Squalus naribus ori vicinis,
foraminibus exiguis ad oculos, Art., Ichthyol. , Gen., p. 68;
Syn., p. 97.

Squalus Galeus, Lin., Syst. Nat., ed. X, tom. I, p. 234;
Blnvlle ( Galeorhinus ), Bull. Sc. Soc. Pliilom. 1816, p. 121;
Cuv. (snbg. Galeus ), R'egn. Anim., ed. I, tom. II, p. 127;
Nilss. {Squalus), Prodr. Iehth. Scand ., p. 115; Sundev.,
v. Wk., Skand. Fisk., ed. 1, p. 185, tab. 45; Couch
{Toper), Fish. Brit. Isl ., vol. I, p. 45, tab. IX; Jord.,
Gilb. {Galeorhinus), Bull. U. S. Nat. Mus., No. 16, p. 21;
Coll., N. Mag. Naturv. Chrnia, Bd 29 (1884), p. 116.

Galeus vulgaris, Flmng, Brit. Anim., p. 165; Yarr., Brit.
Fish., ed. 2, vol. II, p. 509; Kr., Damn. Fisk., vol. Ill,
p. 834; Coll., Forh. Vid. Selsk. Chrnia 1874, Tillaegsh.,
p. 207; 1879, No. 1, p. 102; Day, Fish. Gt. Brit., Irel.,
vol. II, p. 292, tab. CLIII; Lillj., Sv., Norg. Fna, Fisk.,
vol. Ill, p. 612.

Galeus eanis, Bonap., Iconogr. Fna. Ital., tom. Ill, Pesci, tab.
132, fig. 3; Mull., Hle, Plagiost., p. 57; Nilss., Skand.
Fna, Fisk., p. 714; Barb. Boc., Cap., Peix. Plagiost., p.
18; Gthr, Cat. Brit. Mus., Fish., vol. VIII, p. 379;
Winth., Naturh. Tidsk. Kbhvn, ser. 3, vol. XII, p. 56;
Mor., Hist. Nat. Poiss. Fr., tom. I, p. 317; Doderl., Man.
Ittiol. Medit., fasc. II, p. 36; Car., Prodr. Face Medit.,
vol. II, p. 509.

Galeus Linnei , Malm, Gbgs, Boh. Fna, p. 618.

The Tope, known in France as le chien de mer and
in Italy as la lamiola, attains a length of about 2 in."
The form of the body is moderately elongated for a
Shark, but varies rather considerably with age. Old
individuals are more robust, with deeper body, than
young. The greatest depth, just in front of the first
dorsal fin, measures in the young about 1/10 — 1/9, in
the old sometimes 1/1, of the length of the body. The
least depth, just in front of the caudal fin, shows less
alteration, measuring about 1/30 (3'1 — 3’4 %) of the
length of the hody or 17—18 % of that of the head
to the first gill-opening. The ordinary fusiform shape
is laterally compressed in the anterior abdominal re-
gion, further back more terete or of a rounded quad-
rangular section, the dorsal margin being depressed,
and the ventral almost plane. In front of the first
dorsal fin, however, the back is bluntly sharpened
(fastigiate or a so-called hog-back), which gives the
section of the body in front of the pectoral fins an
almost triangular form. Even the form of the head
partakes in this modification, approaching in old spe-
cimens to that of a three-sided pyramid, but with a

a From Dublin Blake-Knox states (Zoologist, Dec. 1866, p.
ft. (21 dm.) long.

6 According to Krcyer sometimes 52.

longitudinal swelling behind the eyes and a flattened
snout, the lateral margins of which form a parabola.
In younger specimens the head is flatter forward from
the occiput, shallower, and behind the eyes of a more
quadrangular section.

The length of the head to the first gill-opening is
somewhat less than 1/5 (18 — 19 %) of the length of the
body. The length of the snout from the preorbital
margin is about 4(1 — 48 %b of the length of the head.
The eyes themselves are indeed round (with round
iris), in the young with round, interiorly acute-angled
pupil, in old with elongated, slit-like pupil; but the
orbits are oblong, their longitudinal diameter measuring
in specimens 1/2 m. long about 18 — 19 %, in specimens
1 73 m. long about 12 — 13 %, of the length of the head,
the vertical diameter only 1/2 — 2/3 of the longitudinal.
The suborbital margin has been turned inwards to form
the fold known as the nictitating membrane. The use
of this membrane we have noticed in the Blue Shark.
It is for the most part shagreened, like the skin of the
body, but at the fold itself (below and at the corners of
the eyes) naked and soft. The interorbital width is
about 2/5 of the length of the head. The spiracles be-
hind tlie eyes are fairly large and distinct in young
specimens; in old they are contracted into small, elon-
gated slits and simultaneously removed farther from
the eyes, the distance increasing from 1/3 to ’/4 of the
longitudinal diameter of the orbit. Spiracular gills are
wanting or are extremely vestigial, forming a row of
10 — 11 small papillae, situated rather far in. The nos-
trils are somewhat obliquely set slits, directed inwards
and backwards from the edge of the snout, on the
under surface thereof, and their length is about equal
to the vertical diameter of the orbits. Their anterior,
overlapping margin is double, being divided into an
outer (lower) and an inner (upper) lobe, and each of
these lobes is furnished, about half-way or two-thirds of
the way along the nostril, with a small, pointed, tri-
angular flap (valvule). Their outer, somewhat expanded
corner, which lies close to the edge of the snout, is
separated from the tip thereof by a distance of about
7/10 (72 — 88 %) of the length of the same. Their inner
angle lies at a distance from the anterior margin of
the mouth that is equal to the height of the gill-

19) that he has taken Tope, which are common there, as much as 7

1134

SCANDINAVIAN FISHES.

openings or about 1/7 of the length of the head. The
cleft of the mouth is horse-shoed. Its breadth is about
equal to the postorbital length of the head. Both cor-
ners of the mouth are surrounded by an impressed
fold of skin and are sharply defined, the distance from
the tip of the snout to this posterior limit of the lips
varying between about 2/s and 3/4 (66 — 77 %) of the
length of the head, and that from the anterior margin
of the mouth to the same point being about 45 %a of
the same length. The teeth are set so unevenly in 3
or 4 rows within one another that their number in
one of these rows on each side was estimated by Kr0-
yer at 7, by Muller and Henle at 17, the former
having only counted the teeth exactly in a line with
one another, omitting those close to (within) these at
the intervening spaces. Both the upper and the lower
jaws are furnished with a thick, but narrow fold (ve-
lum) behind the rows of teeth, with lobes answering
to the dentition. The tongue is broad and fleshy, but
flat. Of the five gill-openings on each side the last
two are situated above the base of the pectoral fin.

The two dorsal fins are of rather similar form,
but differ widely in size. Their form may be described
as a triangle with the posterior corner raised above the
base and strongly prolongated. The basal length of
the second dorsal fin varies between about 63 and 53 %
of that of the first, and the anterior margin of the
former measures from about 58 to 48 % of that of the
latter, which last-mentioned margin increases with age
from about 8 to dlj2 % of the length of the body.
The distance between the first dorsal fin and the tip
of the snout is about 1/s (32 — 34 %), that between the
second dorsal fin and the same point about 2/3 (63 —
67 %), of the length of the body. The anal fin is si-
milar to the second dorsal, but somewhat smaller6 and
situated a little further back'd The caudal fin resem-
bles in form that of the Blue Shark, but is shorter,
its length from the beginning of the inferior lobe to
the extreme tip of the fin decreasing during growth
from about 24 to 22 1/2 % of the length of the body.
The inferior lobe is so deeply forked that it almost
seems to consist of two fins, an anterior, prolongated
forward so as to form a triangular lobe, behind shal-
low and of almost uniform depth, and a posterior, which

is triangular, and together with the elongated shallow
upper caudal lobe, gradually ascending behind, forms
the extreme tip of the fin. The paired fins are tri-
angular. The pectorals are of much the same form as
the dorsals, but the posterior (inner) corner is only
slightly prolongated. The distance between them and
the tip of the snout measures about 23 — 22 %, their
anterior margin about 13 — 15 %, of the length of the
body. The ventral fins, which surround the anus, have
the form of almost right-angled triangles with conti-
guous hypotenuses. These sides, which are about 1ji
greater than the others, measure about 6 — 7 % of the
length of the body. The distance from the beginning
of the ventral fins to the tip of the snout is about 50
- — 53 %, to the beginning of the pectoral fins about
27 — 30 %, of the length of the body.

The whole skin of the body — with the exception
of triangular patches behind the cloacal region and
the dorsal, the anal, and the pectoral fins — the
surface of the fins, and the cavity of the mouth, are
finely shagreenecl with small three-spined placoid scales,
so small that, when stroked from in front, the body
is smooth and slippery to the touch, but in the oppo-
site direction feels harsh and rough.

The external difference of sex is that normal in
the Sharks. In the young male represented in our
figure the pterygopodia had not yet grown beyond the
tips of the ventral fins.

The back, together with the dorsal and caudal fins
and the upper surface of the paired fins, is of a steely
gray, sometimes with a bronze lustre. The ventral
side is white, sometimes with a nacreous lustre, or
yellowish. The iris is white, with a narrow ring of
cupreous lustre round the black pupil.

The Tope is known from the Atlantic, Pacific, and
Indian Oceans. In the Atlantic it has been found from
the Cape of Good Hope to the Orkneys and the neigh-
bourhood of Bergen. In the Mediterranean and on the
coasts of France, the south of England, and Ireland, it
is common. In the North Sea it becomes less frequent
of occurrence to the north; but even in the Cattegat
it cannot be described as rare. North of Bergen it has
never been met with; but in Christiania Fjord and on
the coast of Bohuslan, as well as in Danish waters, it

a According to KR0YER sometimes 50.

b Base about 90 — 91 % of that of the second dorsal; anterior margin about 80 — 86 % of that of the second dorsal.
c Distance between the second dorsal and the tip of the snout about 98 — 99 % of that between the anal and the same point.

TOPE.

1135

is frequently taken. In Liim Fjord a specimen was
caught by Feddersen off Struer in the summer of 1876.
A female 15 dm. long was taken off Skummeslof (Hal-
land) in November, 1881, and forwarded to the Royal
Museum by Mr. A. v. Moller. Another, not much
inferior in size, was secured in the north of the Sound
during January, 1835, and was described by Sundevall
in the former edition of the present work. It is other-
wise usual that only young Tope are met with near
land and enter the Sound (Krdyer and Nilsson), pe-
netrating to the shallows round Saltholm (Winther).
Older individuals generally confine themselves, except
during the breeding season, to deeper water. Their
haunts are soon learnt by the fisherman, who finds the
snoods of his Fladdock-lines torn off, and perhaps hauls
in a Tope that has eventually been hooked, and in its
desperate struggles has twisted the line round its body,
in the same manner and with the same result as we
have just related of the Blue Shark. A Tope 13 dm.
long, which had been forwarded from Bohuslan to the
Royal Museum, had in its stomach two Haddocks about
4 dm. long, each with a hook and a bit of the snood
in the mouth. The great strength of the Tope appears
from a statement made by two fishermen who were
oat fishing for Ling in September, 1887, about seven
miles off Langesund. In 10 fathoms of water a Tope
seized the bait, and the line being strong, he towed
the boat behind him for half an hour. The fish was
a metre long and weighed 55 kilo.

In summer, from June till September, the Tope

gives birth to its young, about 20 — 40 being excluded
from the female at a time, and these already about
4 dm. long. Through the winter the young stay near
the coast, while the old retire to deep water, and ac-
cording to Yarrell they reach their full size the fol-
lowing year; but no precise information on this head
is forthcoming. Couch says that on the English coast
they are about 5 dm. long in January; but a Tope of
this size was taken off the coast of Bohuslan by Mr.
C. A. Hansson in September.

The Tope does more harm than good to the fisher-
man. The oil extracted from its liver and the fine
shagreen prepared from its skin hardly compensate
the damage it causes. Its flesh is poor, but in spite
of this it is eaten in many places. At the Danish fishing-
villages, according to Kroyer, it is boiled fresh, or
dried and sold in this condition at the markets of
Northern Jutland. According to Moreau it also finds
consumers in the French departments bordering on the
English Channel.

The Swedish name of hdstorje (lid, haj -= Shark) is
derived, according to Sundevall, either from the re-
semblance of the head to that of the Sturgeon (storen)
or from the size of the fish (stor = great, large), Swe-
dish fishermen meaning by lia or haj , when used with-
out qualification, the Picked Dog-fish. Similarly they
call the Tunny Makrilstorje (Great Mackerel). The
name of hethaj (Bait-Shark) is conferred upon it in
Bohuslan for its habit of stripping the long-lines and
Haddock-lines of the bait.

Fam. LAMNIDiE.

Two dorsal fins and one anal, the first dorsal opposite to the space between the pectoral and ventral fins. Eyes
without nictitating membrane. Spiracles obliterated or minute. All the branchial apertures situated in front of

the pectoral finsa.

This family too contains large, not to say the
largest Sharks, and has the same pelagic distribution
as the preceding one, but is not so rich in forms. Gun-
ther includes in his Catalogue only 9 ascertained spe-
cies. Most of these are characterized by large — one
genus by exceedingly large — gill-openings. Some are
formidable predatory fishes, with dentition as powerful

as that of the preceding family or even more so;
others — among them the largest Sharks — live on mi-
nute animals, and have extremely small teeth in pro-
portion to the size of the body. These differences led
Gunther to divide the family into two subfamilies,
which are represented in the Scandinavian fauna by
the following three species:

a The last-mentioned character is adduced as being generally valid and distinct, and especially serviceable in cases where the presence
or absence of a nictitating membrane cannot be controlled. Yet it happens, as in the Fox-Shark, that the character may easily lead to a
misconception, the hindmost gill-opening approximating its lower end close to the beginning of the base of the pectoral fin, and being di-
rected, as well as the penultimate opening, obliquely backwards and upwards above the said base.

Scandinavian Fishes.

143

1136

SCANDINAVIAN FISHES.

A : Teeth large or middle-sized. The gill-
openings extend only slightly, if at all,
above the middle of the sides. — Sub-
family Lamnince.

a: Length of the caudal fin about half
that of the body or even greater.

Peduncle of the tail not keeled Alopias vulpes.

b: Length of the caudal fin much less

than half that of the body. Sides
of the peduncle of the tail furnished
with a longitudinal carina in front
of the caudal fin Isurus cornubicus.

B: Teeth extremely small, numerous, of
a simple conical form. The gill-open-
ings extend up the greater portion of

the sides. — Subfamily Selachince Cetorhinus maximus.

Genus ALOPIAS.

Caudal fin usually occupying more than half the length of the body. A transverse depression at its base above
and beloiv. No lateral carina on the tail. Teeth middle-sized, of a compressed triangular form, smooth-margined.

Of this genus, which possesses, in its so singularly
elongated caudal fin, an unmistakable characteristic
among all the Sharks, only one species is known.

This is strictly a stranger to the - Scandinavian fauna,
but has been met with three or four times within the
limits thereof.

THE FOX-SHARK (sw. rafhajen).

ALOPIAS VULPES.

Fig. 328.

Body, apart from the caudal fin , of a thick fusiform shape, almost clavate , the least depth being only slightly less
or even more than half the greatest, which is contained about 4 times in the length of the body , excluding the
caudal fin. Length of the short conical snout about 1/G — 1/5 of the interorbital width. Diameter of the eyes about
half the length of the snout. Spiracles extremely small, situated in a groove exactly behind the eyes, from which
they are separated by a distance equal to or half as great again as the diameter thereof. Gill-openings middle-
sized (small, in comparison with those of the two following species ). First dorsal fin situated about half-way
between the tip of the snout and the base of the caudal fin, triangular , with the lower inferior corner but slightly
prolongated. Second dorsal and anal very small, the former situated about two-thirds of the way between the first
dorsal and the caudal and with its base in front of the perpendicular from the beginning of the anal fin. Pec-
toral and ventral fins rather large, especially the former , which are of a broad scythe- shape, their length being
about V4 of that of the body to the beginning of the caudal fin; ventral fins rather similar to the first dorsal.
Caudal fin scythe-shaped, with a triangular flap near the tip of the inferior lobe. Coloration above slaty-gray,
underneath white, these tivo colours being interspersed on the lower part of the sides.

Fig. 328. Fox-Shark ( Alopias vulpes ), l/8 nat. size. After Todd, in Brown-Goode.

Syn. NXwnn]^, Aristot., De Anim. Hist., lib. VI, c-app. X et XI.

Singe de mer, Bel., Nat., Divers. Poiss., p. 88. Vulpes,

Rondel., De Pise., p. 387. Squalus cauda longiore quam
ipsnm corpus, Art., Ichthyol., Gen., p. 68; Syn., pag. 96.

FOX-SHARK.

1137

Squalus Vulpes , Gmel., Syst. Nat. Lin., ed. XIII, tom. I, p.
1496; Mitch., Trans. Phil., Lit. Soc. N. York, vol. I (1815),
p. 482; Blnvlle ( Carcharinus ), Bull. Sc. Soc. Philoin.
1816, p. 121; Cuv. (subg. Care h arias)-, Rcgn. Anim., ed. 1,
tom. II, p. 126; Yarr., Brit. Fish., ed. 1, vol. II, p. 379;
Bonap. ( Alopias ex Rafin.), Iconogr. Fna Ital., Pesci , tab.
134, fig. 1; Mull., Hle ( Alopecias ), Plagiost., p. 74; Couch
(G1 archarias'), Hist. Fish. Brit. Isl., vol. I, p. 37, tab. VII;
Dum. (Alopias), Hist. Nat. Poiss. (sit. a Buff.), tom. I, p.
421; Barb. Boc., Cap., Peix. Plagiost.., p. 14; Gthr (Alope-
cias), Cat. Brit. Miis., Fish., vol. VIII, p. 393; Coll., Fork.
Vid. Selsk. Clirnia 1874, Tillasgsh., p. 208; 1879, No. 1, p.
102; Mor. (Alopias), Hist. Nat. Poiss. Fr ., tom. I, p. 287;
Doderl., Man. Ittiol. Medit., fasc. II, p. 52; Day, Fish. Gt.
Brit., Irel., vol. II, p. 300, tab. CLVII; Petersen, Vid.
Meddel. Naturh. For. Kbhvn 1884, p. 160; Day, Fish. Bidia
(Suppl.), p. 810; Lillj., Sv., Norg. Fna, Fisk., vol. Ill, p.
627; Car., Prodr. Fna ? Medit., vol. II, p. 507.

Alopias Macrourus, Rafin., Caratt. Ale. Nuov. Gen., Spec., p. 12.

The Fox-Shark attains a length of at least 5 m.
(Moreau) and a weight of 300 kilo. (Doderlein.)
Buckland cites" a specimen, taken by Mackerel-fisher-
men from Yarmouth, 44 dm. long, 1 772 dm. in girth,
and weighing 500 lbs. (2263/4 kilo.), and another* *
measuring 45 dm., taken by Herring-fishermen from
Folkestone.

Its geographical extension includes the Atlantic,
Indian, and Pacific Oceans. Between the Cape of Good
Hope and the Shetland Islands lies its Atlantic home,
and in the Mediterranean it has been known time out
of mind. Its comparatively large eyes suggest that it
is more of a deep-sea fish than the other members of
the family; but throughout its range it attracts atten-
tion at the surface in its pursuit of schooling fishes.
The long caudal fin, which often projects out of the
water on such occasions, has not only conferred upon
the species its name, but has also given rise to the
most fabulous accounts of its ravages. With its tail
it lashes the waves and deals blows on the surface of
the water that resound like a cannon-shot. Dolphins
frightened by this noise have been seen to take to pre-
cipitate flight. It is quite credible that a whale may
now and then have shown the same timidity. But from
these observations, and owing to a confusion with the
grampus ( Orca gladiator), whose dorsal fin also appears
above the surface, it has been concluded that the Fox-
Shark is the relentless persecutor of whales and dol-
phins, though its dentition is far too weak for the
achievements ascribed to it. “It, is sometimes,” says

Buckland, “called the ‘thresher shark,’ because it is
said to inflict chastisement on the whale, though I can
find no real authority for this being the case; nor can
I see why the shark should flog the whale. The tail
of the skate can, as I know to my cost, inflict severe
wounds, and the tail of the sting-ray contains a, dagger
which is even more formidable still; I fancy, therefore,
that the tail of the fox-shark is used by the owner as
a weapon of defence, for if he chooses to use it, it,
would act like a waggoner’s whip. Again, I think it
just possible that, whereas we all know we can drive
fish by long poles dashed suddenly into and about in
the water, so the fox-shark may use his tail to splash
about and drive the frightened herrings or mackerel
into the position where they will be most handy for
him to catch and swallow. I should imagine an un-
frightened mackerel to be a difficult thing to catch
when swimming at liberty in the sea. If, however,
the fox-shark comes lashing about with his tail, he and
his comrades would become confused, and while making
up their minds what to do and where to go, they
would suddenly find themselves going down the shark’s
throat. One thing is quite certain, his long tail would
enable him to turn smartly round the corners and as-
tonish unsuspecting natives.”

In the stomach of a Fox-Shark 41 dm. long Buck-
land found 27 Mackerel; and Herrings are devoured
with equal avidity. Accordingly it happens often enough
that the Fox-Shark gets entangled in the drift-nets shot
for Mackerel and Herrings or in the tackle of the Pil-
chard-fisherman. On the other hand, it never takes a
hook, which apparently proves that on the whole it is
less rapacious than the other Sharks.

In the Mediterranean the Fox-Shark gives birth to
its young during summer. In August Doderlein se-
cured a young specimen 27 cm. long. During October
Buckland found in a large female from Folkestone an
egg 76 mm. long and 63 mm. broad, on its backward
passage through the oviduct, while the ovaries con-
tained thousands of smaller eggs, from the size of a
hazel-nut to that of a pin’s head.

The Fox-Shark is, as we have mentioned, a stran-
ger in Scandinavian waters. A female 429 cm. long
was taken near Bergen on the 31st of August, 1868,
and is described by Lilljkborg. In Bunde-Fjord, about

“ Land and Water, July 6, 1869, p. 608.

* Nat. Hist. Brit. Fish., p. 221.

1138

SCANDINAVIAN FISHES.

a league from Christiania, a specimen 16 dm. long was
caught, according to Collett, on the 1st of February,
1878. Petersen cites a specimen “nearly 4 Danish
ells (83/4 ft-) long,” that Avas cast ashore in December,
1886, off Tidsvilde on the north coast of Zealand.
Kroyer Avas besides informed on trustworthy authority

that the Fox-Shark had once been met Avith on the
Avest coast of Jutland.

The flesh of the Fox-Shark is “moderately good,”
says Doderlein, “though rather hard;” but at Cette
it is sold, according to Moreau, under the name of
White Tunny ( TJion blanc).

Genus ISURUS.

Peduncle of the tail marked above and below ivith a transverse depression and furnished with a longitudinal Ca-
rina on each side. Caudal fin crescent-shaped, its length much less than half that of the body. Teeth middle-
sized, entire-margined , pointed., in adult specimens witha or without lateral cusps at the root.

By its smooth-margined teeth, in the upper jaw
too of a narrow triangular form or even subulately
pointed, this genus is distinguished from Carcharodon,
the most dreaded Shark of the Mediterranean and the
tropics, often confounded Avith the Blue Sharks under
the name of man-eater. In both these genera the

spiracles, at least of adult specimens, are very minute
or even obliterated.

In deference to the current rules of nomenclature
the generic name of Isurus, coined by Rafinesque in
1810c, must supersede the more generally employed
Lamna, Avhich Avas first proposed by Cuvier in 1817''.

THE PORBEAGLE (sav. habranden).
ISURUS COKNUBICUS.

Plate LI, fig. 1.

Snout conically pointed , its length e about /10 (29 — 31 %) of that of the head (to the first gill-opening ), which
measures about x/5 (18 — 22 %) of that of the body. Teeth of a narrow triangular form, with a cusp on each
side of the expanded root. Third tooth, counting from the middle, smaller than those beside it. Distance betiveen
the pectoral fins and the tip of the snout about i/5 (80 — 83 %) of that betiveen the first dorsal fin and the same

point. Coloration above slaty-gray, underneath white.

Syn. Porbeagle, Borlase, Nat. Hist. Cornwall (1758), p. 265,
tab. 26, fig. 4. Haae-Brand, Str5m, Sondm. Beskr., part. I,
p. 281. Touille-Boeuf ou Loutre de mer, Duh., Tr. Peches,
part. II, sect. IX, tab. XX, fig. 4. Beaumaris Shark , Penn.,
Brit. Zool., ed. 1776, vol. Ill, p. 104, tab. XVII. Le
Nez, Broosson., Mem. Acad. Sc. Paris 1780 (ed. 1784), p,
668. Haabrand, Ascan., Icon. Eer. Nat., tab. XXXI.

Squalus glaucns ( Haabrandsunge ), Gunner., N. Vid. Seisk. Skr.,
Trondhj., vol. IV (Kbhvn, 1768), p. 1 (nom. erron.);
StrSm ( Haamcer ), N. Sami. N. Vid. Seisk. Skr. Trondhj.,
vol. II (Kbhvn 1788), p. 335, cum tab.; Retz., Fna Suec.
Lin., p. 307.

Squalus cornubicus , Gmel., Syst. Nat. Lin., ed. XIII, tom. I,
p. 1497; Cuv. (subg. Lamna), Regn. Anirn., ed. 1, tom. II,
p. 127; Nilss. (Squalus), Prodr. Ichtli. Scand ., p. 116;
Id., Observ. Ichthyol., p. 13 (Disp. Lund, 1835); Yarr.,
Brit. Fish., ed. 1, vol. II, p. 384; Bonap. (Lamna), Iconogr.
Fna ltal., Pesci, tab. 134, fig. 2; Sundev., v. We. (Squalus),
Skand. Fisk., ed. 1, p. 135, tab. 30; Mull., Hle (Lamna),
Plagiost., p. 67 ; Gray (Isurus), Cat. Chondropt. Fish.
Brit. Mus ., p. 58; Kr. (Lamna), Damn. Fisk., vol. Ill,
p. 852; Nilss., Skand. Fna, Fisk., p. 718; Dum., Ilist.
Nat- Poiss. (su. a Buff.), tom. I, p. 405; Gthr, Cat.
Brit. Mus., Fish., vol. VIII, p. 389; v. Ben., Poiss. Belg.,

a Subgenus Lamna.
h ,, Oxyrhina.
c Caratt. Ale. Nuov. Gen., Spec., p. 11.
d Regn. Anim., ed. 1, tom. II, p. 126.

e According to Malm this length, “measured along the surface” (not, as here, in a straight line), may amount to -/- of that
of the head.

PORBEAGLE.

1139

p. 8 (Mem. Acad, Roy. Sc. Belg., tom. XXXVIII, 1871);
Coll., Forli. Vid. Selsk. Clirnia 1874, Tilltegsli., p. 208;
Malm, Gbgs, Boh. Fna, p. 618; Mob., Hist. Nat. Poiss.
Fr., tom. I, p. 296; Doderl., Man. Ittiol. Medit., fasc.
II, p. 60.; Mela, Vert. Fenn ., p. 365, tab. X; M5b., Hcke,
Fiscli. Osts., p. 151; Jord., Gilb., Bull. U. S. Nat. Mus.,
No. 16, p. 30; Day, Fish. Gt. Brit., Irel., vol. II, p. 297,
tab. CLVI; Storm, Norsk. Vid. Sels. Skr. 1883 (Trondhj.
1884), p. 44; Lillj., Sv., Norg. Fna, Fisk., vol. Ill, p. 620;
Car., Prodr. Face Medit., vol. II, p. 505.

Squalus nastts, Walb., Ichthyol. Art., Gen., p. 517.

Squalus monensis , Shaw., Gen. Zool., vol. V, p. 350; Cuv.,
1. c. ; Yarr., 1. c., p. 387.

(?) Isurus Oxyrinchus , Rafin., Caratt. Ale. N. Gen. e Spec.,
p. 12, tab. XIII, fig. 1 (fide Doderl., figura tainen pessima
fere potius ad Isurum Spallamanii referenda).

Fig. 329. Teeth of the upper and lower jaws in a Porbeagle (Ism

teeth behind the gap (diastema) on tl

Squalus Selanonus, Leach, Mem. Wern. Nat. Hist. Soc., vol. II,
p. 64, tab. II, fig. 2; Flmng (Squ. selanoneus), Brit. Anim.,
p. 168 = Selanonius Walkeri, p. 169.

The Porbeagle ranks among the larger Sharks. Its
ordinary length in Scandinavia, however, does not ex-
ceed 17,-27, m. In the Mediterranean it attains a
length of at least 6 m.“

The body is almost terete, slightly compressed,
rounded above and below, and pointed at each end.
The greatest depth varies between about 1/6 and x/7 of
the length. The skin is everywhere rough as a tile
when stroked from behind, but not exactly rasping; in
the opposite direction it is soft to the touch. This is

due to the small, dense scales, which in a Porbeagle
13 dm. long are only x/2 mm. broad. On the upper
parts of the body they are semicircular and thick, with
5 elevated, convex s trial, the ends of which project at
the margin of the scale, forming rounded denticulations.
On the under side of the body they have only 3 such
ridges. On the tins they are set with equal denseness,
but are smooth, as it were worn.

The head, like the body, is without prominent side-
margins. The snout in front of the eyes and mouth
is conical, its depth at the mouth being almost equal
to its length. It is densely punctated (tig. 330) with

a

us cornubicus) 213 cm. long, ]/3 nat. size; a, the first three pairs of
: right side of the upper jaw, nat. size.

distinct, impressed, muciferous dots, the mouths of
Lorenzinian ampulla?, distributed in 6 broad, multiplex
rows, which are separated by the same number of im-
punctate stripes, one in the median dorsal line, one
vertically below this, and two on each side, all con-
verging at the tip of the snout, which is also without
punctures. Below the eye lies another row of similar
dots. The eyes are round, closely bounded by the
slightly oblong orbital margins. Their diameter is some-
what less than V3 (27 — 31 %) of the length of the snout.
In a dead specimen the pupil appeared to be somewhat
oblong and obliquely set. There is no nictitating mem-
brane. The nostrils are ^-shaped and transversal, with

“ Bonaparte even states “12 up to 20 or 24 feet,” perhaps confounding the species with Carcharodon Rondelelii.

1140

SCANDINAVIAN FISHES.

a very small flap (valvule) at the anterior margin.
They lie so far in front of the perpendicular from the
preorbital margin that the distance between them and
the tip of the snout is about 80 — 90 % of that between
the mouth and the same point, which latter distance
is rather less than or sometimes equal to the length
of the snout. The mouth is large, its breadth at the
corners being somewhat more than 1j 3 of the length of
the head. The cleft of the mouth is sharply curved,
so that the lower jaw is rather long, and the distance
from the corners of the mouth to the middle of the
anterior margin of the upper jaw is about 3/10 of the
length of the head. Externally no dermal folds are
visible on the jaws, for the small creases that lie above
and below each corner of the mouth are concealed by

Fig. 330. Head of a Porbeagle (Isarus cornubicus) 213 cm. long,
seen from below, ft nat. size.

the skin of the upper jaw. The teeth (tig. 329) are
set in three rows. The anterior ones are the largest,
measuring in a specimen 24 dm. long somewhat more
than 12 mm. or 1/3 of the diameter of the eyes. They
are set rather far apart, of a narrow triangular form or
conical, terete on the inner, but somewhat flattened on
the outer side. The base of the broad root is emarginate,
so that the tooth apparently has two fangs. In old
specimens the root has a small conical cusp on each
side; but this cusp is wanting in young specimens,
sometimes even at a length of 9 dm. In both jaws
we And at the extreme front a small gap,, without
middle tooth. The upper jaw has 14 or 15 (1G) teeth
in each row on either side, the number being alike
both in young and old. The first two teeth are large,

the third is rather small — and behind this lies a wide
gap ( diastema ) — the fourth and the following ones are
again large, but gradually decrease in size, the hind-
most teeth being comparatively minute. In the lower
jaw there are 12 or 13 (14) teeth on each side. The
gap at the extreme front is somewhat wider than in
the upper jaw. The first and second teeth are the
largest, the others decreasing in size behind. All the
teeth are slightly recurved.

The spiracle of the Porbeagle is extremely small,
resembling a hole pierced with a coarse pin. It lies
exactly behind the eye, separated therefrom by a dis-
tance of twice the diameter thereof or somewhat more
in large specimens. It is consequently situated rather
further back than in most of the other Sharks, and has
many times defied detection, being probably obliterated.
All five branchial apertures lie in front of the pectoral
tin, the lower end of the hindmost opening terminating
close below the margin of this fin. This opening is,
however, obliquely set, being considerably approximated
below to the penultimate aperture. In the dermal fold
that covers each opening, lie three parallel cartilaginous
rays (radiating extrabranchial cartilages, see above),
originating from the middle of each arch and extend-
ing to the edge of the opening.

The body itself is almost terete, with indistinct
lateral line, which is best marked in dried skins. From
the temporal region, above and behind the eyes, the
line forms a slight curve above the gills and then pro-
ceeds in a straight direction to the lateral carinae of
the tail. These occupy, immediately in front of the
caudal tin, one-fifth of the length of the body behind
the head. They have an almost sharp, curved outer
margin, and are so large that their breadth is 2/3 of the
greatest breadth of the body, arid that the tail just in
front of the caudal grooves is more than four times as
broad as deep. The caudal grooves above and below are
deep and clothed with finer scales than the rest of
the skin.

The first, dorsal tin is rather large, its base mea-
suring about V9, the length of its anterior margin 1/7
— V6, and its height about •■/ 8, of the entire length of
the body. Its beginning lies at a distance from the
tip of the snout measuring about 30 — 32 % of the length
of the body. In form it resembles an almost equilateral
triangle, with a large rounded incision above the elon-
gated posterior angle. The posterior dorsal and the
anal fins are almost similar in form and size, and lie

PORBEAGLE.

1141

opposite to each other — the former, however, a little
further back, at a distance from the first dorsal measur-
ing' about 28 — 26 % of the length of the body. They
are rather small, their bases measuring about V7 — Ye °f
that of the first dorsal fin. Their anterior (upper) angle
is rounded, the posterior acutely elongated. The distance
between the anal fin and the tip of the snout is nearly
2/3 of the length of the body. The superior lobe of
the caudal fin, which is of the crescent-shaped form
that characterizes most of the members of this family,
measures about V4 of the remaining length of the body,
and is strongly curved in an upward direction, with a
pointed and exceedingly small terminal lobe. The in-
ferior lobe, on the other hand, is larger than in most
of the other Sharks, at, least somewhat more than half
as long as the superior. The pectoral fins are from
1 Yjj times to nearly twice as long as the anterior mar-
gin of the first dorsal fin, and their length is rather
more than 7c — Vs °f the entire length of the body or
about equal to the distance from their base to the cor-
ner of the mouth. Their breadth is somewhat more
than half their length. The ventral fins are small, with
the outer angle obtuse and the posterior acute. Their
length throughout the inner margin (the base together
with the posterior lobe) is about S 1/2 % of the length
of the body. They are set a little behind the middle
of the body.

Besides the pterygopodia of the males — which in
a specimen 22 1/i dm. long were 247 mm. in length
and 25 mm. in diameter — Sundevall remarked that
it appeared to him as if another external sexual cha-
racter were expressed in the form of the second dorsal
and the anal fins, which seemed to have the pos-
terior tip much more elongated in the males than in
the females.

The coloration is above of a blue-black gray, un-
derneath white, the posterior angles of the dorsal fins
being also whitish. The iris is dark brown.

The Porbeagle is somewhat variable in form. In a
number of cases, both male and female, the body is
rather slender (greatest, depth 1/1 — 1/e of the entire
length), the dorsal fin comparatively high and rectilinear,
the snout short and narrow (length of the snout in adult
specimens about 1/5 of the distance between its tip and
the pectoral fins), the inferior caudal lobe % as long as
the superior, or even more, and the hindmost gill-
opening distinctly oblique. This form is apparently
the commoner. Other specimens present the appearance

shown in our figure (Plate LI, fig. 1). They are thicker
(greatest depth up to ;1/5 °f the entire length), with
somewhat larger head and thicker, longer snout (length
of the snout up to 1/i of the distance between its tip
and the pectoral fins), with the inferior caudal lobe little
more than half as long as the superior, and with the
hindmost gill-opening somewhat, less oblique. This latter
form seems in general to be characteristic of youth.
Our figure was drawn in Bohuskln from a young spe-
cimen 13 V3 dm. long. But that this form has repre-
sentatives even among full-grown Porbeagles, seems
probable from the descriptions given by the authors
cited above (in the synonymy) of Squalus monensis,
whereof Yarrell mentions a specimen 29 dm. long.

The remaining alterations of growth consist prin-
cipally in the relative reduction with age of the snout
and eyes and the furnishing of the teeth with basal
cusps, which are wanting in the smallest specimens, but
in Porbeagles 13 dm. long are already quite distinct.

The Porbeagle is the only one among the large
Sharks that appears with any frequency in the south
of the Cattegat; but it, is far from common on the coasts
of Sweden. It- occasionally makes its way through the
Sound into the Baltic, and it has been met with, ac-
cording to Mela, even in the vicinity of Aland. Along
the Norwegian coast it, occurs up to Finmark, where it
is found rather often, according to Collett, except to
the extreme north. Its ordinary name in Norway, as
in Bohuslan, is ffdbrand; but it, is said to be some-
times coupled with the Greenland Shark under the name
of Hdmcir. As it had no Danish name, Kroyer called
it Sillhaj (Herring Shark). The Porbeagle is dispersed
throughout the North Sea, though it is not so common
there as on the west coasts of the United Kingdom and
France; and its greatest development seems to be at-
tained in the Mediterranean. On the other side of the
Atlantic it has been found off the east coast, of the
United States; and according to Muller and IIenle’s
determination of the jaws sent, by Burger from Japan
to the Museum of Leyden, the Porbeagle also occurs in
the Pacific off the Japanese coasts. Haast includes
the species among the fishes of New Zealand.

The appearance of the Porbeagle is even more re-
pulsive than that of other Sharks. It emits an ex-
tremely disagreeable, fetid smell, and the surface of the
body, which is yielding and at several spots flabby,
acquires a dirty look from the secretion of tough mu-
cus. This is especially copious at all the orifices, and

1142

SCANDINAVIAN FISHES.

it was this uncleanness, which is shared by the whole
phalanx, that suggested the ancient name of Squalus%
Furthermore, the profile of the Porbeagle indicates far
more stupidity in combination with ravenousness than
that of other Sharks. The Porbeagle is said to be a
more rapid swimmer than most of them, a statement
apparently borne out by the powerful caudal fin and
the keeled, robust tail, which in nearly all fishes is the
principal organ of locomotion.

The food of the Porbeagle consists chiefly of fishes
and Cephalopods. The Herring-shoals in particular
suffer greatly from its depredations; and several Por-
beagles often join company in the chase. Like the
preceding species these Sharks too sometimes become
entangled in Herring or Pilchard nets. Deep-sea lines
do not escape their ravages (Thompson and Day); and
many a narrative relates how the Porbeagle dashes up
to secure the Whiting, Cod, or other fish which the
fisherman has hooked.

Of the breeding of the Porbeagle not much is
known. The young are considerably developed at birth.
The foetus described by Gunnerus, which was taken
out of the mother-fish in summer, measured “nearly
three-quarters of a Zealand ell’’ (47 cm.) in length.

Sundevall examined a young specimen 74 cm. long,
and found that every external trace of the navel, which
is situated in the Sharks between the pectoral fins, had
disappeared. Hence he concluded that the length of
the young at birth must be between these two dimen-
sions. In a female cited by Pennant only two foetuses
were found.

The Porbeagle is not much sought after by Scan-
dinavian fishermen, for the liver is small and lean, with
a scanty yield of oil, and it is only for this sake that
the large Sharks are taken. By accident, however, or
in the absence of better fish, the Porbeagle is caught
by the Norwegian Shark-fishermen. In the south of
the Cattegat and in the Sound it is scarcely ever taken
except when it entangles itself in the nets set for other
fishes. The skin is fairly well adapted for the polishing
of articles of fine workmanship, in wood or horn for
instance, and it is also applied to this purpose. The
hard scales "withstand a considerable amount of wear.

The flesh of this species, like that of many other
Sharks, has a disagreeable smell, and it is hardly eaten
in Sweden; but in Italy and Spain it enjoys a better
reputation.

(Sundevall, Smitt.)

Genus CETORHINUS6.

Teeth numerous , small, conical. Branchial arches furnished with baleen-like , fine , long , and dense gill-rakers.
Anterior gill-openings extended across the greater part of the sides of the bodg. Tail with transverse grooves in
front of the caudal fin and with the sides longitudinally carinated. Caudal fin crescent-shaped.

The genus of the Basking Shark occupies within
the Lamnoid family a position analogous to that of the
Hounds in the preceding family; but in the Basking
Shark, the solitary species of the genus, the reduction
of the jaw-teeth has advanced still further, these organs
having scarcely retained any function, at least in the
capacity of Shark-teeth. The genus consequently does
not belong to those commonly known as predatory
fishes; but it contains the most gigantic piscine forms.
As we have mentioned above, it shares with another
genus of huge fishes (Rhino don) a manner of life si-
milar to that of the large tvhales. Notwithstanding its
own great size, it lives exclusively on minute creatures,

which it strains off from the seawater by means of its
gill-rakers. This manner of procuring food calls for a
wide mouth and a capacious pharynx, with room for
the filtering apparatus; and herewith is connected the
great width of the branchial arches and apertures. In
the remaining characters we easily recognise a form
akin to the Porbeagle. The most important external
difference to be found on comparison is that in the
Basking Shark the second dorsal fin stands entirely in
front of the perpendicular from the beginning of the
anal fin or nearly so.

The name of Cetorhinus means Whale-Shark, and
has reference in the first place to the dimensions of

a From the Latin squalor, filth.
b Blnvlle, Bull. Soc. Philom. P.aris, 1816, p. 121.

SHARKS.

1143

THE BASKING SHARK (sw. brcgden).

CETORHINUS MAXIMUS.

Fig. 331.

Form of the head in old specimens fairly normal , ivith short , bluntly pointed , conical snout , in the young ( up to
a length of 3 — 5 m.) contracted in front of the mouth into a semicylindrical, inferiorly flat forepart ( orbitorostral
portion ), ivith dors ally pointed tip of the snout. The small spiracles situated high up, just behind the perpen-
dicular from the corners of the mouth or further back , sometimes about lialf-way between the eyes and the top
of the first gill- opening. Coloration above of a brownish or bluish black, underneath lighter, shading into white.

The soft snout , with its numerous pores, reddish.

Fig. 331. The Basking Shark ( Cetorhinus maximus ): A and B, a young specimen in the Museum of Genoa University, after Pavesi, 1 , 3
nat. size; C , the specimen described by Blainville ( Squale pelerin, Nov. 21, 1810, Dieppe), about */54 nat. size, after Todd’s

copy of Blainville’s figure.

Scandinavian Fishes.

144

1144

SCANDINAVIAN FISHES.

Syn. Brygde , Strom, Sonelm. Beskr., pt. I, p. 273. Beinhaakall,
Olafs., Reis. Isl., p. 988.

Squalus mawimus , Gunner., Trondhj. Selsk. Skr., vol. Ill, p.
33, tab. II; vol. IV, p. 14, tab. IV, fig. 1 ; Lin., Syst.
Nat., ed. XII, tom. I, p. 400; Penn. ( Basking shark),
Brit. Zool., vol. Ill (ed. 1776), p. 89, tab. XIII; (?) Fabr.,
Fna Groenl., p. 130; Mitch., Trans. Litt., Phil. Soc., N.
York, vol. I, p. 486; Cuv. (subg. Selaclie), R'egn. Anim.,
ed. 1, tom. II, p. 129; Fab., Fisch. Isl., p. 20; Nilss.
(Squalus), Prodr. Iclithyol. Scand., p. 114; Yarr. (Se-
lachus), Brit. Fish., ed. 1, vol. II, p. 396; Mull., Hle
(Selache), Plagiost., pp. 71 et 191; Dek. ( Selachus ), Zool.
N. York, pt. Ill, p. 357, tab. 63, fig. 208; Kr. (Selache),
Damn. Fisk., vol. Ill, p. 932; Nilss., Skand. Fna, Fisk.,
p. 720; Couch, Fish. Brit. Isl., vol. I, p. 60, tab. XIV
(+ Polyprosopus Rashleighanus, p. 67 + Polypr. macer , p.
68, tab. XV et vol. IV, p. 421, tab. XV*); Dum., Hist.
Nat. Poiss. (su. a Buff.), tom. I, p. 413, tab. 3, fig. 18;
Stor. (Selachus), Mem. Amer. Acad. Arts, Sc., n. ser., vol.
IX, p. 229, tab. XXXVII, fig. 3; Gthr (Selache), Cat. Brit.
Mus., Fish., vol. VIII, p. 394; Strp (Selachus), Overs. D.
Vid. Selsk. Forh. 1873, p. 47, tab. II; Pav., Ann. Mus. Civ.
Genova, vol. VI (1874), p. 5, tabb. I — III; vol. XII (1878),
p. 348, tab. Ill; Coll. (Selache), Forh. Vid. Selsk. Chrnia
1874, Tillsegsh., p. 209; (Selachus) ibid. 1879, No. 1, p. 103;
Gerv., P et H. (Squalus 1. Cetorhinus), Journ. Zool., tom. V
(1876), p. 319; Ltkn (Selachus), Vid. Meddel. Naturh. For.
Kbhvn 1879 — 80, p. 62; Mor. (Selache), Hist. Nat. Poiss.
Fr., tom. I, p. 305; Doderl., Man. Ittiol. Medit., fasc. II,
p. 70; Day, Fish. Gt. Brit., Irel., vol. II, p. 303, tab. CLVIII,
fig. 1; Lillj., Sv., Norg. Fna, Fisk., vol. Ill, p. 633.
Squalus Gunnerianus, Blnvlle, Journ. Phys., tom. LXXI (1810),
p. 256, tab. II ( + Squ. pelegrinus + Squ. Homianus, p.
257); Id., Squ. Pelerin, Ann. Mus. D’Hist. Nat., tom. XVIII
(1811), p. 88, tab. 6; Id., Cetorhinus Gunneri + Cet. Pere-
grinus + Cet. Sliavianus + Cet. Homianus (?), Bull. Sc. Soc.
Philom. 1816, p. 121.

Squalus isodus, Maori, Att. Accad. R. Sc. Borbon., Nap., vol. I
(1819), p. 55, tab. I, fig. 1; tab. II, fig. 2 (+ Squ. ro-
stratus, p. 76, tab. 1, fig. 2). Squalus (Cetorhinus) rostratus ,
Cornish, Zoologist, vol. XXVIII (1870), p. 2259.

Squalus elephas, Lesueur, Journ. Acad. Nat. Hist. Philad., tom.
II (1882), p. 343.

The Basking Shark attains a length of at least about
13 or 15 m. Larger specimens are cited; but the ac-
counts of these do not admit of satisfactory verification.
Gunnerus states, for instance, that specimens have been
met with measuring 16 fathoms (30 m.) in length
and 3 fathoms (oV2 m-) in breadth. A specimen of
this size would yield over 28 hectolitres of liver; and
according to a statement in Collett a Basking Shark
was harpooned in West Fjord about 1868 that con-
tained 23 tunnor (27 hectolitres) of liver. Nilsson
was informed by the fishermen on the west coast of
Norway ivith whom he conversed in 1826, that they
harpooned Basking Sharks every year considerably more

than 40 ft. (12 m.) long, and that the liver of each
fish filled 10 — 12 tunnor. The largest Basking Sharks
they had seen, had yielded 14 tunnor of liver.

Apart from the above-mentioned characters, the
Basking Shark in form of body rather closely resembles
a Porbeagle, being of a terete, fusiform shape, on the
back rather flat, behind with some lateral compression,
though broadened by the lateral carinse of the tail,
which render the breadth of this part greater than
the depth. The greatest depth of the body measures
in young specimens, according to Pavesi, about 10 — 12 %
of its length, in old, according to Blainville, about
18 %, according to Doderlein as much as 22 %, of
the same.

The different form of the head in the young and
old we have remarked above. The alteration is essen-
tially due to the position of the cleft of the mouth,
which in the young is transverse, as in the Rays, in
the old curved, as in other Sharks. The corners of the
mouth, which during youth are directed straight out-
wards, form the rounded protuberances on the sides of
the head behind the eyes. In the jaws the recurved,
conical, but sharp-edged, pointed teeth are set in 4 — 7
transverse rows, but are so small that they “are often
concealed by the gums and accumulated filth’’ (Gun-
nerus). In a Basking Shark 39 dm. long they are
3 mm. high, according to Lilljeborg. Storer counted
1,400 teeth in the lower jaw. The nostrils lie on the
under surface of the snout, but close to either side-
margin, much nearer to the eyes than to the tip of the
snout. The relative length both of the snout and the
entire head is considerably greater in the young than
in the old. In the former, according to Pavesi, the
length of the snout is about 1/10 or 1/i3 of that of the
body, in the latter 1/30 of the same or less. The length
of the head to the first gill-opening is in the former
about V4 — V5, in the latter at most less than 1/G, of
the length of the body". The eyes, which are round
and small, undergo, as usual, the same alteration of
growth. According to Pavesi the diameter of the orbit
in the young is about 1/6 of the breadth of the head
at the base of the snout; according to Storer this dia-
meter measures in an older specimen '/8 °f the inter-
orbital width. The gill-openings extend across so con-
siderable a portion of the sides (the throat) that the
distance between the tops of the first pair is only twice

According to Storer the length of the head in a specimen 9 m. long was 15 7 % of that of the body.

BASKING SHARK.

1145

the diameter of the orbits, between those of the second
pair three times the same diameter, of the third pair
live times, of the fourth pair seven times, and of the
fifth pair nine times, in each case according to Storer’s
measurements. The gill-rakers (fig. 332) have claimed
a special chapter in literature. They are long and line,
apparently corneous, transversal setae, uniserial on the
first and last branchial arches, biserial on the others.
Even Gunnerus described them" as forming a kind of
strainer; and a very characteristic indication of their
structure is his opinion that, it was by clinging to
them that the prophet Jonah escaped being engulfed in
the maw of his captor — a Basking Shark, not a whale,
being the monster that swallowed this remarkable man.
In 1867 6, without knowing what the objects were
or whence they had come, but guided by their micro-
scopical texture, Hannover determined some dry frag-
ments of these gill-rakers, preserved in the Museums of
Copenhagen, Kiel, and Christiania, as a kind of squa-
mous or spinous growth belonging to some Ray. The
same texture, answering to that of dentine, tvas detected
by v. Beneden in 1871° in similar substances from the
Antwerp Crag, and he assigned them to a Ray otherwise
unknown, which he called Hannover a aurata. Steen-
strup (1. c.) and, after him, Gervais (P. and H., 1. c.)
have since maintained both that these gill-rakers afford
one of the most important characters of the genus Ce-
torhinus, and that v. Beneden ’s find carries the existence
of the genus back at least to the Tertiary epoch.

The fins are very like those of the Porbeagle; but
their outer (posterior) margin is less incurved. The first
dorsal begins at a distance from the tip of the snout
measuring about 40 — 36 %, and its base occupies about
10—9 * i/2 of the length of the body. Its height some-
times measures, according to Storer, 14V2 % of the
length of the body. The distance between the two
dorsal fins is about 1 81/2 — 1 9 % of the said length.
The second dorsal and the anal fins are about 1/s as
large as the first dorsal. The superior caudal lobe
measures about 19 — 21 % of the length of the body,
and the inferior lobe is about 2/3 as long. The median
breadth of the caudal fin, from the upper transverse
groove, is about equal to the length of the base of the
first dorsal. A little within the tip the under margin

of the superior lobe has the same notch as in the Por-
beagle. The pectoral fins are rather more than twice
as long as broad. Their length is about 18 % of that
of the body. The ventral fins, according to Pavesi’s
figures, are obliquely quadrangular. Their length is
about the same as the breadth of the pectorals. The
preabdominal length is about equal to the distance be-
tween the beginnings of the two dorsal fins.

The thick skin is armed with sharp and strong
spines — small in comparison to the size of the body,

Fig. 332. Gill-rakers of the Basking Shark. A, an excised branchial
arch, on a reduced scale, after E. Perceval Wright ( Nature , vol. XIV
(1876), p. 313); B, part of a row of gill-rakers, nat. size, after
Hannover and Steenstrup.

“ . . . “fine, stiff, lustrous black strings, resembling bristles or horse-hair, all with one end attached to and hanging over cme side of
a round, cartilaginous arch, of a thumb’s thickness”.

1 D. Vid. Sels. Math. Natur. Skr., 5:te Rseklce, Bd VII, p. 489.

c Bull. Acad. Roy. Belg., ser. 2, tom. XXXI, p. 504, pi. II, fig. 16.

1146

SCANDINAVIAN FISHES.

but large enough to draw blood from the finger, if
incautiously handled — gathered in patches or series,
with interstices between, that give the body an ap-
pearance like that of elephant-hide. Hence the name
conferred upon the species by Lesueur.

The Basking Shark is a pelagic fish of the North
Atlantic®, whose manner of life calls to mind in many
ways that of the whales, which it also rivals in size.
A line between North Africa and Virginia forms the
southern limit of its geographical range, so far as this
has been investigated up to the present time; and the
species roves northwards to Iceland and the extreme
north of Norway, even to Varanger Fjord, principally,
no doubt, within the area of the Gulf Stream. At all
events it does not appear to be, strictly speaking, an
arctic fish; and the old accounts from Greenland of
an immensely large Shark, which was said to devour
dolphins (especially whitefish), rorquals, and humpbacks,
whereon Fabricius based his statement* * * 6 that the Bask-
ing Shark occurred on the coast of Greenland, have
been assigned by Lutken (1. c.) to the category of
fables. The comparatively small eyes and the firm,
thick skin are in themselves sufficient to suggest that
the Basking Shark is no deep-sea fish, properly so call-
ed; and the probability that its life is passed in the
upper levels of the ocean is further increased by the
nature of the food to which it is evidently referred.
As yet, it is true, we know but little of the bathy-
metric distribution in the open sea of those minute
creatures — chiefly lower crustaceans, in general so-
called Entomostraca, and the larvse or even the eggs of
fishes and invertebrates — which compose the diet both
of the large whales and the Basking Shark. Investiga-
tions into the biological conditions at various depths in
the high seas, vigorously pursued as they have been
since first instituted by the SAvedish expedition in 1869
on board the corvette Josephine0, have still much left
to teach us. But Ave already know that the supply of
animal food of this kind in the upper strata of the open
sea is plentiful at depths varying with the changes of
the weather or the set of the currents, and that it
fluctuates at different seasons of the year. Guided
hereby, the Basking Shark too probably shifts its quar-

ters. The course taken by the Gulf Stream affords an
explanation why the Basking Shark is found more fre-
quently off the north of Nonvay than on the south
coast and has never been met Avith in the Skager Rack
or Cattegat. In the North Sea it is rarer than on the
Avest and south coasts of Great Britain and Ireland.
On the Avest coast of France and in Portuguese waters
it has been taken once or tAvice. It penetrates into the
Mediterranean through the Straits of Gibraltar (Doder-
lein). On the east coast of North America it occurs
in the same manner as on this side of the Atlantic.

The Basking Shark is a peaceable and sluggish
creature, harmless to man or other animals of any magni-
tude, and asserting its presence only by roving in quest
of food at the surface, sometimes Avith snout above the
Avater. When it accelerates its pace, cleaving the Avaves
Avith the projecting dorsal and caudal fins, and when
it swims in a company of several, one behind another,
it presents an appearance that may Avell have dictated
an occasional contribution to the history of the great
sea-serpent. Or the same phenomenon may be suggested
to the imagination by a sight of this fish as it lies
during calm Aveather in repose at the surface, often
Avith the belly upAvards, and as the Avaves lap its rotund
body. The name of Basking Shark Avas conferred upon
it by Pennant in exchange for the older name of Sun-
fish employed on the Irish and Welsh coasts, an allusion
to its habit of lying motionless at the surface, as if
basking in the sun. Yet extremes meet, even in the
temperament of the Basking Shark; and it has some-
times been seen to leap several feet out of the water.

In spring the Basking Shark approaches the Irish
coast. Whether this is done for purposes of propaga-
tion, is unknown. Pennant found in a female a foetus
about 3 dm. long; but he does not state the time of
year. On the Avest coast of Ireland, about 100 miles
Avest of CleAv Bay, is a bank long celebrated for its
Basking Shark fishery. Off Tory Island too (N. W.
coast of Ireland) companies of 60 to 100 Basking Sharks
have been seen. In certain years they are commoner;
during others they perhaps do not put in an appearance.
From this locality they seem to migrate northwards
along the Avest coast of Scotland.

a ■ Beown-Goode indeed states (Fisher., Fisher. Industr. U. S., sect. I, p. 669) that the Basking Shark is not unfrequently harpooned

by whalers on the Pacific coast, where a specimen of this species was examined at Monterey (California) by Jordan and Gilbert; but in their

Synopsis of the Fishes of North America the latter writers do not mention a word about this.

6 Fna Oroenl., p. 130.

c See Smitt: De senaste arens under soknin gar om hafsfaunans grcins mot djupet, in the periodical Framtiden for 1870, p. 345.

BASKING SHARK.

1147

“The Basking Shark fishery,” writes Nilsson from
notes taken about seventy years ago, “commences on
the west coast of Norway about the middle of August.
The boats used are about 18- — 20 ells (107 — 119 dm.)
long, carry all the tackle requisite for the fishery, and
are manned by a crew of four. These boats cruise
along the coast, and the Shark generally appears in
their wake and comes close to the boat, but is some-
times found lying still at the surface. One of the
crew then takes the harpoon, which is furnished with
a long shaft (a pole), and plunges it with all his might
into the fish. The others lend a helping hand, ‘but
the moment the bone is touched, he plunges to the
bottom.’ The harpoon-line, which is about 14 — 15
score fathoms (530 — 565 in.) long, and which lies coiled
up ( Jcveilet ) and clear, runs out so rapidly that water
must be poured on the bulwark to prevent it from
catching fire. On reaching the bottom the Basking
Shark swims along until exhausted, towing the boat
out to sea. Sometimes, if lean, it can hold out for
twenty-four hours; if fat, it gives in after three or four
hours. When it is tired out, they haul it up alongside
the boat, and with a long knife cut the tail in front
of the caudal fin, not quite off, for, if so treated, it
would stave in the boat with lashes of the maimed
limb; they leave a part of the fin hanging fast. After-
wards they stab it to death with lances; then turn it
belly upwards in the water, and lash it fast to the mast.

Now one of the fishermen takes a long knife and cuts
out a great piece at the fore end of the liver, where-
upon he inserts his arm and severs all the bands and
ties of the liver. Finally he cuts open the belly with
a single slit, when the liver comes out and floats on
the water like a bolster; but at the same moment the
water rushes into the belly of the fish, and the lashings
must be cut loose with haste, or the boat will sink.”
This fishery was commenced, according to Gunnerus, at
the middle of the eighteenth century, and was still
practised, according to Nilsson, in 1820 — 1830; but it
has now been abandoned as a special occupation, and
for the last forty years the Basking Shark has been
spoken of in Norway almost as a rarity.

The Basking Shark is most sought after for its
liver, which yields an excellent oil; but according to
Gunnerus the poor at least were accustomed in his
time to cut raiding and rafa from the flesh of the
Basking Shark. The outer skin is used for polishing
purposes; and the thick true skin is manufactured into
saddles and shoe-soles.

The Norwegian name of the Basking Shark is pro-
nounced both brugde and brygde. Olafsen cites from
Iceland the names of Ryner and Beinhaakal (Bony
Greenland Shark), the latter having reference to the
greater firmness which its skeleton displays than that
of the Greenland Shark and of the smaller Sharks. In
England too it is known as the Bone Shark.

Fam. SOYLLIIDtE.

Tiro dorsal fins and one anal , the first dorsal situated above the ventral fins or behind the perpendicular there-
from; none of them with spine. Eyes without nictitating membrane. Spiracles open, comparatively large.

Sharks of rather considerable dimensions and some-
times dangerous to man* 6 are indeed not wanting within
the present family, but most of the 25 species are
middle-sized or small. By Gunther the Scyllioids are
distributed among 7 genera. From the preceding Sharks
they differ both in form of body and manner of life.
Instead of pelagic rovers we here meet for the most
part with bottom-fishes and littoral species of nocturnal
habits, hiding themselves by day among seaweeds, on
the sand or under rocks, where they lurk for prey.

The fins are consequently of a more rounded or trun-
cate form. The tail too is straighter, running in a
line with the trunk, and without any marked upward
curvature of the caudal fin, the tip of which forms a
diphycercal lobe, in most cases not very obliquely cut,
the anterior part- of the lower caudal lobe being similar
in shape to the anal fin. Another character of fairly
general validity in this family is the great development
of the nostrils, which is frequently so advanced that
these apertures are confluent, as in the Rays, with the

a See above, p. 415.

6 The Australian Wobbegong ( Crossorhinus ), see Hill in Tenison-Woods, Fish and Fisheries of N. S. Wales , p. 94.

1148

SCANDINAVIAN FISHES.

mouth, being separated from each other and covered
within by a sometimes continuous nasal valvule. The
mouth too of some forms is set very far forward, close
to the tip of the snout, and fringed with dermal flaps.
None of these Sharks gives birth, so far as is known,
to living young.

Within the Scandinavian fauna three species have
been observed, belonging to two very similar genera:

A: Dorsal margin along the beginning of

the caudal fin armed with spines Pristiurus catulus.

B: Dorsal margin without special spiny
armature.

a: Inner posterior corners of the ven-
tral fins acutely prolongated, in the

males coalescent with each other.. Scylliorliinus canicula.
b: Inner posterior corners of the ven-
tral fins obtuse, in the males se-
parated from each other Scylliorliinus stellaris.

Obs. De Filippi and Verani“ have described from the Mediter-
ranean a Scyllium acanthonotum, distinguished by a row of spines on
each side of the back, extending from the middle of the head nearly
to the end of the first dorsal fin. Of the species, however, they had
seen only a young individual with vitelline capsule attached. Gun-
ther* 6 was of the opinion that this young specimen belonged to one
of the above-mentioned Scylliorhini ; Gigliolic referred it to Pristiu-
rus. The correct explanation may be left to the future; but it seems
likely that the spiny armature is a character of youth which dis-
appears entirely in Scylliorliinus , partially in Pristiurus.

Genus PRISTIURUS d.

Nostrils so remote from each other and from the mouth that the distance between their inner angles is perceptibly
greater than that between either of them and the month , somewhat greater than their length , and about equal to
the distance between their outer angles and the orbital margin. Distance betiveen the mouth and the tip of the
snout greater than the postorbital length of the head (to the first branchial aperture ). Anterior part of the upper
caudal margin (fig. 333, a) armed on each side with a row of retral serrated spines.

This genus contains only one species, which is
more nearly approximated by the situation and deli-

Fig. 333. Egg-capsule of the Black-mouthed Dog-fish ( Pristiurus ca-
tulus) from the fishing-bank of Jaderen. Nat. size, a, a part of the
upper edge of the caudal fin in a Black-mouthed Dog-fish 67 cm. long,
superior aspect, X 5 to show the serrated spines and the scale-rows
on the dorsal margin between them.

mitation of the nostrils to the preceding genus than is
the case with the succeeding one. It also has a more
finely shagreened skin than Scylliorliinus . In these Sharks,
as we have seen in numerous Teleosts, e. g. the Gobies
and Salmons, the scales of the tail are larger than those
on the anterior part of the trunk. Here we also find
that the placoid scales of the tail approach more and
more to the form of the jaw-teeth, least so, however,
in Pristiurus. Besides the above-mentioned characters
this genus displays the peculiarity that the tip of the
tail is bent downwards, whereas in the following genus
it is straighter. The egg-capsules of Pristiurus (fig.
333) are distinguished by the absence of horns at the
extremity directed backwards in the uterus, which end
is rounded by the incurvature of the corners and in-
cised at the tip, while the horns at the other end,
one at each corner, are as short as in the egg-shells
of the Rays.

“ Mem. R. Accad. Sc. Torino, ser. 2, tom. XVIII (1859), p. 193, tab., fig. 2.

6 Cat. Brit. Mus., Fish., vol. VIII, p. 403.
c Espos. Intern. Pesca Berlin 1880, Sez. Ital., p. 112.
d Bonap., Fna Ital., Pesci. Established as a subgenus of Scyllium.

SHARKS.

1149

THE BLACK-MOUTHED DOG-FISH (sw. hagalen).

PRISTIURUS CATULUS.

Plate LI, fig. 2.

Coloration above of a reddish or brownish gray, with spots darker than the ground-colour but with a lighter
border, and varying in size and form, usually larger, quadrangular or rounded, next the back, smaller down the sides.
Dorsal fins behind, the other unpaired fins at the base more or less cinereous, at the tops darker. Pectoral and
ventral fins of the same colour as the back or sides, but immaculate, paling outwards. Ventral side white. Mouth
and pharynx, as well as the branchial cavity and peritoneum, black. Iris light blue.

Syn. Haae-Gicele, Strom, Sondm. Beskriv., part. I, p. 283.

Squalus Catulus { Haae-Gcelen ), Gunner., Trondh. Selsk. Skr.,
vol. II, p. 235; Ascan. ( Rodhaae ), Icon. Her. Nat., tab.
XXXVIII; Risso, Ichtli. Nice , p. 30 (p. p. ; feminam Scyllio-
rliini caniculce credidit); Lillj. ( Pristiurus ), Sv . , Norg. Fna ,
Fisk., vol. Ill, p. 657.

Galeus melastornus, Rafin., Caratteri N. Gen., N. Sp., p. 13;
Yarr., Brit. Fish., vol. II, ed. 1, p. 375 ( Scyllium ), ed.
2, p. 495 {Pristiurus)', Bonap. {Pristiurus melanostomus ),
Fna Ital., Pesci, tab. 131, fig. 3; Mull., Hle, Plagiost.,
p. 15, tab. 7; Lowe, Fish. Madeira, p. 93, tab. XIV;
Kr. {Scyllium melastomum ), Danm. Fisk., vol. Ill, p. 832;
Gthr {Pristiurus melanostomus), Gat. Brit. Mus ., Fish., vol.
VIII, p. 406; Coll., Forh. Vid. Sels. Chrnia 1874, Tillasgsh.,
p. 211; N. Mag. Naturv., Bd 29 (1884), p. 117; Malm,
Gbgs , Boh. Fna, p. 623; Winth., Naturli. Tidskr. Kbhvn,
ser. 3, vol. XII, p. 58; Mor., Hist. Nat. Poiss. Fr., tom.
I, p. 284; Day, Fish. Gt. Brit., Irel., vol. II, p. 314,
tab. CLX, fig. 1; Storm, N. Vid. Sels. Skr. 1883, Trondhj.,
p. 43.

Scyllium Artedi, Risso, Fur. Merid ., tom. Ill, p. 117; Barb.
Boc., Cap. {Pristiurus), Peix. Plagiost., p. 11.

Squalus annulatus , Nilss., Prodr. Ichthyol. Scand., p. 114;
{Scyllium), Skand. Fna, Fisk., p. 713.

The Black-mouthed Dog-fish attains according to
Storm a length of 80 cm., according to Moreau of 90
cm.; but it is commonly smaller. Our specimens from
Scandinavia measure between 33 and 68 cm. The largest
are females. The body is of the elongated Shark-form,
with flattened snout, more terete, though somewhat de-
pressed head, behind which the vertical section is more
or less triangular (hog-backed), and more or less com-
pressed tail, almost flattened on the sides, the depth
just in front of the caudal fin being about twice the
thickness. The greatest depth, at the pectoral fins, is
in the males about 9 %, in the females about 10V2
of the length of the body. The greatest breadth falls
within the head, just in front of the first branchial
aperture, and varies, as usual, with the distension of
the jaws and branchial apertures, being somewhat less

Fig. 334. Ventral side of a Black-mouthed Dog-fish {Pristiurus ca-
tulus), 9) from the fishing-bank of Jaderen. Nat. size. The anterior
outer lobe {lae) of the right nostril is raised to show the anterior
inner {lai) and the posterior outer {Ipe) lobes.

or greater than the greatest depth. The least depth
of the tail, just, in front of the caudal fin, is about
4 — 4x/2 % of the length of the body.

1150

SCANDINAVIAN FISHES.

The head occupies about, 17 (17 1/2) — 15 (14'8) %
of the length of the body, and differs essentially from
that of the following genus in the length of the snout,
which measures nearly half (49 1/2 — 46 %) of that of
the head. The snout is translucent, and its skin is
pierced with numerous ampullar orilices, arranged above
in series so as to form oblong patches on each side of
the median line, below in the manner illustrated in
fig. 334. The nostrils, which are directed obliquely
inwards and backwards from the edge of the snout,
have the anterior overlapping margin double (cf. above,
on the Tope), and the outer lobe (lae) has a pointed
triangular flap, the base of which is continued inwards,
the inner ( lai ) an obtuse, more or less rounded flap,
belonging to the outer part of the nostril. Exactly
opposite the last-mentioned flap there projects a more
deep-seated, more tubercular lobe (Ipe) from the pos-
terior margin of the nostril. In this manner the nostril
is incompletely divided, as that of the Rays, into an
outer (anterior) and an inner (a posterior) part; but
the latter is not confluent either with the nostril of
the opposite side or with the mouth. The distance
between the inner ends of the two nostrils is somewhat
more than 1/3 (in the younger among our specimens
about 36 %, in the older about 34 1/a %) of the length
of the snout; and the distance from either of these
ends to the mouth varies between % and 3/i of the
former distance. The orbits are oblong, the vertical
diameter being about 2/5 of the longitudinal or even
less, and the longitudinal diameter measuring about
half the length of the snout, somewhat more or less,
or about 2/s of the interorbital width. Exactly behind
the orbits — at a distance from them rather greater
than their length — lie the spiracles, whose diameter
is about 1/4 of that of the former. The mouth is semi-
circular, the lower jaw being somewhat more pointed
than the upper, and is so situated that the anterior
margin of the upper jaw lies in a line with the pre-
orbital margin or a little behind the perpendicular there-
from, the tip of the lower jaw almost beloAv the centre
of the eyes. Lips are wanting, but the corners of the
mouth are elevated and posteriorly bounded by a rather
deep semicircular groove. The teeth are pointed, with
one or two lateral denticulations on each side of the
main cusp. They are set in a quincunx of three — five
series, which gives them the appearance of being dis-
tributed in some thirty longitudinal rows running
obliquely backwards and outwards across the jaws. The

tongue is fleshy and flat, as in most of the Sharks.
The gill-openings are middle-sized, the hindmost being
set above the beginning of the pectoral fin.

The dorsal fins are characterized, as in the follow-
ing genus, by their similarity to each other both in
form and size; but in these fins, as in the other un-
paired fins, it should be observed that the forward
extension of the anterior margin varies in different in-
dividuals, the measurement of the basal length being
thus rendered uncertain. The distance from the tip of
the snout to the first dorsal fin is about 45 (44 l/2 —
47 7$) %, to the second about 63 (62 — 64) %, of the
length of the body; the interval between the two fins
measures between 13 and 14 % of the said length.
Their form is obliquely quadrilateral, with the posterior
corner almost rectangular. The length of their base
varies between 4 and 4V2 % (sometimes nearly 5 %),
and their height between 5 and 6 1/2 %, of the length
of the body. The first dorsal fin lies opposite to the
anterior part of the space between the ventral and anal
fins, the second to the posterior part of the anal fin.
The last- mentioned fin, which has an unmistakable re-
semblance to the anterior part of the lower caudal lobe
(an obliquely trapezoidal form, gradually attenuated
behind, with pointed infero-posterior corner and slightly
concave under margin), is here distinguished by its
length — about 15 — 16 % of that of the body, and in
the females 4 times or less, in the males up to 5 times
the greatest height of the fin. The distance between
the tip of the snout and the beginning of the anal fin
measures in full grown females perceptibly more than
v2 (about 54 %), in males and young females about
7 2 or less (49 — 51 %), of the length of the body. The
caudal fin is expanded at the end, obliquely truncate,
with somewhat rounded corners. Its length from the
beginning of the anterior lobe is more than 1/i (27 1/2
to nearly 33 %) of the length of the body.

The pectoral fins are broad, obliquely truncate,
with rounded corners. Their length at the anterior
margin varies between 10 and 1 2 1 /2 %, their base be-
tween 61/2 and 8 %, of the length of the body. The
ventral fins are quadrangular, low but long, with the
posterior corner pointed. Their base is longer than that
of the pectorals — in the females about 872 %, in the males
about 9 %, of the length of the body — but their height
(breadth) is only about x/3 or 3/g of the length of the base.

The lateral line is quite distinct, from the very
temples, and at first runs about half as far from the

ROUSSETTES.

1151

dorsal margin as from the ventral, but on the tail at
about the middle of the sides, until it reaches the an-
terior part of the caudal fin, where it bends down-
wards in a loop to the lower third of the depth of
the body.

The Black-mouthed Dog-fish is most known in the
Mediterranean and on the coasts of Portugal and Nor-
way, this being probably due to the circumstances that
it generally lives at a depth of about 100 — 250 fthrns.,
and that fisheries at this depth are most developed in
the said localities. But it also roves into higher strata,
being met with, though seldom, in the North Sea, and
penetrating in the Skager Rack and Cattegat into 40

fathoms of water (Malm) off Bohusl&n and in Christia-
nia Fjord, or even, as once happened, in March, 1847,
according to Nilsson, into the Sound off Raa. Lowe
has described it from Madeira; according to Collett
it has been found off’ Tromso. It thus has a wide
geographical range in the North Atlantic.

The food of the Black-mouthed Dog-fish consists
of fish and crustaceans. It takes the ordinary bait of
deep-sea lines. During spring and summer the female
deposits her singular eggs (fig. 333), two at a time,
as described by Gunnerus. The life of this species is
otherwise little known ; but where it is taken in any
number, the flesh is considered eatable.

Genus SCYLLIORHINUS".

Nostrils approximated behind to the month or even meeting the anterior margin of the upper jaw , the distance
between which margin and the tip of the snout is less than the post orbital length of the head. Upper edge of

the caudal fin without prominent serrations.

As has been hinted above, the difference from the
preceding genus is rather inconsiderable. The most
important distinction consists in the advancement of the
mouth and the more or less marked prolongation of the
nostrils back towards the mouth cavity. Consequently
we also find in the genus Scylliorhinus that the tip of
the lower jaw is in a line with or even in front of the
preorbital margin. The genus is indeed without serra-
tions at the upper edge of the caudal fin , though these
may sometimes be traced in the difference between the
large spiny scales in the upper rows on each side and
the smaller and smoother ones in the median dorsal
line at the said spot; but it has a compensation for
them in the stronger development of the spiny scales
on each side of the body above and behind the anal
fin, which almost exactly resemble jaw-teeth. These
scales literally form a rasp, which the fish uses as a
defensive weapon.

To the characteristics of the genus also belongs
the form of the egg-capsules (fig. 335). These are
similar to those of the Rays, oblong, rectangular, and
flattened, but with one half thicker than the other.
At each corner they are furnished with a long (mea-
suring sometimes half a metre) filamentous appendage.
When the first pair of these tendrils emerge from the

cloaca, the female coils them round a branch of sea-
weed or coral, where the egg is left hanging. Two
eggs, as a rule, become simultaneously ripe for exclu-
sion, one in each uterus; and the female deposits them

Fig. 335. Egg of Nurse Hound ( Scylliorhinus stellaris ), cut open to
show the enclosed foetus, and with the posterior (the first excluded)
end turned downwards. Nat. size. After Moreau, who further states
that the ova of this species differ from those of the Rough Hound
in that the thickened side-margins are transversely striped.

a 2'/dhov in Aristotle, elsewhere G'/xXaS, ( canicula in Gaza), = whelp.

145

Scandinavian Fishes.

1152

SCANDINAVIAN FISHES.

in pairs, at certain intervals. In the course of a month
a female Nurse Hound has been seen to lay 18 eggs.
The eggs deposited early in April were hatched at the
beginning of December (Coste, Coinptes Rendus, Ja-
nuary 21, 1867).

The systematic name of the genus, Scylliorhinus,
was coined by Blainville in 1816 and must be re-
cognised, as Jordan and Gilbert3 have proposed, in
deference to the accepted laws of nomenclature, although
the Cuvierian name of Scyllium has hitherto been in

general use. The Swedish name of Rodliajar (Red
Sharks) is originally a Danish rendering, introduced
by Kroner, of the French Roussettes, an allusion to
the red coloration prevalent in the family, which cha-
racteristic is further distinguished by its maculation
from the plainer dress of other Sharks. Nilsson call-
ed these Sharks doggfiskar, an adaptation of the Eng-
lish word.

Of the genus 8 or 9 species are known from tro-
pical and temperate seas.

THE NURSE HOUND.

SCYLLIORHINUS STELLARIS.
Fig. 336.

Nasal valvules separated exactly in front of the mouth by a space equal in width to the height of the hindmost
gill-opening. Length of the head about 19 — 17 % of that of the body and greater than the distance betiveen the
anterior margins of the two dorsal fins. Distance betiveen the pectoral fins and the tip of the snout at least 2/5
of that between the first dorsal and the same point. Distance between the ventral fins and the tip of the snout
somewhat greater than that between the first dorsal and the eyes; their inner posterior corners obtuse, scarcely
elongated behind. Length of the base of the anal fin greater than or at least equal to the distance between the
two dorsal fins. Length of the caudal fin from the beginning of the inferior lobe more than 1/i of the length
of the body. Greatest depth of the body about 10 — 12 %b ', least depth , in front of the caudal fin, about — 5 %,
of the length of the body. Coloration above grayish brown, sometimes shading into yellow or red, below white.
Back, sides, and fins, the paired ones even underneath, strewn with comparatively large roundish spots — some
( more constant) blue-black, others {sometimes indistinct or wanting) whitish — about equal in size to the gill-

openings, smaller on the head and the forepart of the back.

Syn. Galeus stellaris minor , Belon., Nat., Divers. Poiss., p. 65.

Canicula saxatilis, Rondel., De Pise., “p. 383. Catulus
maximus, Willughb.. Hist. Pise., p. 63; Ray, Synops.
Method. Pise., p. 22. Squalus cinereus, pinnis ventralibus
discretis, Art., Ichthyol., Gen. Pise., p. 69; Syn. Pise.,
p. 97.

Squalus stellaris, Lin., Syst. Nat., ed. X, tom. I, p. 235;
Flmng (Scyllium), Brit. Anim ,, p. 165; Bonap., Iconogr.
Fna Ital., Pesci, tab. 131, fig. 2; Thomps., Nat. Hist. Irel.,
vol. IV, p. 247; Gthr, Cat. Brit. Mus., Fish., vol. VIII,
p. 402; Malm., Ofvers. Vet. Akad. Fork. 1875, No. 10,
p. 33; Gbgs , Boh. Fna, p. 622; Lillj., So., Norg. Fna,
Fisk., vol. Ill, p. 652; Car., Prodr. Fnce Medit., vol. II,
p. 508.

Squalus canicula, Brunn. (nee Lin.), Ichthyol. Massil., p. 4;

Bl. (p. p.), Naturg. Ausl. Fisch., part. I, p. 16, tab. CXII.
Squalus' ( Scyllium ) catulus ( + Sq. stellaris ), Cov., Regn. Anim.,
ed. 1, tom. II, p. 124; Yarr. ( Scyllium catulus, nec Lin.),

Brit. Fish., ed. 1, vol. II, p. 373; ed. 2, vol. II, p.
493; Mull., Hle, Plagiost., p. 9, tab. 7; Dumer., Hist.
Nat. Poiss. (su. a Buff.), tom. I, p. 316; Barb. Boc.,
Cap., Peix. Plagiost. Portug., p. 11; Mor., Hist. Nat.
Poiss. Fr., tom. I, p. 280; Day, Fish. Gt. Brit., Irel.,
vol. II, p. 312.

Nurse Hound, Couch, Fish. Brit. Isl., vol. I, p. 11, tab. I.

The Nurse Hound attains a length of at least
12 (A dm. (Bocage and Capello; according to Day
15 dm.); but so large specimens appear to be rare.
As a rule the length varies between about 5 and 7
dm. It is strictly a Mediterranean fish, dispersed in
the Atlantic to Ireland and England. Sometimes it
strays north to the Orkney and Shetland Islands; and

a Bull. U. S. Nat. Mus., No. 16, p. 869.

6 Sometimes, according to DoDerlein (Man. Ittiol. Medit., fasc. II, p. 23), about 14 °/.

NURSE HOUNI).

1153

on a single occasion it has been met with in Scandi-
navian waters. In November, 1875, Malm received a
female 622 mm. long that had been taken on a Had-
dock-line north-west of Hallo, in the central portion
of the island-belt of Bohusl&n. The favourite haunts
of the Nurse Hound are the deepest algal zones or

tebrates, but also of fishes. As mentioned above, it takes
the bait of long-lines set for Haddock.

From April to September ripe egg-capsules have
been found in the uterine dilatations of the female.
The development of the foetus in the egg after deposi-
tion is said to take nine months.

Fig. 336. Nurse Hound ( Scylliorhinus stellaris ), c 1 2 na^- size- From Nice, Giguoli.

lower levels, on a stony or rocky bottom; its coloration
too indicates that it does not usually frequent the re-
gion of the green algae, even if the females repair
thither, as Thompson states, to attach their egg-capsules
to Laminaria. Its food is, no doubt, the ordinary diet
of the small Sharks, consisting principally of inver-

The skin of the Nurse Hound is considered to be
one of the best polishers — it bites marble and iron —
and after being smoothed it is employed in cover-
ing the hilts of sabres and small-swords. Its flesh
is eaten freely by the poor of Southern Europe. In
Scandinavia its only value is that of rarity.

1154

SCANDINAVIAN FISHES.

THE ROUGH HOUND.

SCYLLIORHINUS CANICULA.

Plate LI, fig. 4.

Nasal valvules in front of the mouth almost confluent , forming a broad dermal fold interrupted only for a very
short distance at the middle. Length of the head about 12 1/2 — l31/2 % of that of the body and. less than the
distance between the anterior margins of the two dorsal fins. Distance between the pectoral fins and the tip of
the snout about 1/3 (31 — 37 ?6) of that between the first dorsal and the same point,. Distance between the ventral
fins and the tip of the snout about i/5 (77 — 83 %) of that between the first dorsal and the same point and less
than that between the latter fin and the eyes. Inner posterior corner of the ventral fins acutely elongated , the
two fins coalescent behind in the males. Length of the base of the anal fin about 3/4 — ij5 (71— 83 %) of the
distance between the two dorsal fins. Length of the caudal fin from the beginning of the inferior lobe less than
7 4 (21 — 24 %) of the length of the body. Greatest depth of the body about 8x/2 — 11 %, least depth , in front of
the caudal fim, about 3 — 33/4 %, of the length of the body. Coloration above russet , on the sides grayish yellow,
paling downwards towards the white ventral side. Fins more or less reddish. Body more or less densely strewn
above and on the sides, as well as the fins (the paired ones only on the upper surface), with roundish brown
spots, densest but smallest on the head and bach, more scattered but larger (though not so large as the gill-open-

ings) on the sides of

Syn. Galeus stellaris major , Belon., Nat., Div. Poiss., p. 64. Cani-
cula Aristotelis, Rondel., De Pise., p. 380. Catulus major
vulgaris ($ adult.) + Catulus minor (juv. et o71), Willughb.,
Hist. Pise., pp. 62 et 64. Catulus major vulgaris 4- C.

minor vulgaris, Ray, Syn. Meth. Pise., p. 22. Squalus ex

rufo varius, Pinna ani medio inter anum et caudam pinnatam
+ Squ. dorso vario; Pinnis ventralibus concretis, Art., Ich-
thyol., Gen. Pise., pp. 68 et 69; Syn. Pise., p. 97. Spotted
Dog Fish + Lesser Spotted Dog Fish, Penn., Brit. Zool., vol.
Ill (ed. 1776), pp. 99 et 101, tab. XIX.

Squalus Canicula, Lin., Syst. Nat., ed. X, tom. I, p. 234;

Gmel., ed. XIII, p. 1490; Retz., Fna Suec. Lin., p. 305;

Blainv. ( Scylliorhinus ), Bull. Sc. Soc. Philom. 1816, p. 121;
Cuv. ( Scyllium , subg.), R'egn. Anim., ed. I, tom. II, p. 124;
Nilss., Prodr. Ichthyol. Seand., p. 113; Bonap. ( Scyllium ),
Iconogr. Fna Ital., tab. 131, fig. 1; Mull., Hle, Plagiost.,
p. 6, tab. 7; Ku., Damn. Fish., vol. Ill, p. 814; Nilss.,
Skand. Fna, Fisk., p. 710; Thomps., Nat. Hist. Irel ., vol.
IV, p. 247 ; Dum., Hist. Nat. Poiss. (su. a Buff.), tom. I,
p. 315; Barb. Boc., Cap., Peix. Plagiost. Port., p. 11;
Fedders., Tidsskr. Fisk. Kbhvn, Aarg. 2 (1868), p. 123;
Aarg. 4 (1870), p. 339; Gthr, Cat. Brit. Mus., Fish., vol.
VIII, p. 402; V. Ben., Poiss. Belg. (Mem. Acad. Sc. Belg.,
tom. XXXVIII), p. 3; Coll., Forh. Vid. Selsk. Chrnia 1874,
Tillaegsh., p. 210; 1879, No. 1, p. 104; N. Mag. Naturv.
Chrnia, Bd. 29, p. 116 ; Malm, £)f vers. Vet. Akad. Forh. 1875,
No. 10, p. 33; Gbgs, Boh. Fna, p. 619; Winth., Naturh.
Tidskr. Kbhvn, ser. 3, vol. XII, p. 57; Mor., Hist. Nat.
Poiss. Fr., tom. I, p. 278; Doderl., Man. Ittiol. Medit.,
fasc. II, p. 24; Day, Fish. Gt. Brit., Irel., vol. II, p. 309,
tab. CLIX, fig. 1 ; Petersen, Vid. Meddel. Naturh. For.
Kbhvn 1884, p. 160; Lill.t., Sv., Norg. Fna, Fisk., vol.
Ill, p. 645; Car., Prodr. Fnce Medit., vol. II, p. 508.
Squalus Catulus, Lin., 1. c., p. 235; Brunn., Ichth. Massil.,
p. 5; Bl., Naturg. Ausl. Fisch., pt. I, p. 21, tab. CXIV ;

he body and the fins.

Donov., Brit. Fish., tab. LV; Blainv. ( Scylliorhinus ), Fne
Franc., Poiss., p. 69, tab. 17, fig. 1 (+var. Sc. Canicula,
p. 71).

Rough Hound, Couch, Hist. Fish. Brit. Isl., vol. I, p. 14,
tab. II.

Obs. Willughb y and Ray regarded the male and female as
distinct species. Artedi and Linnaeus were hardly more than copyists
of their predecessors; but Gmelin, Retzius, and Blainville paved the
way for a correct apprehension of the case. At present there need
be no risk of ambiguity in dropping the Linnasan name of Catulus
here and applying it instead, as Gunnerus has done; to the Black-
mouthed Dog-fish.

The Rough Hound never quite attains the maxi-
mum size of the preceding species. The largest speci-
mens on record were about 1 in. long (10 1/g dm. ac-
cording to Day). The ordinary length of both species
is, however, the same, 5 — 7 dm. In the terete form of
the body, anteriorly somewhat depressed, and broadest
at the posterior part of the head, they are also much
alike; but the Rough Hound approaches nearer to the
Black-mouthed Dog-tish in the usually greater elongation
of the body and the anterior compression of the back.

The difference in the form and delimitation of the
nostrils (fig. 337) shows that the Rough Hound comes
still nearer to those members of the family which are
yet more singularly equipped in this respect. Not only
is the outer lobe (lae) at the anterior margin of each
nostril expanded forwards and prolongated inwards, so

ROUGH HOUND.

1155

close to the corresponding lobe of the other side that
an almost continuous nasal valvule is produced, not
unlike that of the Rays — the posterior margin of the
nostril is also furnished internally in both species with
a dermal lobe, which in the Nurse Hound, however, is
simple and thin, projecting like a leaf into the back-
ward narial groove, but here develops into two barbel-
like processes, one of them (above the dotted line from
Ipi) jutting into the said groove, the other (below the
dotted line from Ipi) directed backwards and laid within
the mouth, outside the margin of the upper jaw. The
lower jaw is furnished in both species, but in the Nurse
Hound only behind, in the Rough Hound throughout
the greater part of its length, with a pendent labial
fold; and the skin on the margin of the upper jaw
swells towards the corners of the mouth in a labial form.

As a compensation for the absent nictitating mem-
brane these Sharks possess the faculty of raising the
lower eyelid so high that the orbits become shut; and
Nilsson as well as Malm has described hotv the Rough
Hound can close the pupil by contracting the iris, till
the latter leaves open only a narrow longitudinal slit,
dilated at each end.

The fins of the Rough Hound are most like those
of the Black-mouthed Dog-fish; but the dorsals are set
somewhat further back, the first beginning about half-
way along the body, the second just- behind the second
third thereof, and the anal and caudal fins are shorter,
the base of the former measuring only about 9 — 10 %
of the length of the body.

The shagreened skin is fairly alike in both the
Roussettes, but generally rougher in the Nurse Hound.

The Rough Hound has about the same geogra-
phical range as the Nurse Hound, but is somewhat-
more addicted to northern rovings. On the English
coast it- is far commoner than the latter, and it is con-
sidered to be one of the commonest Sharks in British
waters. In Scandinavia it- is indeed scarce, but not
among the rarest fishes, for it has repeatedly been met
with on the west coast of Norway, south of Bergen,
and in the Cattegat — even in the Sound, between
Malmo and Helsingborg, one specimen has been taken
(Lilljeborg). Off Mount Kullen, according to Nilsson,
it is familiar enough to bear a special name ( Haskdrf -
vincj ); and at- Agger, on the west- coast- of Jutland, it-
is known, according to Feddersen, as the Tusliaj
(Toad Shark). The original of our figure (Plate LI,
fig. 4) is also from the west coast- of Jutland, and was

presented to the Royal Museum in November, 1889,
by Mr. Fredericksen, a Copenhagen merchant-.

The Rough Hound prefers a sandy bottom with
its growth of seaweed, as indicated by the reddish co-
loration the fish generally displays. When it- lives on
a clayey bottom, the body has a different ground-colour,
darker (grayish or blackish) with still darker spots
(Thompson). In its reproduction, temperament, and
general habits, this species resembles the preceding one.
Both have something of the serpent about them, as

lac
lai \ \

Fig. 337. Ventral side of a Rough Hound ( Scylliorhinus canicula),
cA from Messina, 6 — 10 fthms., stony bottom, Feb. 5, 1891, C.
Bovallius. V2 nat. size. The anterior outer lobe ( lae ) of the right
nostril is raised to show the anterior inner (lai), the posterior outer
(Ipe), and the posterior inner (Ipi) lobes.

they stvay their whole body in sinuous movements;
and both are gregarious, each species by itself, as they
chase schooling fishes. To the Herring and Pilchard
they are dangerous enemies, and they cause the fisher-
man trouble by tangling and tearing his nets. The
long-line fishermen too complain of them. In the Eng-

1156

SCANDINAVIAN FISHES.

lish Channel it has happened" that fishermen who had
shot long-lines for Cod have found a Rough Hound on
almost every hook; and the hooked Cod had been de-
voured down to the head and a bit of the backbone.
The Rough Hound eats, besides fish, both crustaceans
and mollusks, seeming to be especially fond of whelks
(Buccinum undatum ) and lugworms ( Arenicola ).

As human food the Rough Hound also finds con-
sumers among the fishing population and the poor.
In Scandinavia it is far too rare to possess any eco-
nomical importance. Not so in France and the rest

of Southern Europe. Its skin is used in polishing
various substances. Its hard flesh and musky odour
are disagreeable, but the latter is said to be removed
by boiling. The liver, which is described as having
an abominable taste, is considered poisonous. One Doc-
tor Sauvage of Montpellier related in 1745 that a fa-
mily who had eaten the liver of this fish were over-
powered by heavy drowsiness, from which they first
recovered on the third day, and which was attended
with a skin disease, causing the whole epidermis to
peel off in fragments.

CYCLOSPONDYLI.

Sharks with two dorsal fins, hut no anal.

Hasse has shown* that the Sharks whose external
character is expressed in the absence of an anal finc,
in their internal characters stand lowest, next the Cow-
Sharks ( Notidanidce ) — which indeed have an anal fin,
but only one dorsal — and nearest to the primordial
forms common to the Sharks and Rays. A comparison
between our two figures 296 and 297 (see above, pp.
1068 — 69) shows the great difference in the structure
of the spinal column between a Cyclospondylous and
an Asterospondylous Shark. In the former the ver-
tebra? are far less differentiated. The bodies of the
vertebrae form a continuous canal, only imperfectly
coarctated by the constrictions answering to the middle
parts of fully developed vertebra?. Frequently, though
irregularly, so-called diplospondylism occurs, two pairs
of apophyses (two neurals and two inter neurals), or at
least two pairs of foramina for spinal nerves on each
side, appearing in a single vertebra. The basal parts
of the dorsal fins also exhibit a lower grade of diffe-

rentiation; but in compensation most of the Cyclospon-
dyli are equipped at the anterior margin of each dorsal
fin with a spine (ichthyodorulite), which in the species
our figure (296) represents, however, is considerably
reduced, not projecting above the skin.

All the Cyclospondyli have open and rather large
spiracles. Their eyes are without nictitating membrane.
All their gill-openings lie in front of the pectoral fins.
Their caudal fin is generally less heterocercal than that
of other Sharks. Most of them are characterized by
the singular form of the jaw-teeth. One genus ( Acan -
thorhinus ) differs from all the other known Sharks in
having the duodenum furnished at its commencement
with a pair of cgecal appendages'*. The great majority of
them are known to be viviparous; but one species of the ge-
nus just mentioned, the Greenland Shark, has been sus-
pected in recent times to form an exception to this rule*.

Hasse has divided the Cyclospondyli into three
families: Lcemargfi, without externally visible dorsal

“ Brehm, Thierleben , 2:te Aufl., Abth. 3, Bd 2, p. 377.

b Naturl. Syst. Elasmobr., Allgem. Th. p. 41; Besond. Th., p. 55.

c Squales anhypopteriens Moreau, Hist. Nat. Poiss. Fr., tom. I, pp. 276 and 340.

d This observation may be traced even in Gunnerus (Trondhj. Selsk. Skr., vol. II, pi. X, fig. 2), but is unnoticed in the text.
e See Lutken, Vid. Meddel. Naturh. For. Kbhvn 1879 — 80, p. 56. A female Greenland Shark 23 dm. long was taken in the North
Sea during January, 1891, and presented by Mr. Fredericksen of Copenhagen to the Royal Museum. The oviducts lay, as Turner (Journ.
Anat., Physiol., vol. XII [1877 — 78], p. 604) also found them, extended from the diaphragm to the cloaca, straight and thin-walled, gra-
dually expanding in the hindmost part alone to the thickness of a goose-quill, and here with somewhat stouter walls than in front, but with-
out any special dilatation whatever throughout their extent. From this circumstance it is, however, impossible to decide with certainty the
appearance presented by the oviducts after the act of copulation and the development within them of the impregnated eggs.
f Containing the genera Acanthorhinus , Isistius , Euprotomicrus , and Scymnus.

SPUR-SIIARKS.

1157

fin-spines; Echinorhinia with the first dorsal fin oppo-
site to the ventrals and with large dermal spines, si-
milar to the aculei of the Rays, but without dorsal fin-
spines; and Spinacidceb, with externally prominent spine
before each dorsal fin. Two of these families are re-
presented in the Scandinavian fauna.

A : At the anterior margin of each
dorsal fin a more or less prominent
spine. — Fam. Spinacidce.

a: Distance between each nostril
and the tip of the snout con-
siderably more than half of that
between the former and the

mouth Squalus acanthias.

b: Distance between each nostril
and the tip of the snout less
than half of that between the
former and the mouth Etmopterus spinax.

B: No externally prominent dorsal fin-

spines. — Fam. Scymnidce Acanthorlrinus carcharias.

Fam. SPIN AC 11) M.

First dorsal fin set half-way between the perpendiculars from the pectorals and
At the anterior margin of each dorsal fin a more or less prominent spine.

\

Skin uniformly shagreened.
ventrals or further forward.

The intermediate position between the Cow-Sharks
(. Notidanidce ) and the Roussettes (Scylliidee) occupied in
many respects by the Cyclospondyli , is especially mani-
fested in the form of the jaw-teeth. The primitive
Notidanidan type is characterized partly by the diffe-
rence of the teeth in both jaws — longer (broader) and
coarser, like serrate disks, in the mandible — partly by
the obliquity in the growth of the denticulations, espe-
cially in the lateral teeth of the lower jaw, with the
tips of all the denticles directed outwards (towards the
corners of the mouth) and with the innermost (or one
of the innermost) denticles largest, so that the others
evidently answer to accessory cusps. In most of the
Cyclospondyli , and especially within the present family,

this difference between the jaws, as well as the obli-
quity of the teeth, has persisted — only one genus, the
Greenlandic and North American Centroscyllium, has
teeth similar in both jaws and resembling those of the
Roussettes — or the obliquity has advanced yet further,
the original inner edge of the largest denticle being
turned entirely aside and having become an incisive
upper edge; and this form of dentition may finally
develop not only in the lower jaw but also in the
upper. The differences in the dentition besides af-
ford the best distinctive characters between the 5
genera into which the family of the Spur-Sharks
has been divided, together including about 20 known
species.

Genus SQUALUS.

Teeth of the upper and loiver jaivs almost similar in form , obliquely shaped incisors. Nostrils middle-sized , their
width much less than the distance from the tip of the snout to the middle of a transverse line crossing their an-
terior margins. Pectoral fins triangular ( following the ordinary Shark type), with the outer posterior margin

concave. Dorsal fin-spines without lateral grooves.

In the works of most recent writers the Linnsean he employed this generic name, Linnaeus had first in
generic name of Squalus has been dropped entirely, or view the species which is commonest in Scandinavian

applied, as in Bonaparte, to the Blue Sharks, as re- waters, and which he ranged first in his systematic

presenting the highest development of the Shark type. enumeration of the Sharks.

Jordan and Gilbert, however, have advanced the opi- Among the Sharks destitute of anal fin the genus

nionc, previously maintained by Rafinesque* that, when Squalus is the most differentiated or, so to say, mo-

a Containing a solitary species, Ecliinorhinus spinosus, from the Atlantic between the North Sea and the Cape of Good Hope and
from the Mediterranean.

6 Containing the genera Centrophorus , Centrina , Centroscyllium , Squalus , and Etmopterus.
c Bull. U. S. Nat. Mus., No. 16, p. 16.
d Caratt. N. Gen., Spec., p. 13.

1158

SCANDINAVIAN FISHES.

dernized, most nearly resembling in external form the
preceding Sharks. This is best shown by the character
drawn from the shape of the pectoral tins, and might
also be expressed by the relative dimensions of these
fins, but for the fact that the alterations of growth
within the genus approximate the fry, or at least the
almost viable foetuses, too nearly to the following ge-
nera. The same remark essentially applies to the form
and dimensions of the two dorsal fins. How closely
the genus Squalus is allied, however, to the remaining
genera of the family, appears from the rank long as-

signed to a Mediterranean form, the so-called Squalus
uyato of Rafinesque (1. c.), as an independent species
of this genus, until Bellotti and Doderlein" disco-
vered it to be a young form of Centrophorus granu-
losus, an inhabitant of the neighbouring parts of the
Atlantic6. Even between the two species left within
the genus Squalus after this elimination there would
appear to obtain some developmental relation, one of
them ( Squalus Blainvillei ) representing in its cha-
racters the juvenile and male characters of the
other.

THE PICKED DOG-FISH (sw. pigghajen).

SQUALUS ACANTHIAS.

Plate LII, figs. 1 and 2.

Distance between each pectoral fin and the tip of the snout at most about V3 (67 — 58 %) of that between the
first dorsal and the same point. The ventral fins begin half-way along the body or somewhat further back.

Syn. 'M.VMvd'lag yaleog, Aristot., Zool., lib. VI, cap. X (= Mu-
stellus spinax, Gaza). Galeus acanthias, Rondel., De Pise.,
p. 373. Cams Acanthias , Gaz® spinax, Schonev., Ichthyol.
Slesv. Hols., p. 29. Squalus pinna Ani nulla, ambitu cor-
poris subrotundo, Art., Ichthyol., Gen. Pise., p. 66; Syn.
Pise., p. 94; Spec. Pise., p. 102; Lin., Fna Suec., ed.
I, p. 100; It. Wgoth., p. 174; Mus. Adolph. Frid., part.

I, p. 53 ( Hundfi.sk ). Haae, Str6m, Sondm. Beskr., part. I,
p. 280. Haafur, Olafs., Reise Isl. , part. I, p. 359; part.

II, p. 989.

Squalus Acanthias, Lin., Syst. Nat., ed. X, tom. I, p. 233;
Bl., Naturg. Fisch. Deutschl., part. Ill, p. 74, tabb. 75
(fig. 1) et 85; Retz., Fna Suec. Lin., p. 305; Blnvlle
( Acanthorhinus ), Bull. Sc. Soc. Philoin. 1816, p. 121; Cuv.
( Spinax , subg.), Regn. Anim., ed. 1, tom. II, p. 130;
Nilss. (Squalus), Prodr. Ichthyol. Scand., p. 117; Bonap.
(Spinax), Iconogr. Fna Ital., tab. 139; Sundev., v. We.
(Squalus), Skand. Fislc., ed 1, p. 187, tab. 46; Jord.,

Gilb., Bull. U. S. Nat Mus., No. 16, p. 16; Lillj., Sv.,

Norg. Fna, Fisk., vol. Ill, p. 665.

Picked Dog Fish, Penn., Brit. Zool. (ed. 1776), vol. Ill, p.
88; Yarr., Brit. Fish., ed. 1, vol. II, p. 400; Couch, Fish.
Brit. Isl., vol. I, p. 49, tab. XI.

Squalus fernandinus, Molina, Chili, p. 229; Guichen. (Spinax)
in Gay, Chile , Zool., tom. II, p. 365.

Acanthias vulgaris, Risso, Eur. Merid., tom. 3, p. 131; Mull.,
Hle, Plagiost., p. 83; Schleg. in Sieb., Fna Japon ., Poiss.,
p. 304, tab. CXXXV ; Kr., Damn. Fisk., vol. Ill, p. 868;
Nilss., Skand. Fna, Fisk., p. 731; Dum., Hist. Nat. Poiss.

(uouv. su. a Buff.), tom. I, p. 437 ; Barb. Boo., Cap., Peix.

Plagiost. Port., p. 21; Gthr, Cat. Brit. Mus., Fish., vol.
VIII, p. 418; Coll., Forh. Vid. Selsk. Chruia 1874, Tilliegsh.,
p. 211; 1879, No. 1, p. 104; Winth., Naturli. Tidskr.
Kbhvn, ser. 3, vol. XII, p. 58; Mor., Hist. Nat. Poiss.
Fr., tom. I, p. 342; Doderl., Man. Ittiol. Medit., fasc. II,
p. 86; Mob., Hcke, Fisch. Osts., p. 152; Day, Fish. Gt.
Brit., Irel., vol. II, p. 315, tab. CLX, fig. 2; Storm, N.
Vid. Sels. Skr. Trondhj. 1884, p. 45; Car., Prodr. Fnce
Medit., vol. II, p 503.

Acanthias americanus, Stor., Mem. Amer. Acad. Arts, Sc., vol.
II, p. 506; vol. IX, p. 2.32, tab. XXXVIII, fig. 1.

Acanthias Linnei, Malm, Gbgs, Boh. Fna, p. 624.

The Picked Dog-fish belongs to the smaller Sharks.
According to Day it attains a length of 12 dm. on the
English coast; but in Scandinavia it does not exceed,
so far as we know, one of about 9 dm.c New-born
fry with obliterated navel measure about 22 — 25 cm.
In the external form of the body the Picked Dog-fish
is not unlike the Tope, but it is usually rather more
elongated. The greatest depth of the body, just in front
of the first dorsal fin, varies irregularly between about
9 and \\ % (according to Kroner sometimes 8 or 12 %)
of its length, the least depth between 2 and 2 1/2 %
(according to Kroyer sometimes nearly 3 %) of the
same. The latter depth measures between about 13 %

a Man. Ittiol. Medit., fasc. II, p. 118, Nota.

6 This might besides be taken for granted partly from Rafinesque’s description of the teeth of the upper jaw (denti minuti ed acuti),
partly from ESarboza de Bocage’s and Capello’s figures of old and young specimeus of Centrophorus granulosus.
c The Vega Expedition brought home a male 9V2 dm. long from Behring Island.

PICKED DOG-FISH.

1159

(according to Kroyer 16 — 17 %) of the length of the
head to 'the first gill-opening. Like the Tope, the
Picked Dog-fish has an almost terete body, but the
anterior part of the trunk is of the so-called hog-backed
form, with a rounded triangular section. Behind the
body is slightly compressed, the breadth being about
equal to the depth, and the section almost quadran-
gular, though rounded (convex) above and laterally
sharpened below by a dermal carina, running on each
side behind the perpendicular from the second dorsal
fin and some way out above the beginning of the in-
ferior caudal lobe. The sides of the tail are, hoAvever,
without true median carina, nor are there any caudal
grooves, though a trace thereof may sometimes be
found above, at the dorsal margin, in front of the
caudal fin. The head, in particular the translucent
snout, is depressed, behind somewhat broader than the
trunk, with breadth about equal to the depth of the
latter, and with parabolically pointed snout.

The length of the head to the first branchial aper-
ture varies between 17 (in the nearly viable foetus 18)
and 15 % of the length of the body. The length of the
snout to the eyes measures about 2/5 (40 — 44 %) of the
length of the head. Both the eyes and the orbits are
oblong, the latter most so, the vertical diameter of the
eyes being always more than 1/2 (53 — 70 %), that of
the orbits sometimes 1/3 (41 — 33 %a), of the longitudi-
nal diameter. This measures in the case of the latter,
which are besides sharply pointed at each end, about
30' — 20 % of the length of the head. The pupil is
either round, but acute-angled below, or transversely
oblong. Behind the eyes, but higher (further in) than
these, lie the rather large spiracles, their diameter being
only slightly less than or even equal to the vertical
diameter of the orbits, from which they are separated
by a distance of the same magnitude. Their anterior
margin is folded double, the fold being sometimes so
deep as to give them the appearance of being really
double. The interorbital width is generally about equal
to the length of the snout, somewhat greater or less,
usually the latter. The prone nostrils are set about
half-way along the snout, but the distance from the
mouth to the transverse line drawn through their pos-
terior margins is greater in the young than that from
the tip of the snout to the transverse line drawn through

“ According to Kroyer down to 28 %.

6 In a nearly viable foetus rather more than V3.

their anterior margins. In the old the said distances
are about equal, the tip of the snout in front of the
nostrils being even relatively prolongated with age.
The nostrils are, as usual, obliquely transversal, with
a small pointed valvule at the middle of the anterior
margin. Their width is about 1/3 of the distance be-
tween them. The mouth is less curved than in most
other Sharks, with deep folds around the corners. The
distance from the tip of the snout to the anterior mar-
gin of the mouth is about x/2, to the corners thereof
about 60 — 70 of the length of the head. The teeth

are densely set, without gaps or unpaired median tooth.
They are merely attached to the skin, as in all the
Plagiostoms, not to the cartilage, and are consequently
mobile. The first two rows of teeth stand upright,
apart from the rest, the inner row leaning backwards;
the remaining 3 or 4 rows have the cusps turned inwards
(backwards), covered by the transverse fold (velum) in the

Fig. 338. One of the right teeth of the upper jaw ( a ) and the lower
jaw (b) in the Picked Dog-fish ( Squalus acanthias), anterior aspect. X 5.

front part of each jaw. The last-mentioned rows are,
however, erected in the old, where the anterior rows
have disappeared. Each tooth (fig. 338) is small — in
an adult fish only about 2 mm. high — flat and sharp-
edged, with a deep jag in the outer (posterior) margin,
under the keen, laterally directed cusp. They have the
same form in both jaws, but are somewhat larger in
the lower than in the upper. In front they have a
basal cusp running into the gum, and reminding us of
the median ridge and the shaft with which the der-
mal scales are furnished. In a young female Sundevall
counted 26 teeth in each transverse row above and
22 below. The tongue is not free, and consists mere-
ly of a skin investing the rather large hyoid carti-
lage, which occupies almost the whole mandibular arch.
The branchial apertures are comparatively small, the
middle ones smallest, the hindmost opening somewhat
larger than the first, the height of which is about Vs
to 7 2 greater than the vertical diameter of the orbits.

Scandinavian Fishes.

146

1160

SCANDINAVIAN FISHES.

The two dorsal tins in their relative position to
each other resemble those of the Tope; but their form
approaches more nearly to the square, and each of
them is furnished at the anterior margin with a fairly
stout and strong spine. As a rule the second dorsal
tin is perceptibly smaller than the first, about 3/i as
large; but its spine is the larger, being even higher
than the fin itself, whereas the spine of the first dorsal
extends only about half-way up the anterior margin
thereof. The spines are pointed, somewhat curved, and
triangular in section, with the whole posterior surface
concave, but with rather convex side-margins. The
first dorsal fin commences at about the end of the first
third of the length of the body, the second dorsal at
the termination of the second third thereof, somewhat
further forward in the adult males than in the females,
and further back in the young of both sexes than in
the old. The length of the base of the first dorsal is
about equal to the interorbital width. The base of the
second dorsal measures about 4/5 — ®/4 of that of the
first. The caudal fin is distinguished partly by the

a b

Fig. 339. A scale of the Picked Dog-fisli ( Squahis acantluas), seen
from above (from without, a, with the retral tip directed upwards in
the figure) and from the side ( b , with the retral tip turned to the
right of the figure). About 37 times the natural size.

slight upward curvature of the tip of the tail within
the fin, partly by the greater relative breadth of the
upper lobe than in the preceding Sharks and the ab-
sence of incision in the hind part of the inferior mar-
gin of the fin. The length of the upper anterior mar-
gin of the fin is in the males somewhat more, in the
females somewhat less, than x/5 of the length of the
body. The lower anterior margin is about half as long
as the upper or a little longer.

The pectoral fins indeed approach to the more
spatulate form they exhibit in the following Sharks,
but they remind us of the preceding species in the
marked prolongation of the posterior inner corner, the
hind margin being thus rendered concave. The distance
between them and the tip of the snout is about x/5 of
the length of the body, and in the males about 2/3 , in
the females about 4/5, of that between the first dorsal
fin and the same point. The length of their anterior

margin measures about 80 — 95 % of that of the head
— relatively less in the young — - and the breadth of
their base about 34 — 40 % of their length. The ven-
tral fins are obliquely lozenge-shaped, or resemble an
obtuse-angled triangle, when the fins are laid back, in
which case the base is continued, without a break, by
the inner posterior margin. The distance between them
and the tip of the snout in the males is about 50 —
52 % , in the females about 53 — 54 %, of the length of
the body. Their base together with their inner pos-
terior margin measures about 9 or 10 %, their anterior
margin about 5 or 6 %, of the length of the body. In
adult males the distally flattened pterygopodia project
far beyond the ventral fins, and are furnished on the
upper surface with a broad dermal groove, in which
tAvo spines are concealed, the one long and hamatelv
curved at the tip, the other, which is visible in our
figure (PI. LII, fig. 1), pointed, grooved, and serrated.

The skin is uniformly shagreened with scales of a
singular form (fig. 339), elevated on a terete shaft, at
the top of which they are rectangularly bent and ex-
panded behind in a three-pointed foliate form, with the
middle point longest and most powerfully strengthened
by a median ridge, passing into the anterior margin
of the shaft, on Avhose sides the lateral ridges are also
continued.

The coloration is above grayish broAvn, more or
less dark, shading doAvn the sides into violet, with a
feAv small, roundish, Avhitish spots set in a roAv along
the anterior half of the lateral line and in another row
along the back, Avhich latter series is much sparser, but
is continued in very young specimens back to the tail.
The belly is pale and whitish, but finely punctated with
dark dots. After death the colour of the back soon
fades and becomes more grayish, but it may be better
preserved by depositing the specimen at once in strong
spirit.

The external difference of sex may be traced in
unborn foetuses 1 8 cm. long. The males among these
have fairly distinct pterygopodia, Avhich are, hoAvever,
only half as long as the inner margins of the ventral
fins. These young specimens are almost similar in form
and coloration to the adult individuals, only that the
tip of the snout is someAvhat shorter, the distance be-
tween the nostrils .and the extreme tip of the snout
being only about 2/3 of that between them and the
mouth, and that the dorsal fin-spines are short. The
posterior of these spines measures only 2/3 of the height

PICKED DOG-FISH.

1161

of the fin, and the naked tip of each spine projects
only a very little way above the skin. The breadth
of the caudal and pectoral fins is also rather greater
in proportion to their length. Somewhat smaller foe-
tuses, 15 — 16 cm. long, with the large vitelline sac still
attached in the form of a pear 7 cm. long, with a stalk
measuring 272 cm., have a deep groove, like a seam,
along the under surface of the snout. Their fin-spines
are so short that the tips have not emerged from their
scaly dermal envelope, which occupies, however, as
great a portion of the fin-margin as in adult specimens.
In the males only quite indistinct rudiments of the
pterygopodia, not yet protruding from the skin, are
visible. The coloration is fully developed. The young
are not born until the yolk has been entirely absorbed,
and may then have attained a length, as mentioned
above, of 25 cm.

The Picked Dog-fish has a very extensive geogra-
phical range. In the North Atlantic this includes the
most northerly parts of Europe, Iceland, and the east
coast of the United States, as well as the west coast
of Europe and the Mediterranean. To the south the
species has been found off the Cape of Good Hope, the
Mascarene Isles (Dumeril), and on the coast of Australia
(Richardson and Gunther). In the Pacific it is also
known from Chili (Molina), Japan (Sen leg el), and
Behring Island (the Vega Expedition). It is conse-
quently to be regarded, we may almost say, as a cos-
mopolite. Throughout the west coast of Scandinavia
it is common, and it penetrates, though more rarely,
into the Baltic to the east coast of Riigen (Mobius and
Heincke) and to the north-east of Scania (Wallengren®)-

In Scania, Bohuslan, and everywhere in Norway
the Picked Dog-fish is called Ha ( Ilaa ). In Sweden
the name is written liaj , probably from some German
or Dutch dialect. Other, less usual names are: Pig glia
or Hdfisk , in Iceland Hafr or Haafur , on the Faroe Is-
lands Haavur, in Scania Hdkatt or Hafskatt. The name
of lia (Shark) seems to have been everywhere applied
among the Gothic nations by preference to this species,
the commonest Shark; and the other forms have been
distinguished by the addition of some suffix, as Ha-
brand (the Porbeagle), Hdstorje (the Tope), Hamcir,
Hakdring, or Hakal (the Greenland Shark).

The Picked Dog-tish has its constant habitat on a
soft and oozy bottom, but, like most of the Sharks, is

a confirmed rover. It swims in shoals, and is exceed-
ingly voracious. Its food consists both of lower marine
animals and of fishes, especially those of gregarious habits,
such as Clupeoids, Garpike, and even Cod or Haddocks,
which it bites in two with ease, though they be not
much thinner than itself. When Picked Dog-fish appear
in shoals, they are a great nuisance to the fisherman,
for they plunder and damage nets set for other fishes,
make off with hooked fish or get caught themselves
instead, a poor exchange, and often bite off the snood
above the hook, which is thus lost. They are said to be
most numerous early in spring, when they approach the
coast in enormous shoals, whose multitude, it is alleged,
rivals that of the Herring. A boat’s crew, it is stated,
can then take several boat-loads in a day. From Norway
Sundevall was told that the Herring-shoals are some-
times hemmed in near shore by Picked Dog-fish, which
keep watch outside. They are as plentiful on the Eng-
lish coast as in Norway, but in Bohuslan they do not
seem to occur in quite so great numbers, when the
Herring is not there. Strom, in his description of Sond-
more, says that they withdraw from the shallows at the
beginning of April, shortly after the Cod have come in,
after which very many are hooked, about Whitsuntide,
in the deep fjords. On the coast of Bohuslan they are
again taken towards autumn, first on hand-lines and
Haddock-lines, thus in somewhat deeper water, during
August and September, when the catch is said to consist
almost exclusively of males, and later on, in October,
when they are said often to penetrate far up the fjords
and to remain there until the frost sets in. During
winter they are not met with on the said coast, except
when the Herring is there, possibly because they are not
sought after in the deep water where they pass this season.

Aristotle stated that on the coast of Greece this
fish copulates in August and brings forth its young
from May till August. The case is apparently as a
rule the same in Scandinavian waters. Ekstrom con-
cluded from his observations in Bohuslan that the
breeding is performed in shoals during August or Sep-
tember in rather deep water. According to many cor-
roborative statements the young are born most plenti-
fully at the end of April and the beginning of May
and afterwards, in less number, throughout the summer.
W. v. Wright states that during spring the females
assemble in large companies near land, in small pools

Ofvers. Vet. Akad. Forh. 1866, No. 1, p. 5.

1162

SCANDINAVIAN FISHES.

3 — 6 fthms. deep, which are known in Bohuslan as
hdljor, and are then taken in the seine, without the
catch of a single male. When drawn into the boat at
this time of year, they often give birth to young ones,
the probable origin of the assertion that young fish
15 — 17 dm. long swim in shoals together with their
elders on predatory excursions. The Swedish fishermen
declare that the fry never accompany the old, but live
by themselves in shallow water. There they keep to
the bottom, and are often taken on hand-lines during
autumn. As they are never caught in numbers at the
same time, it would appear that they do not gather in
shoals, but lead a more solitary life. Each female
seems to bring forth only about 10 young — in Ireland
Thompson was told that the number is always odd —
and the fishermen state that the female seldom contains
more than 5 or 6 “ full-grown ” young at a time. Sun-
devall found all the young taken out of the same
female to be exactly similar in size and development,
whence he concluded that they are probably born
shortly after one another. As we have mentioned above,
the new-born fry are 22 — 25 cm. long. Those taken
in autumn are hardly any larger, and even in January
a few specimens of about the same size are now and

then caught. These are probably born late in the year,
and Thompson states from Ireland" that even at Christ-
mas-time he found a female Picked Dog-fish with 9
nearly mature foetuses (5 male and 4 female) in the
oviducts.

The flesh of the Picked Dog-fish is quite white
and free from the disagreeable smell with which the
larger Sharks are tainted. It is said to be fairly pa-
latable, and it is eaten in Bohuslan both fresh, either
boiled or fried, and cured, either salted or dried, in
the last case usually after lyeing* 6. So too in Norway,
where it is, however, not much esteemed. In Scotland
and in Southern Europe the Picked Dog-fish is also
frequently used for food. In a dried state it is a mar-
ketable commodity. If left uncooked for some days in
summer, it begins to give off a rather powerful and
exceedingly pure ammoniacal vapour, without any per-
ceptible putrid smell, such as that usually emitted by
decaying flesh. The caudal fin, which is a common
article of trade in the island- belt of Bohuslan, under
the name of ha-rump, is used by cabinet-makers in
polishing wood and metal. In many localities the eggs
are made into pancakes. The liver is boiled down
into oil, and gives a fair yield. (Sundevall, Smitt.)

Genus ETMOPTERUS.

Teeth of the upper jaw symmetrical in form, multicuspid; those of the lower jaw obliquely shaped incisors.
Nostrils large, their width being greater than the distance from the tip of the snout to the middle of a transverse
line crossing their anterior margins. Pectoral fins ovally spatulate. Dorsal fin-spines with lateral grooves.

It is true that Rafinesque", when he established
this genus in 1810, based it partly on an erroneous
observation — that it possessed only three branchial
apertures — partly on a fortuitous character — that its
dorsal fins Avere split at the margin d. But no ambi-
guity is thereby involved, and the current rules of no-
menclature therefore demand that his name for the genus
should supersede the generally adopted, but younger
name of Spinax, which Cuvier" and, after him, Bona-

parte, in accordance with an unfortunate practice, bor-
roAved from the Linmean specific name.

Gunther indeed recognises three species of this
genus, one from Europe, another from Madeira and
the West Indies, and a third7 from the southern
coast of South America; but the distinction be-
tween them is apparently rather dubious, and the
former two are identified Avith each other by Du-
meril.

wing, fin), which ought strictly to be written Etlimopterus. Still we

a Nat. Hist. Irel., vol. IV, p. 254.

6 See above, p. 1107, note f.
c Caratt. N. Gen., Spec., p. 14.

d Hence the generic name (Gr. rfigog, strainer and tcveqov.
retain here the orthography of its author.

e Regn. Anim., ed. I, tom. II, p. 129 (1817).
f Spinax granulosus, Shore Fishes, Rep. Challeng. Exped., Zoo!., 6, p. 19, tab. II, fig. (J.

SHARKS.

1163

THE SAGRE (sw. blakaxan).
ETMOPTERUS SPINAX.

Plate LI, fig. 3.

Skin shagreened with curved , setiform spines , expanding into three or four arms at the base. Coloration above

blackish blue , underneath coal-black.

Fig. 340. Teeth and scales of the Sagre ( Etinopterus spinax). a , two teeth from the upper jaw; b, the four middle, longitudinal rows
of mandibular teeth, removed from the jaw and seen from in front, the descending compensatory teeth being visible through the mucous
membrane; c, the same rows of teeth, seen from behind (from within); a — c, from a female 42 cm. long, X 4; d and e, scales, anterior

and lateral aspects, X about 45.

Syn. Galeus Acanthias seu spinax fttscus , Willugbb., Hist. Pise.,
p. 57. Squalus pinna ani carens; naribus in extremo rostro,
Art., Ichthyol., Gen. Pise., p. 67 ; Syn. Pise., p. 95.
Squalus Spinax, Lin., Syst. Nat., ed. X, tom. I, p. 233 5
Martin, Vet. Akad. Hand]. 1761, p. 227; Lin., Fna Suec.,
ed. 2, p. 107 ; Gunn. ( Sortliaaen ), Trondhj. Sels. Skr., vol.
II (1763), p. 313, tab. VIII; Ascan. ( Blaataske ), Icon.
Rer. Nat., tab. XXXVII; Retz., Fna Suec. Lin., p. 305;
Risso, Ichth. Nice, p. 41; Id. ( Acanthias ), Eur. Me'r ., tom.
Ill, p. 132; Nilss., Prodr. Ichthyol. Scancl., p. 118.
Etmopterus aculeatus, Rafin., Car. N. Gen., Spec., p. 14,
tab. XIII, iig. 3.

Spinax niger, Cloquet, Diet. Sc. Natur., T. I (Suppl.), p. 93;
Agass., Poiss. Foss., tom. Ill, pp. 61, 93, tabb. A, fig. 3;
B, figg. 4, 5; Bonap., Iconogr. Fna Ital., Pesci, tab. 141;
Mull., Hle, Plagiost., p. 86; Kr., Damn. Fisk., vol. Ill,
p. 893; Nilss., Skand. Fna, Fisk., p. 729; Dum., Hist.
Nat. Poiss. (N. su. a Buff.), tom. I, p. 441, tab. 4, figg.
13 et 14; Gthr, Cat. Brit. Mus., Fish., vol. VIII, p. 424;
Coll., Forh. Vid. Selsk. Chrnia 1874, Tillsegsh., p. 212;

1879, No. 1, p. 104; Gigl., Espos. intern, pesca Berlino

1880, Sez. Ital., Cat., p. 113; Mor., Hist. Nat. Poiss. Fr.,
tom. I, p. 348; Doderl., Man. Ittiol. Medit., fasc. II, p.
96; Storm, N. Vid. Selsk. Skr. Trondhj. 1883, p. 44; Car.,
Prodr. Face Medit., vol. II, p. 504.

Spinax Gunneri, Rhdt, Overs. D. Vid. Selsk. Forh. 1824 — 27,
p. XVI; Coll., N. Mag. Naturv. Chrnia, Bd 29 (1884), p.
117; Lillj., Sv., Norg. Fna, Fisk., vol. Ill, p. 673.

? Acanthiclium pusillum , Lowe, Fish. Mad., p. 37, tab. VI.

Spinax Linnei , Malm, Gbgs, Boh. Fna, p. 626.

The Sagre is the smallest European Shark. It
would scarcely appear to attain a length of half a
metre: Gunnerus’s largest specimen was “somewhat
more than half an ell in length” (between 30 and 40
cm.), and the largest female obtained by Storm from
Trondhjem Fjord was 47 cm. long. The body is of a
terete, fusiform shape, with rather indistinct hog-back
flatly convex on the forepart of the back and on the
belly, somewhat flattened at the middle of the sides,
deepest at the first dorsal fin, and gradually passing
forwards from this point into the flatly depressed snout,
though with a depression at the occiput, between the
spiracles, and a corresponding swelling below, at the
lower jaw and the hyoid bone. The peduncle of the

a In old females, with belly distended by ova or foetuses, the hog-backed form (rounded triangular section) of the body is more
pronounced.

1164

SCANDINAVIAN FISHES.

tail is of a rounded quadrangular section, and is more
or less deeply impressed in the median line above and
below. Where the body is shallowest, just in front of
the caudal fin, the breadth (thickness) is about 3/4 of the
depth. The greatest depth measures about 1372 (?) —
1 1 1/2 (cf) %, and the least depth about 272— 3 %, of
the length of the body. The greatest breadth occurs
in the head, just in front of the gill-openings, and is

Fig. 341. Anterior part of the ventral side in a Sagre ( Etmopterus
spinax ), ok 4 dm. long, from Stromstad Fjord. Nat. size.

about 11 — 9 % of the length of the body. Across the
gill-openings, on the other hand, the body is somewhat
contracted, forming, as it were, a neck.

The length of the head to the first branchial aper-
ture measures in full-grown specimens about 1 7 J/2 —
17 %oa of the length of the body. Above the head is
almost flat, underneath convex. Between the eyes the

forehead is coursed by a longitudinal groove on each
side of a terete median carina, which forks behind into
two divergent branches. The interorbital width (in-
cluding the dermal rim of the supraorbital margins) is
about 45 %, and the length of the elongated, anteriorly
and posteriorly pointed orbits about 31 %, of the length
of the head. The eyes themselves are also elongated,
the height of the iris being about 2/s of its length,
which is nearly 7s of that of the head. The length
of the snout is about equal to that of the orbits, and
in form the snout differs considerably from that of the
preceding species, advancing with uniform breadth,
until it is abruptly rounded off at the nostrils to a
very blunt tip. The nostrils (fig. 341) are apparently
double on each side, the small, pointed nasal valvules,
one from the anterior margin, one from the posterior,
laying their tips together, so that each nostril seems to
consist of a circular aperture, situated at the very margin
of the snout, and a proximal, oblong opening, directed
obliquely backwards and inwards. Their entire breadth
(both apertures) is about equal to the distance between
their inner extremities, but only 73 °f that between
the latter and the middle of the margin of the upper
jaw, which margin lies below the posterior margin of
the eyes, about half-way along the head. At the middle
of the under surface of the snout is a longitudinal con-
cavity which calls to mind the groove present at the
same point in foetuses of the preceding species. Among
the pores belonging to the system of the lateral line
on the snout, a double row on each side of the median
groove underneath and a similar series on the upper
surface, in a loop above the nostrils, are especially pro-
minent. At an angle to the last-mentioned series a
row crooks over each eye and is continued back towards
the occiput. The broad mouth is slightly curved, its
breadth being about 3/7 of the length of the head, or
somewhat greater than the distance between the nostrils
and the upper jaw. The thin corners of the mouth
are each surrounded by a deep groove, which is conti-
nued backwards by a slit narrowing to a point. The
upper jaw is armed with 3 or 4 transverse rows of
tricuspid or quinquecuspid teeth, with the median cusp
largest, one row close behind the other. The lower
jaw contains only one row of simultaneously function-
ing teeth, essentially similar to those of the preceding
species, and forming, as there, a serrated cutting-edge;

According to Lilljeborg 16 according to Krgyer sometimes 15Y2 %.

SAGRE.

1165

but behind (within) this row lie two or three rows of
compensatory teeth, directed inwards and downwards.
Thick dermal folds form, as usual, transverse curtains
and cover the inner rows of teeth both in the upper
jaw and the lower. The tongue is broad and flat. On
opening the mouth, we can see from within straight
through the large spiracles, which here, as in the pre-
ceding species, have the anterior margin double. Here
too they lie obliquely behind and above (within) the
posterior ends of the orbits, so far inwards that the
distance between them is only about 4/5 of the inter-
orbital width, and so far back that their distance from
the tip of the snout measures somewhat more than 2/3
of the length of the head. Their width is about half
the external diameter of the iris. The gill-openings
are still smaller than in the Picked Dog-fish. They
are scarcely larger than the spiracles, or even smaller
than these. The last three are often set so close to-
gether that at a casual glance one may fail to distin-
guish between them, this being the probable explana-
tion of Rafinesque’s above-mentioned mistake.

All the fins are extremely thin at the outer mar-
gin, without scaly covering, and more or less trans-
parent. They are consequently torn in most cases at
the said margin, and they are seldom seen so entire as
in our figure, which is drawn from a newly caught
specimen in a good state of preservation.

The dorsal fins are distinguished by the posterior
being considerably larger than the anterior and having
about twice as large a spine. The first dorsal fin com-
mences at the end of the first third of the body, the
second somewhat before the end of the second third
thereof. The form of the first dorsal calls to mind
tha/t of the adipose fin in the Salmonoids, the second
is more typically a Shark fin, with concave outer pos-
terior margin. The caudal fin, as a whole, is sickle-
shaped, with the usual sinuses in the under margin
only subindicated. The laterally compressed tip of the
tail forms a slight upward curve within it. From the
beginning of the upper lobe the caudal fin measures
rather more than l/5, from that of the lower lobe rather
less than 1/i, of the entire length of the body.

The insertion of the obtusely rounded pectoral fins
commences at a distance from the tip of the snout
measuring somewhat more than Vs °f the length of the
body. Their length is about 8 — 8x/2 0/0 of that of the

“ See for example above, p. 155, on the Blue-mouth.

body or 43 — 54 % of that of the head; and the breadth
of their base is about 52 — 56 % of their length. The
ventral fins are elongated, with the outer angle strongly
rounded. Their insertion, which measures about 872
— 93/4 % of the length of the body, begins at a distance
from the tip of the snout answering to about 54—56 %
of the said length. The pterygopodia of the fully ma-
ture males — figured by Dumeril, 1. c., Pi. 4, fig. 13 —
have before the tip three curved and pointed spines, two
of which — the outermost being mobile — are set at
the outer margin of the cleft on the upper side of these
organs, the third being hidden in the skin near the top
of the inner tegumentary margin of the cleft. Tile
cloacal aperture occupies about the posterior half of
the space between the ventral fins.

The scaly armature we have already noticed. The
lateral line is sometimes quite distinct, marked by a
black punctuation, and following a rather straight course,
near the back, until it comes within the region of the
caudal fin, where it crooks downwards to follow the
inferior margin of the tip of the tail and pass straight
out over the fin itself.

The coloration is distinguished from that of the
preceding Sharks first and foremost by its being darkest
on the ventral side; and from this black colour a stripe
ascends on each side behind the ventral fins and ex-
pands both forwards, above the said fins, and backwards
on the sides of the tail. Another similar stripe runs
up before the beginning of the inferior caudal lobe and
spreads along the lower part of the tip of the tail. The
termination of the lateral line is also marked by a fine,
coal-black streak along the under margin of the tip of
the tail. The blackish blue colour of the back and
sides acquires along the middle of the latter a more
or less distinct silvery band, due to the grayish lustre
of the setiform scales. The iris has a greenish metallic
lustre; the pupil is green. The dorsal fin-spines and
the teeth, as well as the outer parts of the eyes, are
conspicuously marked by their white colour.

The Sagre is strictly a deep-sea fish, and it con-
sequently shares with several other fishes" the pecu-
liarity of having been found in widely separated loca-
lities without being discovered in the interjacent regions.
Thus it was first described by Willughby from the
Mediterranean, where it was already known — more
than three hundred years ago — to the Genoese fisher-

1166

SCANDINAVIAN FISHES.

men under the above name. It is certainly not very
rare in the Mediterranean, for in 1879 Giglioli took
40 specimens on one occasion from depths of 800 —
1,000 m. The stated occurrence of the Sagre on the
coast of Portugal is doubted by Barboza and Capello;
but in the Bay of Biscay it is found, though seldom,
according to Lafont (in Moreau). On the coasts of the
United Kingdom and in the North Sea it has not yet
been met with ; but on the Norwegian coast it is com-
mon, occurring northwards at least to the 70th degree
of latitude. In Trondhjem Fjord, according to Storm,
is is one of the most usual catches on deep-sea lines.
But it also ascends into comparatively shallow water,
and the original of our figure, a male, was taken in
November, 1881, on a Cod-line, at a depth of 40 — 50
fthms. in Stromstad Fjord. In Christiania Fjord, ac-
cording to Collett, it is common. As mentioned above,
it can hardly be distinct in species from Acantliidium
pusillum , a Madeiran fish described by Lowe, which also
lives, according to Gunther, in Cuban waters; and re-
membering the wide distribution of the preceding spe-
cies, we do not consider it impossible that with the
Sagre should be also identified in species Gunther’s
Spinax granulosus from Chili.

The Sagre swims in shoals and, despite its insigni-
ficant size, pesters the fisherman after the same fashion,

if not in the same degree, as the Picked Dog-fish. It
often bites on the long-line, and is a disappointing sub-
stitute for the expected catch. Its diet is similar to
that of the Picked Dog-fish, though it must, of course,
confine itself more exclusively to small creatures. In
a female 43 cm. long, from Bohusl&n, the contents of
the stomach comprised fish-scales (some Scopeloid spe-
cies), a small cuttle-fish (? Octopus ), almost digested,
feet of crustaceans (some shrimp or Pandalus), several
specimens of an indeterminable Annelid, fragments of
worm-tubes, and a quantity of clayey sediment. The
same female contained in the left ovary 8 eggs, in the
right 5, about 14—17 mm. in diameter; but no eggs
had passed into the oviducts. During April Storm
found in Trondhjem Fjord a female with 14 young in
the oviducts, and at the middle of July Collett ob-
tained off Namdalen females with 7 or 8 fully deve-
loped foetuses. In the Mediterranean, according to
Risso, the Sagre breeds all the year round, giving
birth to 15 — 20 young about 1 dm. long.

The flesh of the Sagre is not used for food; but
the liver yields a good oil.

Frequently the Sagre is infested with an ectoparasite,
a Cirrhiped, Anelasma ( Alepas ) squaticola , Loven, which
forms at the dorsal fins and on the under surface of the
head tumours, first described and figured by Gunnerus.

Fam. SCYMNIDiE.

Skin uniformly shagreened with tubercular or conical spinulce. First dorsal fin situated half-way betiveen the
perpendiculars from the pectorals and ventrals or further forward. No externally protruding dorsal fin-spines.
Teeth of the upper and lower jaivs different from each other , but all smooth-margined and without lateral cusps.

The two genera that compose this family, Scymnus
and Acanthorhinus, come nearest both in the dentition
and the coloration to Etmopterus. Hasse has suggested"
that the obscure pelagic forms Isistius brasiliensis and
Euprotomicrus Labordii should also be ranged within
this family, in which case the above character for the
position of the first dorsal fin must be altered, this
fin being removed rather far back in the two last-

o

mentioned genera, more as in the family Echinorhi-
nidce. In either case we are apparently confronted
here with one or more series of reduction-forms, which
have originated from the Picked Dog-fish type, and
vdiose genealogical relations to one another and to the
primordial forms still require elucidation. The Scan-
dinavian fauna possesses only one genus.

Nat. Syst. Elasmobr., Allgem. Th., p. 43.

SHARKS.

1167

Genus ACANTHORHINUS.

Teeth of the upper jaw conical , carinated on the anterior surface, and set in several simultaneously functioned
transverse rows. Teeth of the lower jaw obliquely shaped incisors, set in severed transverse rows, of which the

foremost two or three are simultaneously erect and set
the eyes as from the mouth. Length of the caudal fin

Of this genus two species have been described, one
of which, however, Acanthorliinus rostratus, from the
Mediterranean and the Atlantic outside, is extremely
little known, and has been founded on characters that
seem rather uncertain. The name of the genus was
coined by Blainville, who in 1816 conferred it upon
all the Sharks without anal fin with especial reference,
it is true, to the Picked Dog-fish, but also enumerating
among the species our Greenland Shark. Subsequently
the genus has received several names, which must be
rejected, however, for different reasons. In 1817 Cuvier
included the Greerdand Shark in the subgenus Scymnus

close together. Length of the snout about the same from
at the upper edge less than t/5 of the length of the body.

( les Lwiches ), from which it was afterwards removed
on account of the dentition. Lesueur (1818) gave the
genus the name of Somniosus, a translation of the
Sleeper, as the Greenland Shark is called by American
fishermen. In 1841 Muller and Henle bestowed
upon it the name of Lcemargus , which Ivroyer had
conferred shortly before (1837) on a genus of crus-
taceans. The oldest available name is Blainville’s,
and as it cannot be applied to either of the two pre-
ceding genera, both of which have earlier names, its
employment here is fully justifiable.

THE GREENLAND SHARK (sw. hakaringen or haskardingen).

ACANTHORHINUS CA RCHARIAS.

Plate LII, fig. 3.

Skin rather sparsely shagreened with thorn-like ( conical ) spines, in an unworn state pointed , with grooved base and
arcuately recurved tip. Length of the snout about 4/5 of the postorbital length of the head or more. Greatest
depth of the body about 15 — 17 %, its least depth ( just in front of the caudal fin) about 3 V2 — 4 %, of its length.
Coloration brown or black, shading on the sides into violet; iris blue, pupil green.

Fig. 342. Upper ( a ) and lower (b) jaw-teeth of a Greenland Shark (Acanthorliinus carcharias ) 3'/2 In- long, nat. size; c, d, and e, three
dermal spines (scales) of different form from the same individual as a and b, magnified.

Syn. Hay, Egede, Gronl. Perlustr., p. 49. Cards marinus , Cam's § 14. Haae-Kierring, StrOm, Sondm. Beskriv., p. 284.

Carcharias, Crantz, Gronl. Hist. (ed. suec.), lib. II, cap. 2, Haakall, Olafs., Reis. Isl., pp. 359, 597, 839.

a Rafinesqoe had indeed collected all these Sharks into one genus ( Dalatias ) previously (Car. Ale. N. Gen., Spec., p. 10); but he
characterized them by the absence of spiracles, and besides he was not acquainted with any species of the present genus.

Scandinavian Fishes.

147

1168

SCANDINAVIAN FISHES.

Squalus Carcharias ( Ha a- Slier ding), Gunn., Trondbj. Sels. Skr.,
vol. II (1763), p. 330, tabb. X et XI; Lin. (p. p.), Syst.
Nat., ed. XII, tom. I, p. 400; Fabr., Fna Groenl., p. 127;
MOll., Zool. Dan. Prodr., p. 38; Rink, Gronl., vol. I, i, p.
135; vol. II, p. 212.

Squalus microcephalus, Bl., Schn., Syst. Ichth., p. 135; Kr.
( Scymnus ), Damn. Fisk., vol. Ill, p. 914; Mgrn, Ofvers.
Vet. Akad. Fork. 1864, p. 536; 1867, p. 264; Coll.,
Forb. Vid. Selsk. Chrnia 1874, Tillaegsh., p. 212; Malm,
Gbgs, Boh. Fna, p. 626; Winth. ( Lcemargus ), Naturb.
Tidskr. Kbbvn, ser. 3, vol. XII, p. 59; Ltkn ( Somniosus ),
Vid. Meddel. Naturb. For. Kbbvn 1879 — 80, p. 56; Mela
(Scymnus), Vert. Fenn ., p. 366, tab. X; Jord., Gilb. (Som-
niosus), Bull. U. S. Nat. Mus., No. 16, p. 15; Storm ( Scym-
nus), N. Vid. Selsk. Skr. Trondbj. 1883, p. 44; Coll.
(Somniosus), N. Mag. Naturv. Chrnia, Bd 29 (1884), p. 117;
Day (Lamar g us), Fish. Gt. Brit., Irel., vol. II, p. 320,
tab. CLXII, fig. 1 ; Petersen (Somniosus), Vid. Meddel. Naturh.
For. Kbbvn 1884 — 86, p. 160; Lillj., Sv., Norg. Fna,
Fisk., vol. Ill, p. 679.

Acanthorhinus Nonvegianus, Blnvlle, Bull. Sc. Soc. Philom.

1816, p. 121 (4 -A. Microcephalus , ibid.).

Somniosus brevipinna, Lesueur, Journ. Acad. Nat. Soc. Pliil.,
vol. I (1818), p. 222; Stor. (Scymnus), Mem. Amer. Acad.
Arts. Sc., N. ser., vol. IX (Boston 1867), p. 235, tab.
XXXVIII, fig. 2; Mor. (Lcemargus), Hist. Nat. Poiss. Fr .,
tom. I, p. 361.

Squalus borealis, Scoresby, Acc. Arct. Peg., vol. I, p. 538,
tab. XV, figg. 3 — 5; Flmng (Scymnus), Brit. Anim., p.
166; Yarr. (Greenland Shark), Brit. Fish., ed. 1, vol. II,
p. 403; Mull., Hle (Scymnus, Lcemargus), Plagiost., p.
93; Nilss. (Scymnus), Skancl. Fna, Fisk., p. 724; Couch,
Fish. Brit. Isl., vol. I, p. 57, tab. XIII; Dum., Hist. Nat.
Poiss. (N. su. a Buff.), tom. I, p. 455 ( + Scymnus [Lce-
margus'] brevipinna , p. 456); Gthr (Lcemargus), Cat. Brit.
Mus., Fish., vol. VIII, p. 426.

Squalus glacialis , Fab., Fisch. Isl., p. 23; Nils., Prodr. Ich-
thyol. Scancl., p. 116.

Scymnus micropterus, Val., Nouv. Ann. Mus. Hist. Nat., tom.
I (1832), p. 454, tab. 20.

Squalus (Scymnus) Gunneri, Richards., Fna Bor. Amer., part.
Ill, p. 313.

Obs. As we were compelled above to recognise the Linneean
specific name of catulus for the Black-mouthed Dog-fish, although it
was originally intended for another species, here too we must acknow-
ledge the specific name of carcharias, which was transferred by Gun-
Nerus in precisely the same manner to the Greenland Shark, all the
more, since Linn^us himself included Gunnerus’s determination among
the synonyms of his own Squalus Carcharias.

Next to the Basking Shark the Greenland Shark
is the largest Scandinavian fish. Specimens so large as
those known from hearsay by Gunnerus — nearly 8 m.
in length — must be reckoned as exceptions; and the
statement may perhaps depend on a confusion with the

Basking Shark. Both Faber and Rink give 18 feet
(56 dm.) as the greatest length of the Greenland Shark,
and according to the latter its average length on the
coast of Greenland is only 25 — 28 dm. Nilsson as-
sumed the maximum length might be fixed at 15 feet
(4 V2 in.). In Trondhjem Fjord, according to Storm,
the Greenland Shark hardly exceeds a length of 3 m.

The body is of a handsome and regular, com-
pressed fusiform shape, sometimes, like that of the Sagre,
somewhat approaching in front to the so-called hog-
backed form. The head is strongly arched above, the
snout convex both above and below, anteriorly depressed
towards the edges, which meet in a tip parabolically
rounded in the horizontal plane. The back is broad
in front, in uninterrupted continuation of the arched
head; but behind the perpendicular from the base of
each pectoral fin it is compressed, and acquires in the
median line a carina, anteriorly imperceptible, gradu-
ally ascending, and continued by the anterior margin
of the first dorsal fin. The median line is also some-
what keeled between the two dorsal fins and in front
of the caudal fin, the beginning of these fins being
consequently difficult to fix externally. The peduncle
of the tail, however, behind the second dorsal fin and
the cloaca, often — as may occasionally be observed in
the two preceding species — has a longitudinal impres-
sion both above and below, and on the lower part of
each side we find, as in the Picked Dog-fish, a more or
less distinct longitudinal carina. The greatest breadth
of the body, just in front of the branchial apertures,
is about 2/3 of the greatest depth, at the beginning of
the first dorsal fin, where the breadth is more variable,
according to the distension of the abdominal cavity®.
The peduncle of the tail also shows considerable varia-
tion of form, the breadth (thickness) of its shallowest
part varying at least between 71 6 and 82 % of the
depth thereof.

During the growth of the body from a length of
15 to one of 19 dm. the length of the head decreases
from about 201/2 to 18 V2 % of that of the body0. The
eyes are round and rather small, measuring Vs — Vio
of the length of the head and about 1/5 — 1/6 of the in-
terorbital width. They are closely surrounded by the
margins of the orbits, which are not very large. The

a According to Valenciennes the breadth at this point is 7/]0 of the depth.
b According to Valenciennes down to 50 %.

‘ Id a stuffed specimen 3 ]/2 m. long the length of the head is only 17 V2 % of that of the body; but the measurement is, of
course, uncertain.

GREENLAND SHARK.

1169

nostrils are similar to those of the two preceding spe-
cies, but are set at about the end of the first third of
the snout, measured from its tip. The broad and but
slightly curved mouth has free corners and dee]) grooves
outside them. The teeth (fig. 342, a and b) are most
like those of the Sagre. In the upper jaw they are set
in 5 — 7 transverse rows. Lateral cusps are wanting or,
if present, small and tuberous. The base of the tooth
is higher than in the Sagre, more quadrangular, and
on the broad anterior surface has a terete carina, con-
tinued to a greater or less distance on the cusp. In
the lower jaw the first two rows of teeth are commonly
erect, the compensatory teeth, on the other hand, the
five or six posterior (inner) rows, directed downwards.
Here too the lamellar base is higher, in proportion to
the obliquely set, serrature-like cusp. In the lower
margin it has a median sinus and on the anterior sur-
face above the sinus a median carina, with one or more
holes on each side. In comparison with those of the
two preceding species the spiracles are small and set
high. They have the appearance either of triangular
or elongated, narrow openings, their length not exceed-
ing the diameter of the iris. Their distance from the
tip of the snout measures about 2/3 (61 — 68 %) of the
length of the head, and their distance from each other
about half as much (about 30 % of the length of the
head). The gill-openings are hardly middle-sized, the
height of the first being about twice the diameter of
the iris, but their height gradually increases from the
first to the hindmost opening. They differ from those
of the two preceding forms in that the last aperture
lies a little in front of, not quite close to, the base of
the pectoral fin.

All the fins are distinguished by their insignificant
size. The two dorsals are obliquely quadrangular,
longer than high, with the upper anterior angle roundly
truncate and the upper posterior elongated to a point.
The first dorsal begins at about the end of the fourth
and ends at the termination of the fifth tenth of the
body; the second begins just in front of the end of the
seventh tenth and ends at the termination of the third
quarter. The distance between the posterior extremities
of their bases is consequently equal to that between the

second dorsal and the upper tip of the caudal fin. This
distance is also similar to that between the beginning
of the pectoral insertion and the end of the base of
the first dorsal. The caudal fin is comparatively broad,
but short, its form being more nearly approximated
than in any of the preceding Sharks to the Teleostean
caudal fin. A peculiarity almost or entirely absent in
the two preceding species is, however, that the hind
margin is interrupted above by a shallow break, calling
to mind the incision usually present in the other Sharks.
The upper anterior margin of the fin measures about
18 %, its lower anterior margin about 12 %, of the
length of the body. Among the paired fins the ventrals
are not unlike the dorsal fins, but the pectorals are
quadrangular in a different manner, as if the oval form
were truncated and concavely emarginated at the broad
end (the top). The relative position of the ventral fins
to the second dorsal is about the same as in the Sagre;
they begin at a distance from the tip of the snout
measuring about % (63 %) of the length of the body.
The distance between the pectoral fins and the tip of
the snout is rather more than 1/i (26 — 28 %), and
the length of these fins at the anterior margin about
7 — 10 %, of the length of the body.

The shagreen and the coloration are noticed above.
The small spiny scales are of the same type as in
Scymnus, and resemble in miniature the spines of Eelti-
norhinus, but are not so densely grooved. The white
teeth of the mouth stand off sharply, as in the Sagre,
against the dark ground of the body. Soon after the
Greenland Shark is drawn out of the water, the loose
epidermis peels off, and the body assumes a bluish gray
tint. v. Wright’s figure (PI. LII, fig. 3) represents the
fish as it appears immediately after its capture.

The Greenland Shark is really an Arctic species,
its affinity to Acanthorhinus rostratus , a form observed
in the Mediterranean and on the coast of Portugal,
being as yet unelucidated “. Fisheries for the Green-
land Shark have long existed on the coasts of Green-
land, Iceland, and Norway, and in recent times Spits-
bergen has also become a fishing-station. On the North
American coast the species goes south to Cape Cod.
Isolated specimens have been met with in the North

° We need hardly remind the reader of the erroneous conceptions which stuffed specimens either of Sharks or Rays may give. With
the other close resemblances to be observed between Acanthorhinus cavcharias and A. rostratus, it would appear quite possible that the very
strong prolongation of the occiput in the latter, according to Capello’s (Jorn. Sc. Math., Phys., Natur., Lisboa 1869, p. 146, tab. IX, figs. 2,

2 a, 3 h) and Canestp.ini’s (Mem. R. Accad. Sc. Torino, ser. 2, tom. XXI, p. 364, tab. II, figs. 2 — 4) figures, may be due to stretching in
the process of stuffing. The slenderness of the body and the great size of the eyes in Acanthorhinus rostratus may be characters of youth.

1170

SCANDINAVIAN FISHES.

Sea and the Channel (off the mouth of the Seine), as
well as in the Cattegat, even off Mount Kullen, where
Baron Gyllenstjerna secured a specimen during the
spring of 1831, and on the north coast of Zealand,
whence Kroyer and Petersen each adduce a specimen.
On the coast of Bohusl&n the Greenland Shark is not
too scarce to have received a special name among the
fishermen, being called Hamcir (as in Norway) and, on
Koster, Blamauer or Blamag-miir. Strictly, however,
it is a deep-sea fish, with its true habitat at a depth
of some hundreds of fathoms, so that its appearance in
the upper strata and in shallow water must be regarded
as more or less fortuitous.

The Greenland Shark is sluggish and insensitive,
but rapacious. Its acute hearing and keen smell have
always been remarked by the fisherman. From the
depths and from afar it is attracted by the odour of
dead flesh, whether a whale is being cut up, a seal-
hunt in progress, or a piece of carrion be cast out as
a bait. From the carcase of the whale it tears pieces
as big as a man’s head, and cares little if the exaspe-
rated whaler stabs it with his long knife or pierces it
with a lance. It is soon ready again to renew its vo-
racious repast. To a moderate-sized Greenland Shark
a seal is a comfortable mouthful. In the stomach of a
Greenland Shark, according to Gunnerus, the carcase
of a reindeer has been found; and Faber relates that,
when horses belonging to the Icelanders venture on too
thin ice and are drowned, the Greenland Shark makes
its way up the narrowest fjords — where it never puts
in an appearance at other times — to feed on horse-
flesh. But as a rule, no doubt, its prey consists of live
animals; and in the stomach of a specimen 44 dm.
long Faber found a whole seal ( Phoca vitulina), 8 large
Cod, a Ling 1272 dm. long, the head of a large Hali-
but, and several pieces of whale-blubber. It has even
been considered dangerous to man. Fabricius relates
that it attacks the Greenlanders in their kayaks and

bites both boat and kayaker in two. The Greenlanders,
he states, therefore took care never to make a noise or
talk aloud when passing over deep spots, for fear the
Greenland Shark should hear them and come up. Rink,
hoAvever, seems to give no credence to these accounts,
and even says that the Greenlanders are only afraid of
the very large Greenland Sharks, which by their move-
ments and with their sharp skin might tear holes in
the kayaks. Faber states that in Iceland no instance
had been recorded of a Greenland Shark attacking
human beings.

As has been indicated above, ichthyologists are
still doubtful whether the Greenland Shark gives birth
to living young or lays eggs. Fabricius states posi-
tively that it is viviparous, and Faber adds that the
young are born in July and August. A female of
Acanthorhinus rostratus dissected by Cornalia3 had 6
foetuses in the oviducts, and another prepared by MM.
Gal at Nice* 6 contained 12 foetuses. Lutken, however,
pointed out the singularity of the fact that in recent
times, so far as he could ascertain, no foetus had been
found in any Greenland Shark, and also cited express
statements from Iceland, where a general conviction
obtains that the Greenland Shark is oviparous. In sup-
port of this opinion he adduces the circumstance that
the eggs of the Greenland Shark, which have been
found, inside the fish, of the size of goose-eggs, are so
numerous that an adult female sometimes contains a
barrel and a half or two barrels thereof; and such
numerousness, he says, is surely not compatible with
the development of the foetuses within the oviducts,
provided the development be simultaneous or nearly so.
The last-mentioned assumption, however, has not been
demonstrated; and so long as this is the case, we have
good reason to follow Lilljeborg and give the prefe-
rence to the earlier belief that in this respect the
Greenland Shark resembles its nearest relatives, and
gives birth to living young c.

“ See Lutken, I. c.

6 See Moreau, 1. c.

0 Long after the printing of the above in the Swedish edition of the present work, on Christmas Eve, 1894, the Royal Museum re-
ceived a female Greenland Shark that had been taken some days before on the coast of Bohuslan. The length of the body to the tip of
the upper caudal lobe was 172 cm., to that of the lower caudal lobe 158 cm. The length of the abdominal cavity was about 44 % of the
former measurement. The two pointed lobes of the liver, which gradually narrowed behind, were longer than the said cavity, the length of
the left lobe being about lpg times (136 % of) that of the right lobe l'/4 times (126 / of) the length of the abdominal cavity; and they
were therefore bent behind, their posterior extremities being directed forwards. Both were of about the same breadth, which measured, where
greatest (in front), about 1/,0 of the length of the abdominal cavity. The gall-bladder was rather large, twice as long as broad, and about
equal in length to the base of the ventral fins or about V8 as long as the abdominal cavity. On each side of the ventrally median peritoneal
fold joining the gall-bladder to the diaphragm, opened the anterior end of the oviducts, which were of almost uniform thickness throughout

GREENLAND SHARK.

1171

The fishery for the Greenland Shark is fairly lu-
crative in many localities. On the coast of Norway,
and occasionally on that of Bohuslan, the Greenland
Shark is sought after principally for the sake of its
liver, from which oil is extracted; but in Iceland and
Greenland the flesh is eaten both by man and dogs.
A large fish contains, according to Rink, two barrels
of liver, which, when melted down, yields oil to the
amount of about 53 %. The flesh is also oily, but the oil
exudes in process of drying, or may be removed by press-
ing while the fish is fresh. The Icelanders, like the Nor-
wegians in Gunnerus’s time, cut the flesh up into rav
and rdklinga. The Greenlanders prefer to eat it rotten.
When fresh, it tastes like Halibut, according to Rink,
and has an agreeable, white appearance, but is some-
what coarse and tough.

In Norway the Greenland Shark is caught on large
hooks, turning freely on a swivel, and with a snood
of slender iron-chain, which the fish cannot bite asun-
der. The line is a centimetre thick or a little thinner,

and is Avound up on a small Avindlass, made fast to the
bulwark of the vessel. With this tackle the bankers
lie at anchor in the open sea between Norway and
Bear Island or Spitsbergen, in 100 — 200 fathoms of
water. The method practised is the same on the coasts
of Iceland and South Greenland; but in North Green-
land, according to Rink, the Greenland Shark is taken
with far less trouble and much greater success through
holes in the fast ice. The strong-smelling bait is dang-
led only a feAv metres below the surface, or it even
happens that the Shark may be enticed right up to the
hole, where it is captured with a gaff. The sluggish-
ness of the Greenland Shark is such that it sometimes
alloAvs itself to be taken on an ordinary Cod-line.

Among the parasites of the Greenland Shark is
often observed a crustacean, Lernceopoda elongata,
which attaches itself to the cornea of the eye, and
Avhich Avas first figured by Scoresby and, after him,
by Couch in his figure of the Greenland Shark.

their length, and which crooked backwards along the posterior surface of the diaphragm, each to its respective side, below and along the
spinal column. The two ribbon-shaped ovaries were symmetrically suspended on each side in the anterior part of the abdominal cavity and
were of about the same size, 30 cm. (2/3 of the length of the said cavity) long and 2 1/i cm. broad. The eggs were innumerable, and hardly
distinguishable to the naked eye. At no point was there any indication of an earlier development in one part of the egg-mass than in an-
other. This specimen too seemed accordingly to bear out Lutken’s opinion as to the reproductive operations of the Greenland Shark; but
the question cannot be fully solved until older and ripe females have been examined.

° See above, p. 415.

1172

SCANDINAVIAN FISHES.

PISCES CYCLOSTOML

Skeleton cartilaginous only at the head and tail, elsewhere fibrous. Notochord unaltered
throughout life: no complete spinal column. No true maxillary or mandibular parts. No
true gill-arches, the branchial lamellae being serially arranged on the inside of sacs each
with a separate aperture or all having a common aperture on each side of the body. One
nasal aperture only, in a developed state dorsal. No paired fins. No shoulder-girdle, no pelvis.

Hitherto all the fishes under our consideration have
been of indisputable piscine nature. We now approach
the primordial limits of the vertebrate type, where the
differences between the various classes find more or less
imperfect expression in the structure of the several
forms. Our knowledge of the geological evolution of
the vertebrates is indeed deficient as yet in many re-
spects, and fails us when we have to determine the
origin of the classes. Besides, many remains cannot
be expected to be preserved of animal forms which for
the most part were indubitably without firm skeleton.
But morphology and ontology (the development of the
individual) point so clearly to intermediate forms which
probably had representatives once in the living world,
that the imagination has full right to picture an ex-
tensive area of the system, principally tenanted by ex-
tinct animals in which the characters of the different
classes were combined or not yet differentiated. The
Cyclostomous order survives during the modern age in
such a position. The points of resemblance between

these fishes and the rest are indeed so many that none
can well deny the piscine nature of a Myxine or a
Lamprey; but several portions of their structure lend
themselves best to explanation by a comparison with
the organs of batrachian larva?.

A very salient proof of the low rank occupied by
the Cyelostomes as vertebrates lies in the metamorphosis
pointed out by A. Muller" in the transition from Pride
(. Ammoccetes ) to Lampern (Petromyzon). The trans-
formation is here as great in certain respects as that
undergone by many invertebrates, and is also accom-
plished, at least in part, by histolysis (dissolution) of
the organs to be transformed.

The order contains two families, one of which, that
of the Myxine, never acquires any true (discoid) suc-
torial mouth, the name of Cy cl ostomi (with round mouth)b
— which we retain in accordance with the rules of
zoological nomenclature — being hence less appropriate
than the later Marsipobranchii ( with saccate gills)c.

Fam. petromyzonidh.

Seven gill-openings on each side of the forepart of the body (the throat). The nasal duct terminates ccecally
and in a developed state lias its inner ( posterior ) end situated between the anterior part of the notochord and the

(esophagus, under the former and above the latter.

On account of the above-mentioned intermediate well as by the following family, with relation to the

position occupied by the family of the Lampreys, as other classes of vertebrates, its study has possessed

“ Mullers Archiv 1856, p. 323.

b C. Dumeril, Zoologie Analytique, Paris 1806, p. 100.

c Bonaparte, Selachiorum tabula analytica , N. Ann. Sc. Nat., BologDa, Ann. I, tom. II (1838), p. 214, and Systema Ichthyologies,
ibid., Ann. II, tom. IV (1840), p. 277.

LAMPREYS.

1173

great morphological significance. In the structure of
the skeleton of the Lampreys the difference between
the internal (vertebral) and the external (extravertebral)
parts is far more prominent than in the fishes we have
hitherto considered, the latter parts being here endowed
with a more preponderating importance. The purely
vertebral parts appear in the Lampreys merely as the
framework of the cranium and its nearest appendages
and as very imperfectly developed neural arches (fig.
343, arc ) on the sides of the myelon and the unaltered
notochord {etch). In the vertical fins too cartilaginous
rods are developed which are applied to the neural
arches in such a manner that it has been proposed to
interpret them as representing spinous processes both
superior (neural) and inferior (hsemal). The skull itself
is an almost straight continuation of the fibrous me-
dullispinal sheath, which has been dilated, and which

grows, on each side of the brain, to form a more or
less complete roof for the cerebral cavity (tegmen cranii,
t). Laterally and downwards the floor expands in three
places on each side into flat processes. The foremost
pair of these processes projects from the ethmoidal part,
the posterior two pairs issue from the sphenoidal part.
The former is evidently homologous with a palatine arch
(pal), the latter answer to the upper parts in the sus-
pensory apparatus of the mandibular and hyoid arches.
But no mandibular arch proper is developed in these
fishes. The tAvo posterior processes on each side are
confluent at the base; but the anterior of them (ptg)
groAvs outwards and forwards, in a loop below the eye,
until it coalesces Avith the top of the foremost (palatine)
process (pal). In this manner a flat ring is formed,
surrounding an aperture (fenestra snborbitalis or fissura
orbitalis inferior, fo), and composing the bottom of the

etch

Fig. 343. The cartilaginous skeleton of the anterior part of a Lampern ( Petromyzon Jluviatilis ), 3/2 nat. size.
ann, annular cartilage of the oral disk; arc, rudimentary neural arches; aw, cartilaginous capsule of the auditory apparatus; hr , branchial basket;
cell, ceratohyal cartilage; ctr, posterior cartilage of the roof of the mouth ( cornua trabecularum) ; eph, epihynl cartilage; et, ethmoidal part
of the cranial floor; etch, outer sheath of the chorda dorsalis; fo suborbital fenestra; la, anterior lateral cartilage of the roof of the mouth;
lat, lateral labial cartilage; Ip, posterior lateral cartilage of the roof of the mouth; med, median cartilage of the roof of the mouth; n, ol-
factory capsule, with the tubular orifice of the nasal duct; pal, palatine part of the roof of the mouth; phy, posterior hyoid cartilage;
pro, pericardial cartilage; ptg, pterygoid part of the roof of the mouth; si, sublingual cartilage; sph, sphenoidal part of the cranial floor;
spl, symplectic (? hyomandibular) cartilage; t, lower part of the cranial roof ( tegmen cranii).

ann lat

is invested in the developed Lamprey with a more or
less complete cartilaginous capsule (£) originating from
the skeletogenous layer round the anterior end of the
notochord. The part first formed, the cranial floor,
ansAvering to a sphenoidal region (sph) applies itself to
and coalesces Avith the already present auditory capsules
(ait). In a fonvard direction are developed the so-called
cranial trabecula?, around the cerebral appendage (hypo-
physis, posterior nasal duct) and the under-brain
(infundibulum); and after their union in front into an
ethmoidal part (et) they advance to the so-called cornua
trabecularum (ctr), Avhich coalesce into a disk in front
of the nasal capsule (n). UpAvards the cranial floor

cranial orbit. To the top of the hindmost process is
attached the hyoid arch proper. The upper part of
the latter is an unbroken doAvmvard continuation of the
said process, and represents an epihyal bone (eph)-, the
loAver (ventral) part, Avhich is suspended at right angles to
the loAver tip of the upper part, ansAvers to a ceratohyal
bone (cell). A certain similarity to this structure of
the cranial roof of the palate and pharynx Ave have
indeed seen in the Chimeeras (fig. 294, p. 1065), Avhere
a coalescence, suggestive of a development presumably
very primitive, has taken place between the pterygo-
palato- quadrate parts and the cranium. But the closest
resemblance to this structure in the Lampreys meets us

1174

SCANDINAVIAN FISHES.

in batrachian larva? (fig. 344), a reminder that in their
origins these two types were structurally not far re-
moved from each other. The tongue of the Lampreys
contains a long rod-like supporting apparatus, capable
of protrusion and retraction, and suspended under the
ceratohyal. This apparatus consists of two cartilages,
an anterior, semipateriform, excised in front, situated
in the mouth, and a posterior ( phy ), much longer, the
posterior extremity of which projects into the branchial
basket. It is interpreted as homologous with a basihyal,
and the anterior cartilage bears the lingual teeth (tig.
345), at the very tip a curved, transversal dental plate
(aln), with firm, rounded and obtuse teeth, and further
back two plates (pin), lying more loosely in the gums,
meeting anteriorly in an ellipse, and furnished with
small but acute, retral teeth.

The whole oral apparatus of the Lampreys, which
is abundantly supplied with cartilages, paired and un-
paired, pateriform or lamellar and rod-like — one of

Oc pro

Ip pal spl hy fo Inn

Fig. 344. Cartilaginous parts etc. in the head of a tadpole about
1 in. long, at the time when the hind legs begin to appear.

About 9 times nat. size. After Parker.
ctr, n, and et, as in preceding figure; Eh, Thl, and Me, the three an-
terior divisions of the brain; Oc, eye; als (alisphenoid), pro, (prootic),
pto (pterotic), so (supraoccipital), eo (exoccipital), and op (opisthotic),
all answering to t in preceding figure; dr, section of the skin; G, gill-
opening; hr, as in preceding figure; hm and spl, answering to spl in
preceding figure; fo, as in preceding figure; hy, answering to phy in
preceding figure; Ip, lips (dermal flaps); mck, Meckelian cartilage;
mdh, mandible; qu, quadrate bone; max, upper jaw.

them annular — belongs to the extravertebral structures.
Below the cornua trabecularum (fig. 343, ctr) is medially
situated an invertedly pateriform cartilage (the median
cartilage of the roof of the mouth, med, Muller’s “vor-
dere Deckplatte des Mundes”), which has behind it on
each side a flatly convex cartilage of a rounded quad-
rangular form (the posterior lateral cartilage of the
roof of the mouth, Ip, Muller’s “hintere Seitenleiste”)
and at about the middle of its side-margin a narrower,
more terete plate (the anterior lateral cartilage of the

roof of the mouth, la, Muller’s “vordere Seitenleiste”).
Under and before the said median plate lies the annular
cartilage (ann, Muller’s “knorpeliger Lippenring”), the
bearer of the largest teeth, which has behind it a me-
dian plate (the sublingual cartilage, si, Muller’s so-
called “Zungenbein”), situated under the tip of the hyoid
bone, in front expanded and convex, behind acutely
elongated. On each side of this sublingual plate lies
behind the annular cartilage a rod-like cartilage (the
lateral labial cartilage, lat, Muller’s ustielformiger
Anhang cles Lippenring es")\ and three small cartilages,
the bearers of the lateral teeth of the mouth, lie loosely
in the skin on each side of the annular cartilage.

The branchial basket (hr) too is entirely extra-
vertebral, a network of thin flat cartilaginous rods, sup-
porting the outer walls of the gill-sacs. Anteriorly this
network is suspended from the epihyal cartilage; pos-
teriorly the cartilaginous mass developes into a saccate
case for the heart, a pericardial cartilage ( prc ).

The nervous system of the Lampreys has thrown
light, especially in two points, on the morphology of

Fig. 345. Tip of the tongue and its dental plates in a Lampern
(Petromyzon fluviatilis ), X about 5.
aln, anterior dental plate of the tongue; pin, posterior dental plate

of the tongue.

the higher vertebrates. These points have reference to
the interpretation of two organs still involved in great
obscurity, the cerebral appendage and the pineal gland.

At the bottom of the human cranial cavity, as well
as on the under surface of the brain in all other true
vertebrates, lies a glandiform organ whose function is
still doubtful. It is known anatomically as the hypo-
physis cerebri. In man it is about as large as a pea,
oblong in form and transversally set, lodged in the de-
pression on the upper surface of the sphenoid bone
(the so-called sella turcica ) and attached to the brain by
a funnel-shaped prolongation thereof in a downward
direction from the bottom of the third cerebral ventricle.
It consists of two parts, an anterior, the development
of which we shall now describe, and a posterior, whose
development originates from the brain. Great impor-
tance was formerly attributed to it both by physiologists
and psychologists. By Galen, in the second century

PITUITARY GLAND.

1175

A. D., and after his time far into the middle ages, even
if the conception was modified in several manners, the
cerebral hypophysis was explained as an extremely im-
portant organ for the purification of the fluids contained
in the brain. It was also called the gland for the se-
cretion of the nasal mucus ( glandula pituitaria), an error
which was not refuted until the seventeenth century.
Another opinion was advanced, which even at the be-
ginning of the present century was maintained by so
eminent an anatomist as Meckel, namely that it secreted
some kind of important fluid, which was supplied to

foetal life than afterwards and is comparatively larger in
young persons than in old, as is the case with several
other organs whose significance appears to be more
historical than physiologically clear.

Better than its function, however, we know the
origin of the hypophysis". In man it arises as an in-
volution of the embryonic ectoderm in the region which
afterwards becomes the roof of the mouth cavity. This
involution rises, towards the anterior end of the noto-
chord. It makes its appearance before the mouth ca-
vity proper is formed, and long before the mouth has

Fig. 346. Longitudinal section of the forepart of larvse of Petromyzon Planeri , after A. Dohkn. A three days, B six days, C seventeen

days after exclusion.

Ch, chorda dorsalis; Ent, entoderm; Ep , epiphysis; Hy , hypophysis; Inf , infundibulum; M, stomodreum, future mouth cavity; Max , upper
jaw; Mdb, lower jaw; MT, mouth tentacle; N, nasal sinus, future nasal cavity; Thy, thyreoid gland; Tub. cin, tuber cinereum.

the third ventricle of the brain. In modern days phy-
siology does not seem to devote very much attention to
its interpretation; it is most generally referred to the
so-called blood-vascular glands, which it resembles in
the greater part of its structure. Remarkable is the
circumstance that it has a more robust growth during

broken a passage through the pharynx to join the in-
testinal canal. In the form of a tube it grows up to-
wards the rudiment of the brain; and when the floor
of the skull is developed the lower part of this tube
closes, in the region where the presphenoid bone sub-
sequently coalesces with the postsphenoid; the coinmu-

“ The first investigator hereof was Rathke, Ueber die Entstehung der Glandula pituitaria , Mullers Archiv 1838, p. 482.

Scandinavian Fishes.

148

1176

SCANDINAVIAN FISHES.

nication with the mouth cavity is interrupted, and the
hypophysis lies enclosed in the cranial cavity. Such is
the development in all the higher vertebrates. In the
lower, on the other hand, as for instance in the frogs,
the cerebral hypophysis appears still earlier, before the
mouth cavity is developed enough to contain the origin
of this appendage within its region. In Pelobates fttscus
Goette" has shown that the origin thereof is visible
even before the future mouth cavity is indicated in any
way. There, accordingly, the cerebral hypophysis is at
least as primordial an organ as the mouth. In the
Lampreys Dohrn0 and, after him, KupffeiT saw a stage
of development (tig. 346, A and B) in which the nasal
cavity (N), the hypophysis (////), and the mouth cavity
(M) had begun to appear on the under surface of the
rudimentary head in the form of three uniserially (in a
sagittal row) arranged impressions of the ectoderm. But
here the homologue of the hypophysis becomes much
more than, and something quite different from, a mere
appendage of the brain. Such an appendage is indeed pre-
sent in the Lampreys; the said rudiment here too grows
in a tubular form upwards and backwards towards the
tip of the notochord, but it also moves upwards round
the snout, continually growing, and becomes the so-
called nostril of the Lampreys, which throughout their
life opens on the dorsal side of the head and thence
extends in a tubular form backwards and downwards
over the pharynx. During its growth and migration as
described above the nostril draws within its limits the
true olfactory apparatus (fig. 343, n), a capsule imper-
fectly divided internally into two chambers, and deve-
loped from the nasal cavity (fig. 346, B ; N) that once
lay before the rudiment of the hypophysis. In this
manner the Cyclostomes attain a unique position among
the vertebrates. They possess, it is true, a well-developed

olfactory apparatus, but only one nasal capsule and one
nostril. Heckel therefore called them Monorhini.

The hypophysis is accordingly a very primitive or-
gan — at least as ancient as the mouth — but in the
highest animals has lost its most essential significance L
Similar is the history of the pineal gland.

In 1881 Luys published a psychology* 6 as the result
of his works on the brain and its functions, giving us
in a very popular way means to understand the relation
between the different parts of the brain. By numerous
and successful sections he traced the course of the nerve
fibrils to the brain and within this to the connexion
between its countless ganglion cells. Contemporaneously
with these and other anatomical works Ferrier in Ene-
land, Fritsch and Hitzig and, after them, Munk in Ger-
many, had published their famous attempts to discover
in the superficial layers of the brain the so-called psy-
chomotor centres and to define their limits. These and
other later researches in many points have modified the
views advanced by Luys, especially as to the arrange-
ment of the sensorial tracts, but for our present purpose
we may leave those discrepancies out of sight. The
brain of the higher vertebrates, with its rich psychical
life, is, however, of so complex a nature that the mys-
teries of that life would probably have never been
solved by the methods of natural history, had not more
simple material been placed at the investigator’s disposal.
The study of the lowest fishes in particular has led to
the one comprehensive conclusion after the other. Thus
we shall, no doubt, be able at length to trace the ori-
gin of different psychical faculties in connexion with the
different development of some part or other of the brain.
Already in 1840 Johannes Muller foreshadowed this,
when in his work on the nervous system of the Myxines
he chose the cerebral structure of these fishes and of

a Entwickelungsgeschichte der Unke , p. 288, taf. II, figs. 34 — 38.

6 Mittheil. Zool. Stat. Neapel, Bd. IV, p. 172, taf. 18.

c Arch. f. Mikr. Anat., Bd. 35 (1890), p. 537, taf. XXXI, fig. 62.

d In the Quart. Journ. Micr. Sc., N. ser., vol. XXIII (1883), pp. 349, cett., Hubrecht has tried to show that both the hypophysis
and the notochord are inheritances from the worm-like predecessors of the vertebrates. The hypophysis, he contends, represents the proboscis
of the flat- worms and the notochord their proboscidian sheath. At the limit between the proboscis and the sheath- — where the latter sutlers
invagination to receive the former when retracted — lies. the principal mass of the Platyelminth nervous system; and at the corresponding

point in the skull of the lower vertebrates, at the limit between the epicliordal and prechordal parts of the brain, there too do the notochord

and hypophysis meet each other.

Neither the hypophysis nor the notochord would thus be peculiar to the vertebrates; they would have their prototypes in the lowest
worms. The hypophysis lies under our brain in the most protected spot throughout our body, as even Galen remarked, as if it were a most
important and delicate organ; but it lies there, according to Hubrecht, merely as a relic, a vestige of an organ which serves the lowest
worms for purposes of touch, otfence, and defence. In us adults the notochord has entirely disappeared; but during our foetal life and that

of the other higher animals, as well as in the adult state of lower vertebrates, this string is the axis round which the vertebrae are developed

and their bodies chondrified or ossified; — and this string, says Hubrecht, was once a sac within which the homologue of the hypophysis
might find concealment.

e The Brain and its Functions , Intern. Scient. Series, vol. XXXVII, Bond. 1881.

PINEAL GLAND.

1177

the Lampreys as the basis of his explanation of the corres-
ponding structures in the higher vertebrates: — and even he
could declare that “the structure of the brain in the Lam-
prey and Pride is a perfect parallel to the earliest stages
in the development of the brain of the higher vertebrates.”
From the results of the history of evolution Beard"
drew a picture of the worm-like ancestors of the ver-
tebrates, ancestors which have all disappeared from the
earth, so far as we know, without leaving a trace of
their existence. Their body, he says, was anteriorly
furnished with several, at least eleven pairs of gill-clefts,
and above, i. e. at the dorsal termination of each gill-
cleft, lay a sense organ in the skin, probably to warn
the animal of the approach of any dangerous object
towards the gill-bearing parts of the body. The nearest

guide, sense organs of the same kind have originated
the development of all the other higher senses. Even
in the mammals Froriep4 has traced the same course
of development for three of the cranial nerves ( facialis ,
glossopliaryngeus, and vagus) as Beard found to obtain
in Rays and other fishes for all these nerves, including
those of smell, hearing, and taste. In order to obtain
their basal ganglion these nerves grow from the sides
of the rudimentary spinal canal downwards and out-
wards to the inner surface of the embryonic skin and
meet a thickening thereof at a spot answering to a prim-
ordial sense organ of a gill-cleft. The ganglion is form-
ed by the growth and multiplication of the epidermal
cells at this point; but the rudiment of the primordial
sense organ disappears after having thus done its duty

C

Fig. 347. Three sketches of the brain and medulla oblongata in a Pride ( Petromyzon in the Ammoccetes stage), magnified. After Wieders-

heim. A, from below; B , from the right; C, from above.

I — X , cranial nerves in order from in front; 1, first pair of spinal nerves; Cb, cerebrum; Cbl , cerebellum; Ep, epiphysis; Hyp , hypophysis;
L. ol., lobi olfactorii; ME, mesencephalon; Med, medulla; Mob, medulla oblongata; S. v., saccus vasculosus; Thl, thalamencephalon.

parallel to the sense organs of the gill-clefts in their
original form is thus afforded by the lateral-line system
of fishes and batrachians, with their marvellous deve-
lopment to the possession of a sense which we as yet
scarcely understand, a specification of the sense of pres-
sure, enabling these lower vertebrates to appreciate cer-
tain undulations of the water, and thus to detect at a
distance the presence of objects or the approach of
changes which under the same circumstances could
hardly be perceived by the senses of the higher verteb-
rates. But if the history of evolution be a faithful

in the history of evolution, without ever having felt
a sensation. Even the formation of the eye may be
referred, considering the manner of origin shown by
the ciliary ganglion and the lens, to the same scheme.
Originally a number of these sense organs would thus
seem to have been present in the skin of the verteb-
rates; but according to this assumption most of them
have become useless since the development of the senses
in their present form. The course of this process is
known at least in the case of one such organ, the pineal
gland, whose fate has been traced with fair precision.

a Syst. branch. Sense Org., Quart. Journ. Micr. Sc., Nov., 1885, and Develop. Periplier. New. Syst., same periodical for Oct., 1888.
b Ueber Anlagen von Sinnesorgo.ncn am Facialis, Glossopliaryngeus und Vagus etc., Arch. f. Anat. u. Physiol. 1885. Anat. Abtheil.

1178

SCANDINAVIAN FISHES.

When Ahlborn resumed the study of the brain of
the Lampreys in 1883“, its structure appeared in a new
light, different from that shed upon it in Joh. Muller’s
day. In the brain of the lower vertebrates (fig. 347)
a distinction is now drawn in the first place between a
posterior and an anterior part. The posterior, epichor-
dal part — so called because it extends as far forward
as the subjacent chorda dorsalis — contains the medulla
oblongata (prolongated spinal cord or afterbrain, Mob)
and the small homologue of our little brain (hindbrain
or cerebellum, Cbl), but in the Lampreys is almost as

15 6 10

Fig. 348. Diagram of a longitudinal section through the human ce-
rebrum and medulla oblongata, to illustrate the higher functions of the
brain, according to the assumptions advanced by Luys.

7, thalami optici; 2 , corpora striata; 3, course of the propagation of
acoustic impressions (labyrinth of the internal ear). These impressions
arrive in the corresponding centre (4), are radiated towards the sen-
soriurn (3), and reflected at 6 , to the large cells of the corpus striatum,
and thence at 7 and 7', towards the motor regions of the spinal axis.

8. Course of sensitive impressions. These are concentrated (at 9) in
the corresponding centre, radiated thence into the plexuses of the sen-
sorium (10), reflected to the large cortical cells (11), and thence pro-
pagated to the large cells of the corpus striatum, and finally to the
different segments of the spinal axis. 13. Course of optic impressions.
These are concentrated (at 14) in their corresponding centre, then
radiated towards the sensorium (at 15). They are reflected towards the
large cells of the corpus striatum and afterwards propagated to the
different segments of the spinal axis; 18, 19', 19", the antero-lateral
fibres from their point of origin in the corpus striatum, are invested by
the elements of cerebellar innervation which begin to appear in the
peduncles (19), to become considerably thicker at 19' , on a level with
the region called the pons, and to diminish insensibly on a level with
the medullary regions, 19" . — 20, peripheral expansion of the ol-
factory nerves.

large as the whole anterior portion, the so-called pre-
chordal part. All the cranial nerves, except those of

sight and smell, have their roots in this posterior part
of the brain. In the prechordal brain the homologue
of the brain proper (forebrain or cerebrum, Cb) of the
higher animals is merely a small, hollow prominence
on each of the two olfactory lobes (. L . old). The two
parts of the brain which are incomparably most deve-
loped in man — the cerebrum and cerebellum — are thus
the smallest in the Lamprey; and but little, though
somewhat, greater is the development of the cerebrum
in true fishes (Teleosts) and batrachians. Merely for
anatomical reasons it has been possible to prognosticate
that in these animals the cerebrum is simply a gathering-
place and a control-station for the sensations coming
from the organs of smell. In 1868 Goltz successfully
extirpated the cerebrum of frogs, which still lived for
some time ‘‘with their intelligence preserved,” as he
expressed it; and in 1886 the same experiment was
performed on fishes by Steiner at Heidelberg and Vul-
pian in Paris. A Carp which lived six months after
the removal of its cerebrum (forebrain), the other parts
of the brain, even the olfactory lobes, being left intact,
showed no other irregularity of behaviour, according to
Vulpian, after the wound had healed, than that it was
apparently destitute of the sense of smell. Yet in the
higher vertebrates that part of the brain of which it
was deprived, is the foundation for the subsequent de-
velopment of the material substratum in everything that
bears the name of consciousness. Thus in the fish it
is only those ganglion cells with which the conscious
apprehension of the impulses of smell is bound up that
lie there. Another part of the prechordal brain appears
in fishes to occupy exactly the same relations to the
stimuli of sight. This is the postero-superior part,
the so-called midbrain (ME), which on the dorsal side
is closely applied to or even confluent with their little
brain. The first control-stations for the sensations of
sight within the region of the brain, however, lie in
the antero-inferior part of the median portion of the
prechordal brain, the so-called ’tweenbrain (fig. 347,
Till). Originally, as here in the Lampreys, the ’tween-
brain and midbrain enclose that cavity of the brain
known as the third ventricle, and this they also do in
the embryos of the highest vertebrates; but in the adult
brain of the latter they have sunk down to the bottom
of the ventricle; and before this has happened, great
revolutions have taken place in the surroundings of
that ventricle. The midbrain has dwindled into the

Unclers. tib. das Gehirn der Petromyzonten, Zeitschr. f. Wiss. Zool., Bd. 39.

PINEAL GLAND.

1179

corpora quadrigemina, and thus become a part of the
enrolling stations for the nervous action which pro-
ceeds, according to Luys, between the surface-layers
of the cerebral hemispheres and the central gray matter
of the spinal column, i. e. between the psycho- intel-
lectual and the vegetative life. The ’tweenbrain, on
the other hand, has developed into the optic thalami,
which now, hoAvever, have a far greater importance
than that of mere collectors of the visual sensations,
for hither, according to Luys (tig. 348), stream all
the sensations not yet idealised, but destined to the
consciousness. Here Luys distinguished between four
centres, well-detined gangliar masses in each optic
thalamus ( 1 ), the foremost of which receives and trans-
mits the impulses of smell {20), the second those of
sight {13 — 14), the third those of touch {8 — 9), the
fourth those of hearing {3 — 4). Before and outside
the optic thalami lies the primitive floor of the cerebrum

was a time when it was supposed that the soul had a
fixed and special station in some part of the brain, and
Descaiites assumed that the pineal gland was the seat
of the soul: — it had such a remarkably central situa-
tion, and it was solitary, unpaired. Had the soul dwelt
in a paired organ, said Descartes, we should always
have two ideas about the same thing, and there would
be a continual conflict to determine which side should
have the mastery. The researches of modern times
have certainly divested the pineal body of most of its
reputation from this point of view; but its historical
importance claims all the greater recognition.

Before the transformation of the ’tweenbrain into
optic thalami, even before the appearance of the rudi-
ment of the true optic thalami, this part of the brain
grows laterally outwards in a vesicular form and sup-
plies the rudiments of the eyes and optic nerves, and
in a downward direction it forms an expansion known

c stp Ibrs Id A Id

sip c r

Fig. 349. Ripe female of the Lampern ( Petromyzon fluviatilis, forma branchialis ), nat. size. A, from the left; B, forepart, from above; C , the

same, from below.

c, anterior connecting branch between the frontorostral canals; stp, posterior connecting branch; r, rostral branch; so, suborbital branch;
m, mandibular branch; Im , mediolateral line; Id, dorsolateral line; Ibrs, superior brancbiolateral line; Ibri, inferior branchiolateral line.

(forebrain), the so-called corpora striata {2), to which
the manifestations of consciousness that have been de-
rived from the said sources and developed in the sur-
face-layers of the cerebral hemispheres are conducted
{14— 15— 16— 17 ; 9—10—11—12 and 4—5—6) for
their first realisation, with that contribution to the
power and distinctness of the currents which is supplied
by the little brain, whose upper peduncles, according
to Luys, by means of their yellow fibrils join the gang-
lion cells in the yelloAV nucleus of each corpus striatum.
BetAveen the optic thalami and the corpora quadrigemina
lies a small SAvelling, the pineal gland {Ep in tig. 347),
the history of Avhose transformations is among the most
remarkable. In man the nervous matter of the pineal
body probably plays the same part as that of the cor-
pora quadrigemina, but is of less importance. Yet there

as the underbrain {Inf. in fig. 346); but in the roof of
the ’tweenbrain there appear two thickenings of the
cerebral Avail (behind Ep in fig. 347), situated side by
side and usually of unequal size. These collections of
nervous matter become the rudiments of the true optic
thalami. Over the ’tweenbrain the cerebral wall has
grown out into a vesicular SAvelling {Ep in figs. 346,
347), which is at first flattened, but rises, becomes tu-
biform, and grows fonvard over the brain. This is the
first appearance of the pineal gland in the loAver verteb-
rates, and in them it attains a considerable develop-
ment. In the Lampreys it is even externally visible
through the skin as a light, round spot behind the
tubular aperture of the nostril (fig. 349, B).

The pineal gland belongs, as mentioned above, to
that division of the brain with Avhich the origin of the

1180

SCANDINAVIAN FISHES.

at

Fig. 350. Section of the forepart of a Lampern ( Petroinyzon ); about
twice the nat. size; partly after Pakkek, partly after Vogt and Yung.
abr , branchial artery; ally , anterior hyoid cartilage; aln, anterior
linguo dental plate; alt , anterior lateral dental plate; anti, annular car-

tilage of the mouth (severed in two places); at, anterior dental plate;
aurc, auricle of the heart; c, ventricle; Cer I + II, prosencephalon
and tlialamencephalon; Cer III , mesencephalon; Cer IV + V, cerebellum
and medulla oblongata; cli, chorda dorsalis; coel, abdominal cavity
(coelom); ctr, cornua trabecularum; clc, ductus Cuvieri; ebr, outer gill-
openings; hep, liver; Hy, nostril; ibr I — VII, first-seventh inner gill-
openings; med, median upper cartilage of the mouth; Ml, myelon;
mlt, left mediolateral dental plate; myc, myocomma; mym, myomere;
n, olfactory capsule (the severed cartilaginous wall); oes, oesophagus;
ovar, anterior extremity of ovary; pliy, posterior hyoid cartilage; pt,
posterior dental plate of the mouth ; t, tegmen cranii ; v, anterior
part of intestine; vel, velum.

paired eyes is connected, and its manner of origin is
the same, the difference is strictly but local. In its
highest development too, it becomes an eye with optic
nerve, an eye in the middle of the skull, asymmetrical,
for its nerve root would appear to originate from the
left optic thalamus, though before this becomes a true
optic thalamus, and while it still lies, as in the Lampreys,
in the roof of the brain and bears the name of ganglion
habenulce (intermedium) sinistrum. This pineal eye,
whose functional time proper fell under the period known'
by geologists as the Mesozoic, bore within itself, how-
ever, the seeds of its destruction in the vertebrates, for
it was constructed on the ocular type of the invertebrates:
— its lens was an ependymal instead of an epithelial
(epiblastic) growth, and its retinal cells had the base
directed peripherally instead of centripetally into the
eyeball. And after the advent of the Tertiary period this
Polyphemous type disappeared from among the verteb-
rates. In many, however, as in the Lampreys, it endures
in a more or less vestigial condition. To all appearances
the pineal eye was more ancient than the paired eyes,
perhaps originally the only true eye in the worm-like
ancestors of the vertebrates, hardly elevated as yet above
the level of the invertebrates; and all the sensations des-
tined for our consciousness still follow in the optic thalami
(if we may believe Luys, fig. 348) a path laid down
through the realm where the pineal eye once held sway.

The mental faculties of the higher vertebrates
had a long history before they became what they
are. The pineal eye and the sensory organs of the
primitive gill-clefts are examples of structures at dif-
ferent grades of abortion. The first has almost ceased
to functionate at all, and terminates its existence as
a functional organ in the class of reptiles; the latter
do not extend so high in the animal series, and they
nowhere retain their original, preponderating signifi-
cance; but they have a descendant, the system of the

LAMPREYS.

1181

lateral line, which still serves one of the most impor-
tant. senses possessed by fishes and batrachians.

The lateral-line system of the Lampreys retains
perhaps a relic, at least a reminder, of the primitive
condition assumed by Beard, a pore of the system being
present above each gill-opening (fig. 349, Ibrs). In its
extension over the body this system is otherwise not
unlike that of the Teleosts. Behind the eyes run two
transverse series of pores ( c and stp) ; and the lateral
line proper is double (bn and Id). The frontorostral
branches and their subdivisions are most distinct below
and before (so) the eyes and in two parallel longitudinal
rows O') on the upper surface of the snout. Round the
lower jaw runs a mandibular branch (m), which sends
out backwards on each side a sub-branchial series (Ibri)
below the gill-openings. In most cases, however, all
these pores, except the rostral, are very difficult of
detection.

Another primitive condition is manifested, accord-
ing to Langerhans, in the skin of the Lampreys, its
outer epidermal layer (cuticular cells) being frequently
furnished, though irregularly and in patches, with cilia
— calling to mind a very common appearance in the
skin of the invertebrates. But in the Lampreys, according
to Langerhans, these cilia are dead, motionless (sensory
hairs). Neither the Lampreys nor the following fishes
show any sign of squamous growths in the skin.

The intestinal canal of the Lampreys is simple and
straight, the divisions being scarcely distinguishable from
without. Only at the extreme front and back is it attached
by a mesentery to the dorsal wall of the abdominal
cavity; throughout the rest of its course it lies free in
the cavity. The anterior part, answering to an oeso-
phagus and a stomach, is wider but has thinner walls
than the duodenum, which is lined with longitudinal
folds, one of them deeper than the rest and bending at
the middle of ' the duodenum into three or more spiral
coils. Behind the duodenum and separated therefrom
by an annular valve, lies a short but somewhat dilated
rectum. The liver lies as usual in the extreme front
of the abdominal cavity, and is rather small, but firm
and without lobes. There is no gall-bladder. The air-
bladder is also wanting. The kidneys are paired and
ribbon-like, with the sharp inferior margin hanging free
in the posterior part of the abdominal cavity, but ex-
tending forward beyond the middle of its length. Their
efferent ducts have a common opening at the top of a
papilla projecting behind the cloaca! aperture and longer

and more pointed in the males than in the females.
This common outlet, however, has on each side (to the
right and left) an opening (a slit) from the abdominal
cavity, and by this route the sexual products, both ova
and sperma, after being shed into the said cavity, are
expelled through the papillar orifice. Both the ovaries
and the testes are unpaired, suspended between the kid-
neys, without special deferent canals. When ripe, they
fill the abdominal cavity throughout its length, sur-
rounding the shrunken intestine.

Especially interesting is the foremost part of the
intestinal canal, the oesophagus, as it appears during the
metamorphosis of the Lampreys. In their larvae, the
Prides (Ammocoetes), the branchial cavity (fig. 351,
br I — 177) communicates directly on the one hand
(anteriorly) with the pharynx and mouth, on the other
(behind) with the oesophagus ( oes ) and the intestine, so
that all food must pass through the branchial cavity in
company with the water used for respiration. During

Fig. 351. Section through the forepart of a Pride ( Ammocoetes ),

about 3 times the nat. size.

abr, branchial artery, with seven openings, for the arterial branches
supplied one to each gill-opening; br I — VII , first-seventh branchial
sacs; C, heart; Oh, notochord; Fly, nostril; La , anterior lip; Lp,
posterior lip; M, posterior (inner) mouth cavity; Ml, myelon; MT,
mouth tentacles; oes, oesophagus (anterior part of intestine).

the metamorphosis, however, the posterior passage is
closed, and a new oesophagus (fig. 350, oes) is formed
above the branchial cavity, in the form of a string first
solid and afterwards hollow. The food receives a special
passage. The water used for respiration passes neither
through the mouth nor the nose, but must be received
and expelled through the branchial apertures. At the
same time the rostral region is developed, from the
cornua trabecular urn (see above) to the jaws and denti-
tion inclusive.

The family of the Lampreys hardly contains a
score of species, though more have been described.
These are distributed among four or five genera, only
one of which belongs to the Scandinavian fauna.

1182

SCANDINAVIAN FISHES.

Genus PETROMYZON.

The diphycercal caudal fin continuous above with the posterior part of the dorsal. Anterior dented plate of the
mouth furnished only at each end with an elevated tubercle or pointed tooth.

As we have mentioned above, in the Lampreys
we cannot speak of real jaws or jaw-teeth; but within
the suctorial disk of adult Lampreys are set several
horny, pointed teeth or terete protuberances, charac-
teristic of different genera and species, and to these
teeth it has been hitherto customary to give such names
as if they were set in real jaws. The oral aperture
is first surrounded, as we have seen, by an annular
cartilage (fig. 343, ann). This bears both on its an-
terior (upper) and its posterior (lower) margin an hori-
zontal dental plate furnished with more or less pointed
teeth (fig. 352), which may lie called respectively an-
terior teeth (at) and posterior teeth (pt). On each side
of the annular cartilage there lie in the sucking-disk,
next the mouth, three small loose cartilages, each set
with 1 — 3 horny teeth (protuberances), which we entitle
the lateral teeth of the mouth (alt, mlt, and pit). Out-
side these the sucking-disk has within its fimbriated
margin a number of scattered teeth, arranged in rings
or in curved rows, suctorial teeth, as Gunther calls
them (ist and est). Within the mouth itself, between
the anterior and posterior dental plates, appear the
lingual teeth, an anterior ( din ) and two posterior (pin)
curved dental plates, parallel to the posterior dental
plate of the mouth. It must be observed, however,
that when the tongue is protruded to its utmost extent
its posterior teeth project beyond the anterior ones, and
probably serve as the most powerful rasping organ of
the Lampreys. The dental character assigned above to
the genus Petr omy son is intended to express the fact
that the anterior dental plate of the mouth is neither

bipartite, as in the Pacific genus Mordacia, with a
group of three teeth on each part, nor tridentate, as
in the genus Ichthyomyzon from the west coast of North
America". The other character given above differen-
tiates all these genera from Geotria, a Pacific genus,
with the caudal fin separated from the posterior dorsal.
A deep-sea genus, Bathymyzon , “with the suproral and
infroral plates or lamina? destitute of odontoid tu-
bercles,” was distinguished by Gill*' among the fishes
taken by the American fishing-schooner Albatross at a
depth of 520 fathoms within the area of the Gulf Stream
in lat. 49° N. He remarks, hoAvever, that the only
species known (Bathymyzon Bairdii) comes very near
to Petromyzon marinus.

The species of the genus Petromyzon that belong
to the Scandinavian fauna are the same as are found
in other parts of Europe; but considering the variabi-
lity of form which here asserts itself, we distinguish
only the following:

A: Anterior dental plate of the mouth
so short that its two teeth stand close

together — Subgenus Petromyzon Petromyzon marinus.
B: Teeth on the crescent-like anterior
dental plate distant from each other

— Subg. Ammoccetes Petromyzon fluviatilis.

a, var. major: the two dorsal fins

more or less distinct-
ly divided from each
other — Petromyzon
fluviatilis.

b , var. minor: the two dorsal fins

more or less distinct-
ly confluent — Pe-
tromyzon branchialis.

a According to Jordan and Fordice a median tooth sometimes occurs on the anterior dental disk in Petromyzon fluviatilis branchialis ,
so that the genus Ichthyomyzon can hardly be retained.

6 Proc. U. S. Nat. Mus. 1883, p. 254.

CYCLOSTOMES.

1183

THE SEA LAMPREY (sw. hafsnejonogat).

PETROMYZON MARINUS.

Plate LIII, fig. 1 .

Anterior ling no dental plate medially recurved and hollowed from in front. Teeth on the anterior dental plate of
the mouth contiguous at the base. Length of the suctorial disk ivhen closed {in adult fishes ) more than half that

of the head to the first gill-opening .

Syn. Mus tela, Auson., Mos., vers. 107. Lamproye de mer, Belon,
Nat., Div. Poiss., p. 66. Lampetra a , Rondel., De Pise.,
p. 398. Lampreta 1. Lampredab, Gesn., De Aquatil., p.
592; Paralip., p. 22. Petromyzon c maculosus, ordinibus
dentium circiter viginti, Art., Ichthyol., Gen. Pise., p. 64;
Syn. Pise., p. 90. StrOm, Sondm. Beskr., pt. I, p. 297
( Neyenogen ). Sea-Lamprey, Penn., Brit. Zool. (ed. 1776),
vol. Ill, p. 67, tab. VIII, No. 27.

Petromyzon marinus, Lin., Syst. Nat., ed. X, tom. I, p. 230;
Fna Suec., ed. II, p. 106; Gdnn., Trondhj. Selsk. Skr.,
vol. IV (1768), p. 22, not.; P>l., Fische Deutscld., pt. Ill,
p. 38, tab. LXXVII; Retz., Fna Suec. Lin., p. 302;
Born, Zeitsclir. Organ. Phys. (Heusinger), Bd. I (1827), p.
170; Nilss., Prodr. Ichthyol. Scand ., p. 121; Yakr., Brit.
Fish., ed. 1, vol. II, p. 448; Sel. Longoh., Fne Beige,
p. 226; Kr., Danin. Fisk., vol. Ill, p. 1025; Nilss.,
Skand. Fna, Fisk., p. 743; Hckl, Kn., Siisswasserf. Ostr.
Mon., p. 374; Sieb., Siisswasserf. Mitteleur ., p. 368; Malm
( Lampetra ), Gbgs Vet., Vitt. Samh. Hand!., Ny t.idsf ., H.
VIII (1863), p. 87; Mgrn {Lampreta), Finl. Fiskfna (disp.
Helsingf. 1863), p. 75; Couch ( Petromyzon ), Fish. Brit. Isl.,
vol. IV, p. 385, tab. CCXLVII, fig. 1 ; Blanchard, Poiss. d.
eaux douees d. 1. Fr., p. 512; Canestr., Arch. Zool., Anat.,
Fisiol. (Modena), vol. IV, p. 184; Gthr, Cat. Brit. Mus.,
Fish., vol. VIII. n. 501; Coll., Forh. Vid. Selsk. Chrnia
1874, Tilhegsh., p. 218; 1879, No. 1, p. 106; N. Mag.
Naturv. Chrnia, Bd. 29 (1884), p. 122; Malm, Gbgs, Boh.
Fna, p. 630; Winth., Naturh. Tidskr. Kbhvn, ser. 3, vol.
XII, p. 61; Feeders., ibid., p. 94; Mor., Hist. Nat. Poiss.
Fr., tom. Ill, p. 602; Bncke, Fisch., Fischer ., Fischz. 0., W.
Preuss., p. 194; Mela, Vert. Fenn., p. 369, tab. X; Day,
Fish. Gt. Brit., Irel., vol. II, p. 356, tab. CLXXVIII; Fremy,
Coinpt. Rend. Acad. Par. 1883, No. 11 (Mars 2), p. 721;
MSb., Hcke, Fisch. Osts., p. 159; Jord., Ford., Ann. New
York Acad. Sc., vol. Ill, p. 283; Lillj., Sv., Norg. Fna ,
Fisk., vol. Ill, p. 719.

Petromyzon Lampetra, Pall., Zoogr. Ross. Asiat., vol. Ill,

p. 66.

Petromyzon Americanus, Lesueur, Trans. Amer. Phil. Soc. 1818,
p. 373 (+ Petr, nigricans , p. 385); Dek., New York

Fna, pt. IV, Fish., pp. 379 et 381, tab. 66, fig. 216; tab.
79, fig. 247; Stor., Mem. Amer. Acad. Arts, Sc., N. ser.,
vol. IX, pp. 251 et 253, tab. 38, fig. 4; tab. 39, fig. 6.

Ammocoetes bicolor, Lesueur, 1. c., p. 386; Dek., 1. c., p.
383, tab. 79, fig. 248; — forrnam larvalem hujus speciei cen-
suerunt Jordan et Fordice, 1. c.

Petromyzon appendix , Dek., I. c., p. 381, tab. 64, fig. 211.

Ammocoetes unicolor, Dek., 1. c., p. 383, tab. 79, fig. 250;
— form, larval, sec. Jord., Ford.

Petromyzon maculosus, Gronov., Cat. Fish., ed. Gray, p. 2.

The Sea Lamprey attains a length of about 1 m.
and a weight of about 1 x/2 kilo. Still larger speci-
mens, weighing at least 274 kilo., are indeed adduced d,
but are certainly rare. According to Linnjeus* Wahl-
bom caught at Calmar a Lamprey of an arm’s thickness.
In spite of its dimensions this species undergoes its
metamorphosis from the larval stage at a length of
only about 3/4 — lV2 din.'

The body is Eel-like, anteriorly terete or showing
some lateral compression, which increases more and
more behind. The greatest depth, about half-way be-
tween the last gill-opening and the beginning of the
first dorsal tin, is about 7 or 8 % of the length of the
body. Behind the body gradually tapers; but even at
the space between the two dorsal tins the depth is about
5 or 6 % of the length. In front the depth is more
constant forward to the head, the form of which is
determined by the singular snout. When the fish is
not attached to any object, the snout is not unlike an
inverted spoon; but the under surface, the sucking-disk,
can alter its position, when the fish attaches itself, till
the snout terminates in a more or less vertical, broadly

“ “A lambendis petris”.

6 Ex lampen , German. — dependere.

c “A TtstQOQ lapis et uvQio s. Livid oj sugo, quia sugendo lapidibus ad I ue ret in fluviis”, Philos. Ichth., p. 73.
d Pennant, Brit. Zool., vol. Ill (ed. 1776), p. 68; Day, 1. c., p. 359.
e Fna Suec., 1. c.

I Dekay states the length of Petromyzon appendix at 101 — 152 mm. Storer adduces a length of 76 — 178 mm. for Petrom. nigri-

cans. In Scandinavia Schagerstr6m (according to Lilljeborg), Malm, and Collett have found fully developed Sea Lampreys about 14* 1 * 2 cm.
long. The larvae {Ammocoetes unicolor) attain, according to Dekay, a length of at least about 1 dm.

Scandinavian Fishes.

149

1184

SCANDINAVIAN FISHES.

elliptical disk. In its first-mentioned condition, when
the disk can almost be folded together from the sides,
its length is about equal to the depth of the body,
somewhat greater or less, and as a rule more than 4/5
of the length of the snout (to the anterior margin of
the eyes), which varies between 10a and 8 % (excep-
tionally approaching 7 %) of the length of the body
or between 2/s b and 3/ 4 of that of the head to the first
gill-opening.

In the mere size of this suctorial disk, even rela-
tively greater than in the following forms, we have a

est

median line), directed in a more or less sharp crook
towards the mouth; but exactly in the preoral median
line the row is straight, and the rows on the posterior
part of the suctorial disk are only slightly curved, each
row being directed inwards with fair regularity towards
one of the teeth (cusps) of the posterior dental plate
of the mouth. The teeth in these rows (the suctorial
teeth, tig. 352, est and ist ) are unicuspidate, conical,
gradually increasing in size inwards, separated by
grooves in the gums, which are thus divided into lo-
zenges, one to each tooth. Four rows meet the sides

Fig. 352. Oral disk of a Lamprey ( Petromyzon marinus) 8Y2 cm. long, X 2.
aln, anterior linguodental plate; alt, anterior lateral dental plate; at, anterior dental plate; est, outer suctorial teeth; ist, inner suctorial teeth ;
mil, mediolateral dental plate; pin, posterior linguodental plate; pit, posterior lateral dental plate; pt, posterior dental plate.

very good character for the Sea Lamprey; but the most
characteristic peculiarities are shown by the dentition
of the mouth and tongue. The suctorial disk is edged
with a double or multifarious series of densely set
fringes (dermal papillie), and within these appear nu-
merous rows of teeth (12 or 11 on each side of the

of the mouth, and the two innermost (largest) teeth in
these rows (the lateral teeth, alt, mlt, and pit) are so
closely set that they are joined at the base into bicuspid
teeth, each on its own root (lateral dental plate). The
bicuspid anterior dental plate of the mouth (at), situated
between but a little behind the anterior lateral plates,

“ Sometimes 11 in the variety occurring in Cayuga Lake (at Ithaca, N. Y.) ; see Meek in Jordan and Fordice, 1. c.
b In exceptional cases only 63 see Meek, 1. c.

SEA LAMPREY.

1185

is of about the same size as these and of the same
form. The crescent-shaped posterior dental plate of the
mouth (pt), ou the other hand, is considerably larger.
In young specimens it is comparatively higher than in
old, but also has comparatively smaller and usually
more numerous (in many cases 8") and more pointed
teeth. In old specimens the latter (usually 7) are more
conical, of about the same size and form as the large
lateral teeth. All these teeth probably serve merely as
an adhesive organ that holds the Lamprey fast to the
skin of its prey, which is forced to drag it along.
Within the mouth is the rasping lingual apparatus that
opens the skin of the victim and saws into its flesh.
The anterior linguodental plate (lingual plate, (tin) is
indeed simple and curved semilunarly like the posterior
dental plate of the mouth; but at the middle it is deeply
recurved, thus acquiring the form of a 8. Of the two
loops thus produced the left bears 7 or 6, the right 6
or 5 sharp points (tooth-cusps), exclusive of the hind-
most median tooth of the plate. Here too it seems to
be the rule that young Lampreys have more numerous
and more pointed teeth than old* 6. Behind (within) this
plate lie the posterior linguodental plates (supralingual
plates, pin) beside one another, each resembling in form
a reversed ruminant hoof, with the crook directed for-
wards and parallel to one of the loops of the anterior
linguodental plate. The left supralingual plate is usually
furnished with 13 pointed teeth, the right with 12,
these being largest in front, on the crook, gradually
smaller behind.

The eyes are round. Their size varies considerably
with age; but as they are covered by the more or less
transparent skin, the measurements taken depend greatly
on the transparency of the skin and the method of
measuring. In Sea Lampreys 4 — 8V3 dm. long Kroyer
found the diameter of the eyes to be 4/5 % of the length
of the body or about 7 % of the length of the head to
the first gill-opening. Our measurements of the outer
diameter of the iris in so large specimens fix the latter

percentage at about 10 or 9C. The opening of the
nasal duct (the nostril), more or less distinctly ele-
vated into a tube, is about 1/3 as large as the eyes,
and is situated just in front of the interorbital space,
at a distance from the tip of the snout measuring about
2/3 (63 — 70 %) of the said length of the head. Behind
the nasal duct lies the light, elliptical, but indis-
tinctly bounded epiphyseal spot.

The gill-openings are comparatively small (see our
figure in PI. LIII), but when distended they attain a
height not much less than the diameter of the eyes.
They are each covered from in front, partly by the
anterior margin itself, which forms a vertical dermal
flap, partly by two other, somewhat pateriform, rounded
dermal lobes, one of which rises from the inner part
of the lower wall of the gill -opening, the other hanging
down in a similar manner from above. The hind mar-
gin of each gill-opening is furnished with small fringes
and papilla?. The total extent occupied by the seven
gill-openings in a row one after another measures about
9 or 10 % of the length of the body, and in young
Sea Lampreys (2 — 3 dm. long) is in many cases equal
to the length of the snout to the nostril or about 2/s
of the length of the head to the first gill-openingd, in
the old greater, about 4/5 (77 — 84 °%) of the said length
of the head, which in the young is up to 15 %, in the
old only about 11%, of the length of the body.

The first dorsal fin is an elongated, low triangle,
with the greatest depth (75 — ’7 of the length) situated
in the anterior part, but with the anterior end indis-
tinctly delimited from the dorsal edge of the body,
which margin in ripe individuals (oftenest in males,
but sometimes too in females) may rise into a hard
carina both before the first dorsal fin and between it
and the second dorsal. This renders it also difficult
to determine with certainty the distance between the
first dorsal and the tip of the snout*, but the rule
appears to be that the fin commences close behind or
even at the middle of the body, comparatively further

a According to Meek (1. c.) exceptionally 9.

6 Blunter and sharper teeth thus occur in this species as in the following one. In estimating the importance of this difference, which
is by no means exclusively dependent on age, it should be remembered that we have here to deal not with real teeth, but only with points
of horny sheaths, which are changed time after time. In ordinary cases the horny sheath is so loosely attached to the plate that it can be
removed altogether, new, unworn, and therefore (if they be fully developed) more pointed teeth being thus exposed to view.

c Lilljeborg gives 9 or 8 % (*/,, — '/i 2)- According to Meek (1. c.) this percentage is about 16 — 13 in Sea Lampreys 21/2 — 3'/2
dm. long from Cayuga Lake.

d According to Meek (1. c.) sometimes only 55 % of the last-mentioned length.

e According to Meek this distance varies with great irregularity between 55 V2 and 49 % of the length of the body.

1186

SCANDINAVIAN FISHES.

back in the young0. The length of the tin is about
9V2 — 12 % of that of the body. The second dorsal is
considerably higher (sometimes twice as high or a little
more), but essentially of the same form, its length
varying between about 18 and 26 % of that of the
body, with the highest percentage in old specimens.
Behind it meets the caudal tin, or overlaps the upper
anterior margin thereof, but is always sharply defined,
as well as the first dorsal, at its posterior termination.
In front it is separated from the first dorsal by an in-
terval which in adult specimens (over 21/2 dm. long)
varies with age from about 6 to 4 % of the length of
the body, and which, according to Wilder’s measure-
ments of the North American fresh-water form, is less
in the males than in the females. The diphycercal
(almost symmetrical) caudal fin is uniformly widened
behind (tongue-shaped), with short, triangular but-
rounded tip. The entire length of the caudal fin oc-
cupies in adult Lampreys about 9V2 — 8 V3 % of the
length of the body.

The anal aperture lies below the anterior portion
of the second dorsal fin, at a distance from the tip of
the snout measuring 75 b — 72 % of the length of the
body, further back in the young than in the old. The
urogenital papilla is comparatively small. As men-
tioned above, all the Cyclostoines are without anal fin;
but between the vent and the caudal fin a dermal carina
frequently occurs, similar to that of the dorsal margin.

The coloration varies with age, locality, and the
time of year. Our figure represents a nearly ripe fe-
male in a fresh condition, as received by the Royal
Museum from Ringkjobing (Jutland) at the end of May
through Mr. 0. Fredericksen. Here only the white
and blue colours prevailed, the former being pure on
the belly, the latter pale but unmixed on the head, and
composing the pale ground-colour of the sides and back,
which were mottled with a darker blue, shading into black.
The fins were pale, mottled with blue, with a dash of gray.
The pupil was black, the iris of a faint silvery lustre
but not much paler than the prevailing colour. The
gill-openings and mouth were tinged with red. Dono-
van" figures and describes the species as more varie-
gated, with a ground-colour of lustrous green, mingled

with blue, yellow, and red, with a handsome marbling
of black. He also adduces blue and olive varieties, as
well as reddish ones mottled with chestnut-brown in-
stead of black. According to Kroyer the under surface
of the head is yellow, the fins being reddish, of a dark
orange, the dorsal fins besides mottled with black. Ac-
cording to both Malm and Collett the marbling is
absent in young specimens; and according to Lesueur
and Dekay this also applies to the larva?.

The larval stage, so far as is known, has not yet
been observed in Europe; but in North America an
Ammocoetes bicolor was described by Lesueur (1818)
and another form, Amur, unicolor , by Dekay (1842), both
of which have been referred by Jordan and Fordice to
Petromyzon marinus as its larva?. The former of these,
according to Dekay, has not been rediscovered since
Lesueur’s time, but had been named from its colora-
tion, reddish on the back and sides, white on the belly,
with a sharp, undulating limit between these two co-
lours. The latter ( unicolor ) is in all respects still more
like an earthworm, but has developed distinct eyes. Its
coloration is described by Dekay as a nearly uniform
dark or dusky brown, occasionally verging to bluish,
somewhat lighter beneath; the anterior part of the head
darkest. The form of the body cylindrical for two
thirds of its length from the head, becoming slightly
'compressed just anterior to the vent, very much com-
pressed and acuminated at the tail. Greatest depth of
the body 71/'2 % of its length. Surface smooth, with
between 80 and 90 transverse folds (myocommata),
giving the body an annulated appearance. The fins ap-
pear to be nothing more than mere membranous pro-
longations of the skin, without the slightest vestige of
rays. The dorsal commences at the beginning of the
3rd fifth of the length of the body (from the snout), in
a scarcely perceptible furrow, and advances low and
subequally until about the middle of its length, when
it begins to rise, and then rapidly diminishes at the tip
of the tail, where it unites with the caudal and anal,
which latter is obsoletely triangular, and becomes in-
sensibly effaced at a point corresponding to the begin-
ning of the last fourth of the length of the body. The
vent is a large longitudinal aperture between the 2nd

a Our measurements of Sea Lampreys 2 1/2 — 8 1 / dm. long show that the distance from the tip of the snout to the beginning of the
first dorsal fin diminishes with increasing age from 53 to 48 % of the length of the body. Kroyer’s measurements of specimens 4 — 8 f'.j
dm. long indicate that these percentages decrease from 51 'A to 50.
b According to Meek (1. c.) exceptionally 77 or even 80.
c Brit . Fish., pi. LXXXI.

SEA LAMPREY.

1187

and 3rd third of the length of the body. Mouth quad-
rilateral. Opening to the throat very large, but accu-
rately closed by six irregular and ragged subcartilagi-
nous processes, which meet in the centre. Anterior lip
transverse, convex on its outline, and emarginate at each
end, where it unites with the lateral lips; these latter
are wide and convex on their slender margins, uniting
posteriorly below the edge of the lower lip, leaving them
free above. Length of the snout to the eyes 5 %, and
the diameter of the eyes lV4 %, of the length of the body.
Eyes distinct, lateral, covered with the common tegu-
ments, and placed in a depression over the margin of
the lower lip. Nasal orifice large, and surrounded by a
raised margin; contracted, linear in front, circular be-
hind; its distance from the tip of the snout 33/4 % of
the length of the body. The branchial furrow occupies
7 8 of the length of the body, is directed obliquely down-
wards, and extends to a point above the lower angle of
the lateral lips; the apertures are exceedingly small, and
appear to be capable of being closed by their mem-
branous edges. Such was Dekay’s description of this
larva. It is said to be common in the muddy bottom
of most streams in the north and west of New York
State. Jordan and Gilbert explained it as a larva of
Dekay’s Petromyzon appendix; and when Jordan and
Fordice combined this nominal species with Petromyzon
marinus, the larva, of course, had to be referred to the
latter species. If the explanation is correct, the Sea
Lamprey thus undergoes a metamorphosis similar to
that of the following species, though not- in every de-
tail. The mouth apparatus and the snout are entirely
re-formed (see above). The eyes advance further for-
ward from the gill-openings. The nostril loses its an-
terior narrow part. The anal aperture is removed fur-
ther back. The dorsal fin-growth is divided, and its
beginning moved more to the front. A special caudal
fin is developed; but the rudimentary anal fin disappears.
Ammoccetes bicolor, which has also been referred to the
present species, was said to resemble the developed Sea
Lamprey more nearly in the dorsal fins and the position
of the anus, though the eyes were still indistinct.

The Sea Lamprey, as we have seen, has an exten-
sive geographical range, embracing all the seas of Europe.
According to Pallas the same species inhabits the Cas-
pian Sea and ascends the Volga; but Kessler ranged
this form as a new species, Petromyzon Wagneri. It

would also appear uncertain whether we can still rely
on Pallas’s statement that our Sea Lamprey also occurs
in the Sea of Okhotsk. From “West Africa”, however,
according to Gunther, the British Museum has received
the true Petromyzon marinas; and this Lamprey has
perhaps a more congenial habitat in the west of the
Atlantic and the east of North America than in Europe.
There it has even been landlocked in several lakes, and
lives in these without ever finding its way to the sea.
In this species we accordingly find the same state of
things as among the Salmons, and the form landlocked
in Cayuga Lake (Ithaca, N. Y.) was regarded by Wil-
der as a distinct species, Petromyzon dorsatus, its dor-
sal carina being particularly well developed.

In Scandinavia the Sea Lamprey is found every-
where from the extreme north, though of unfrequent
occurrence. From the Vestmannaeyiar, on the south
coast of Iceland, it was remarked by Krdyer, from the
Faroe Islands by the elder Reinhardt. In Greenland it
is unknown. In Norway, as well as in Bohusl&n and
on the other coasts of Sweden, probably too in Denmark,
many years may elapse without the catch of a single
specimen being notified to the museums, and as a rule
only solitary individuals are taken. In the Baltic, where
it is so well knotvn on the coast of Blekinge, according
to Nilsson, that it has a special name ( Sillapipare )
among the fishermen, the species penetrates so far that,
according to Malmgren, “a fairly large and handsome
specimen has been taken in the seine in Gammelstads-
vik” (Southern Finland).

All that is known of the life led by the Sea Lam-
prey in salt water is that it swims like the Eel, bending
the whole body in serpentine movements. Frequently
it is taken from shore in the Herring-seine; and this
would in all probability not happen so often, unless the
Lamprey were moving about among the Herrings. Most
of its time, however, it no doubt passes attached to
stones or floating objects, such as driftwood, or boats,
and to the fishes which it selects as its victims. Gun-
nerus (1. c.) states that Sea Lampreys fix themselves in
great numbers to the body of the Basking Shark and
do not leave it until it is dead". The Sea Lamprey also
attacks finer fish, such as Mackerel, Codfish, etc. Gesner
relates6, on the authority of a Strasburg fisherman, that
the Sea Lamprey attaches itself to Salmon ascending
from salt water, and is thus conveyed far up the rivers.

“ The same statement, given on the authority of Governor Christie of Bergen, appears in Kroyer, 1. e., p. 1038.
b De Aquatilibus, p. 596.

1188

SCANDINAVIAN FISHES.

But the Sea Lamprey also roves into fresh water on its
own account, when it makes its way, early in the year
or at latest in early summer (Feb. — June), to the spawn-
ing, which is performed during summer. On these ex-
cursions, however, the Sea Lamprey proves to be no
very powerful swimmer, capable of stemming the cur-
rent with ease; “but, when the stream is so strong that
the Lamprey lias difficulty in surmounting it, the fish
plunges quickly forwards and hastily attaches itself to
some fixed object, then awaiting an opportunity for a
new plunge” (Yarrell and Iaroyer). Consequently it
does not penetrate far up the stream in rapid rivers;
but in the Gotha Elf it goes at least to Lilia Edet
(Lloyd® and Malm* 6 * *); in the Kjeflinge River (Scania) it
ascends at least a couple of leagues from the sea (Nils-
son); and in the River Helge it was taken by Schager-
strom at Kristianstad (Lilljeborg). In Germany it is
found still farther inland, in the Rhine to Basel, in the
Elbe up to Bohemia (Benecke). In the Loire it has
been met with above Orleans, in the Rhone and Isere
beyond the frontier of Savoy (Blanchard).

The spawning of the Sea Lamprey was observed
by Panizzac at Pavia on the River Po; and Bartlett
wrote to Stored (1. c.) of its breeding operations in
Massachusetts, “They ascend the rivers a little earlier
than the shad, and move mostly in the night. It is not
known by the fishermen when they return, as they are
never seen. There is a notion that they all die. They
are often seen in the summer in pairs at work together,
constructing a little mound of stones. They build this
about three feet in diameter at the base, and about two
feet high, of stones from the size of an ounce bullet to
that of the fish. They often aid each other in carrying

the same stone. This is pretty evidently a labor of love , as
they copulate once in five minutes, or so, during the whole
time. The young go down the river when the water be-
gins to freeze. They are from six to eight inches long”.
But to take the above words of Bartlett to imply that the
metamorphosis from the Ammoccetes stage takes place with-
in the first year, would, no doubt, be too hasty. Strange
to say, no observations are recorded in Europe either of
the development of the Sea Lamprey in its earliest stages.

The Sea Lamprey is a palatable dish, if skilfully
prepared, when caught in spring and before it is ripe
for spawning. In former times it was offered as a
Christmas gift by the city of Gloucester to the sovereign
of the realm. It may be stewed like Burbot, or mari-
naded for preserving. Day gives the advice, however,
to remove the notochord before boiling, as being too
indigestible. Many consider the tvhole fish uneatable.
In many places it is related that the Sea Lamprey is
far rarer now than it was formerly. The once famous
fishery of the Severn has decayed so greatly that, ac-
cording to Day, the usual price of a Sea Lamprey there
is now half-a-crown. In France the Sea Lamprey was
protected, together with the Salmon and Shad, in the
piscatorial laws of ancient times, by fishing prohibi-
tions under certain circumstances, and the trade was
strictly regulated. At the present day, says Blan-
chard, it is, if not exactly rare, so uncommon that
no great attention is paid to it. In Scandinavia this
fishery has never possessed any importance.

All the Lampreys are tenacious of life, and this
adapts them eminently for use as bait. The Sea Lam-
prey, however, can seldom serve the Scandinavian fisher-
man for this purpose.

THE LAMPERN OR RIVER LAMPREY (sw. nattingen).

PETROMYZON FLU VI AT JUS.

Plate LIII, figs. 2—4.

Anterior linguodental plate evenly and slightly curved, with the convexity in front. Anterior dental plate of the
mouth semilunarly curved and so long {in the transverse direction of the body ) that the elevated, dentiform ends
are distinctly separated from one another11 . Length of the oral dish in adult specimens less than half that of the

head to the first gill-opening .

Syn. Lamproye <$ eau doulce + Lamproyon, Belon, Nat., Divers. | fluv., p. 202. Mustela + Alterum genus Lampredce, Gesn.,

Poiss., p. 67. Lampetra parva et fluviatilis, Rondel., Pise, j De Aquat., pp. 595 et 597. Lampetra fluviatilis, Willughb.,

a Scandinav. Advent., vol. I, p. 147.

6 Malm also cites a newspaper paragraph which seems to indicate that the Sea Lamprey sometimes passes the locks at Trollhattan into Lake Wener.

c Mem. Istit. Lomb. Sc., Art., Milano, vol. II (1845), p. 25.

d = Genus Lampetra , Gray, List Spec. Fish., Brit. Mus ., part. I, Chondropt., p. 140.

LAMPERN.

1189

Hist. Pise., p. 104, tab. G. 2, fig. 1 (4- Lamp, media,
p. 106, tab. G. 3, fig. 2 + Lamp, cceca Baltneri , p. 107,
lab. G. 3, fig. 1). Petromyzon unico ordine denticulorum
minimorum in limbo oris, prseter inferiores majores ■+• Petrorn.
corpore annuloso, appendicibus utrinque duabus in margine
oris, Art., Ichtliyol., Gen. Pise., p. 64; Syu. Pise., pp.
89 et 90; Spec. Pise., p. 99. Petromyzon pinna dorsali
secunda angulata (Stiecis Neinoga, Westrobothniensibus Natting)
+ Petr, pinna dorsali seennda lineari, labio oris snperiore
pone lobato, Lin., Fna Suec., ed. I, p. 102. Lesser Lam-
prey, Penn., Brit. Zool. (ed. 1776), vol. Ill, p. 70, tab.
VIII, No. 28 + Pride, p. 71, tab. VIII, No. 29.

A : forma major, pinnis dorsalibus minoribns, remotis.

Petromyzon fluviatilis , Lin., Syst. Nat., ed. X, tom. I, p.
230; Bl., Fisch. Deutschl., part. Ill, p. 41, tab. LXXVIII,
fig. 1; Retz., Fna Suec. Lin., p. 303; Swartz, Svensk
Zoologi, H. 6, No. 33; Pall., Zoogr. Boss. Asiat., tom. Ill,
p. 66; Cuv., Regn. Anim., ed. I, tom. II, p. 118; Nilss.,
Prodr. Ichtliyol. Scand., p. 122; Ekstr., Vet. Akad. Hand].
1834, p. 70; Yarr., Brit. Fish., ed. 1, vol. II, p. 454;
Kb., Damn. Fisk., vol. Ill, p. 1042; Nilss., Slcand. Fna,
Fisk., p. 745; Hckl, Kn., Siisswasserf. Ostr. Mon., p.
377; Sieb., Siisswasserf. Mitteleur., p. 372; Malm, Gbgs
Vet., Vitt. Samh. Handl., Ny Tidsf.. VIII, p. 88; Mgrn,
Finl. Fiskfna (disp. Helsingf. 1863), p. 72; v. Bemm. in
Herklots, Bouwst. Fna Nederl., part. Ill, p. 392; Blanch.,
Poiss. d. eaux donees Fr., p. 515; Canestr., Arch. Zool.,
Auat., Fisiol., vol. IV, p. 185; Fna Ital., Peso., p. 31;
Gthr, Cat. Brit. Mus., Fish., vol. VIII, p. 502; Coll.,
Forh. Vid. Selsk. Chrnia 1874, Tillfeggsb., p. 219; 1 879,
No. 1, p. 107; N. Mag. Naturv. Chrnia, Bd 29 (1884),
p. 122; Ltkn, Arct. Man. (Rupert Jones, 1875), p. 122;
Malm, Gbgs, Boh. Fna, p. 632; Fedders., Naturh. Tidskr.
Kbhvn, ser. 3, vol. XII, p. 94; Mor., Hist. Nat. Poiss. Fr.,
tom. Ill, p. 604; Bncke, Fisch., Fischer., Fiscliz. O., JV.
Preuss., p. 196; Mela, Vert. Fenn., p 370, tab. X; Day,
Fish. Gt. Brit., Irel, vol. II. p. 359, tab. CLXXIX, fig. 1 ;
M6b., Hcke, Fisch. Osts., p. 161; Fatjo, Fne Vert. Suisse ,
vol. V, part. II, p. 512; Lillj., Sv., Norg. Fna, Fisk.,
vol. Ill, p. 693; L6nnb., Bib. Vet. Akad. Handl., Bd 18,
Afd. IV, No. 2.

Petromyzon argenteus , Bl., Ausl. Fisch., part. IX, p. 74, tab.
CCCOXV, fig. 2; Bl., Schn., Syst. Ichtliyol., p. 532, tab.
102, fig. 1; Couch, Fish. Brit. Isl., vol. IV, p. 400.

Petromyzon pricka (ex Daubenton, Encycl. Meth.), Lacep., Hist.
Nat. Poiss., tom. I, p. 18.

Petromyzon Omalii, V. Ben., Bull. Acad. Sc. Belg., ser. 2, tom.
II (1857), p. 549 cum tab. ad p. 554; tom. XX (1865),
p. 46; Malm, 11. cc.

? Petromyzon plumbeus , Ayres, Proc. Calif. Acad. Nat. Sc. 1855,
p. 27. Petr, ayresii, Gthr, 1. c., p. 505. Ammoccetes
cibarius, Jord., Ford., Ann. N. Y. Acad. Sc., vol. Ill (1885),
p. 292.

B: forma minor, pinnis dorsalibus altioribus, approximatis.

Petromyzon Planeri, Bl., Fisch. Deutschl., part. Ill, p. 47,
tab. LXXVIII, fig. 3 (dub.); Lacep., 1. c., p. 30 (+ Petr,
sanguisuga, tom. II, p. 101, tab. I, fig. 3 + Petr, sept-
oeil + Petr, niger, tom. IV, p. 667, tab. 15); Osb., Vet.
Akad. Handl. 1804, p. 181; Swartz, 1. c.; Cuv., 1. c., p.
119; Nilss., Prodr., 1. c. ; Yarr., 1. c., p. 457; Kr., 1. c.,

p. 1052; Nilss., Fna, 1. c., p. 747; Kessl., Bull. Soc.
Natur. Moscou, 1856, p. 390; A. Mull., Miill. Arcliiv
1856, p. 325; Hckl, Kn., 1. c., p. 380; Sieb., 1. c., p.
375; Malm, Gbgs Handl., 1. c., p. 92; Mgrn, 1. c., p.
73; Blanch., 1. c., p. 517; Canestr., Arch., 1. c., p. 186;
Fna, 1. c.; Langerh., Ber. Verh. Naturf. Ges. Freib. i. B.,
Bd VI (1873), H. 3; Fedders., 1. c., p. 95; Mor., 1. c.,
I>. 606; Bncke, 1. c., p. 197; Fatio, 1. c., p. 499; Lillj.,
1. c., p. 708.

Petromyzon plumbeus (= P. septoeil, Lacep.) + Petromyzon
bicolor (= P. niger, Lacep.), Siiaw, Gen. Zool., vol. V, p.
263.

Petromyzon branchialis ( + larva), Gthr, 1. c., p. 504 ; Coll.,
Forh. 1. c. 1874, Tillasgsh., p. 220; ibid., 1879, p. 107;
Malm, Fna, p. 636; Mela, 1. c., p. 371, tab. X; Day,
1. c., p. 362, tab. CLXXIX, figs. 2 (larva) et 3; Jord.,
Ford. ( Ammoccetes ), 1. c., p. 293.

C: forma larvalis.

Petromyzon branchialis, Lin., Syst., 1. c. ; Hi.., Fisch. Deutschl.,
1. c., p. 45, tab. LXXVIII, fig. 2; Swartz, 1. c.; Dum.
(Ammocoetus), Dissert. Poiss. Cyclost. 1808; Cuv. ( Ammo -
coetes ), 1. c., p. 120; Nilss. ( Petromyzon , subg. Ammo-
ccetes), Proclr., p. 123; Yarr. ( Ammoccetes ), 1. c., p. 459;
Kr., 1. c., p. 1060; Nilss., Fna, 1. c., p. 748; Hckl, Kn.,
1. c., p. 382; Cederstr., Ofvers. Vet. Akad. Forh. 1861,
p. 91.

Petromyzon ruber, Lacep., 1. c , tom. II, p. 100, tab. I, fig. 2.

Petromyzon Unnbricalis, Pall., 1. c., p. 69.

Acln. Ammocoetes aureus (Bean, Proc. U. S. Nat. Mus. 1881,
p. 159; Br. -Goode, Fish., Fish. Industr. U. S., Sect. I,
tab. 251), ex Alaska, Petrorn. fluviatilis simillimus videtur.
Caput (fide descriptionis) perparvum, !lVmon longitudinis cor-
poris 1. 187/ln00 distantife inter apicem rostri et originem pinme
dorsalis primse, quod numquam in nostra specie vidi. Figura
tamen citata has rationes non confirmat.

The Lampern attains in Scandinavia a length of
at least 4x/4 din. In Germany and France it is said
sometimes to measure 5 dm. In this respect, however,
there is great difference between the Larnperns of dif-
ferent tracts within the same country, this species too,
in its full development, being an anadromous fish, and
requiring the bounteous resources and saltness of the
sea, or at least of brackish water, to attain its most
typical size and form. How variable the anadromous
fishes are in this respect, we have seen most distinctly
in the Salmons; but the Sea Lamprey here affords a
comparison at closer quarters. Where the Lampern
lives in brooks and other small collections of fresh
water, it usually appears as a smaller form, about 2
dm. long. The same differences may be observed in
the size of the larvae. Sometimes the metamorphosis
from Ammoccetes to Petromyzon takes place at a length
of 1 din., sometimes larvae are met with at least 1 ' 2

1190

SCANDINAVIAN FISHES.

dm.“ long. What are the causes that retard the meta-
morphosis or accelerate the growth of the larvae, we
do not know; and sometimes we find in the same Ava-
ters Prides which are larger than the Lamperns*.

The size, the teeth, the closer proximity of the
fins, and the usually immaculate colour distinguish the
Lampern from the preceding species; “but in all other
relations of form, both external and internal”, Avrote
Lacepede, “the Iavo Lamprey species are so like one
another that they seem to be two copies of the same
model”. The Eel-like body is here too more or less
terete in front, laterally compressed behind, Avith the
same variability as in the Sea Lamprey, though some-
Avhat greater0.

O

The length of the head to the first gill-opening
varies in the true Lamperns between about 14 and a
little over 10 %d of the length of the body, the per-
centage being least in the oldest specimens; but here
too it is as a rule greater than the length of the series
of gill-openings0. In the Prides, on the other hand,
the length of the head is only about 6 7 2 — 7 1/2 % of
that of the body and ahvays less than that of the
branchial region, which occupies about 12- — 1 2* 1/2 % of
the last-mentioned length. The length of the snout to
the anterior margin of the eyes is in the Lamperns
about V3 (60 — 68-r %), in the Prides only 1/2 , of the
length of the head. These differences are external ex-
pressions of the alterations involved by the metamor-
phosis; but hereAvith too is connected the great differ-
ence in the structure and form of the mouth and oral
disk. The Prides (PL LIII, figs. 3 and 4) are toothless
and never attach themselves by suction. They lead a
more worm-like life, buried in sand or mud. Their
mouth is edged in front and on the sides Avith a horse -
shoe-shaped or semiquadratic anterior lip (fig. 351,
La), the lateral parts of Avhich are rounded and pen-
dent, passing evenly or with a shalloAV sinus into the
corners of the anterior side, but sharply terminated be-
hind, Avhere they enclose betAveen them the shallower,

transversal posterior lip ( Lp ). The outer mouth cavity
is forniciform, Avith a patch of small papilke, set in rows,
at the middle of the roof, and is bounded behind by a
coroniform series of concentrically directed processes Avith
verrucose ramifications (papillae, MT). Behind this co-
rona lies an anterior pharyngeal cavity (inner buccal
cavity M), of somewhat smaller capacity than the other
and bounded behind by an annular, contractile valve
(velum, vet), Avhose lateral parts are tumid; and behind
this again lies the still smaller true pharynx, the en-
trance to the tubiform branchial cavity, through which
the food too must pass. In the anterior pharyngeal
cavity the lingual apparatus is developed. The corona
of papilla? disappears. The lips are thickened, but
coalesce into a ring, Avhich becomes the suctorial disk
of the Lampern (PI. LIII, tig. 2). The most important
differences betAveen the suctorial disks of this species and
the preceding one have been already noticed. Here the
disk is both smaller — the dimensions of the snout being
thus also reduced — and more feebly armed. The edge
is fringed Avith papillae here too, but sometimes only in
a single toav. Just Avithin the margin we see a ring of
small horny teeth ( est ), like the other teeth pointed or
blunt, Avhite or yellow*7. Within the anterior part of
this ring and parallel to it, Ave find in ordinary cases
a curved row of similar, simple teeth ( 1st ), and the rule
is that the remaining surface of the suctorial disk, out-
side the composite teeth of the mouth, is smooth or very
finely verruculose. This is one of the most patent char-
acters of the species, in contradistinction to the lozenged
comparting of the suctorial disk in the Sea Lamprey,
Avhere the gums round the bases of the several suctorial
teeth are bounded by grooves intersecting in a diamond
pattern (see fig. 352). Frequently, hoAvever, Ave find
here too, in the Lampern, scattered suctorial teeth on
the anterior part of the disk, and in a pair (c? and 9)
of Lamperns from Archangel (fig. 353), Avhich are other-
Avise quite typical specimens — only that the middle pair
of the lateral teeth of the mouth, a peculiarity Avhich is

a 2 dm., according to Benecke.

6 At Helgevarma Mill, near WexiS, Baron G-. C. CederstrSm caught, in May, 1857 and 1860, Prides 145 mm. long and Lamperns
(forma Planeri ) 120 mm. in length.

0 The greatest depth of the body varies in the Lamperns from about 7^2 to nearly 9 % of the length thereof. In the small Prides,
which as a rule, however, have a more tumid branchial region, the depth behind this is sometimes only 6'/3 % of the length of the body.

d 10'3 is the least percentage we have found for this relation.

e An exception is adduced by Kroyer in the largest Lampern measured by him. The variations may be due, however, to the dif-
ferent degree of contraction in the snout and dorsal muscles at death or in alcohol.

f In the preceding species 68 — 75 %.

S' The dental differences in these respects are so variable that we have not been able to reduce them to any rule ; but it would appear that speci-
mens from the sea or with only intumescent genital organs and not yet shrunken intestinal canal most often have the most pointed and whitest teeth.

LAMPERN.

1191

not uncommon, are bicuspid instead of tricuspid — there
not only appear anterior suctorial teeth (ist, above in
the figure), in almost the same arrangement as those of
the Sea Lamprey and with lozenged frames, but the
male also possesses, on the posterior part of the disk,
a curved though irregular row of teeth (ist, below in
the figure). The specific character in the anterior dental
plate of the mouth (at) we have pointed out above. The
posterior dental plate (pt) is of the same form as in the
preceding species; but its two outermost cusps are larg-
er than the others and oftenest bifid (bicuspid). The
number of cusps between them is as a rule five, excep-
tionally (fig. 353) four, so that the total number of cusps
varies between 7 and 9. The lateral dental plates of
the mouth (alt, mlt , and pit) are similar to those of the
preceding species, but lie farther apart, and the middle
pair on each side (mlt) are, as we have mentioned,
usually tricuspid. The anterior linguodental plate ( aln )
is crescent-shaped, with the median cusp on the uni-
formly curved margin largest, and with 5 or 6 small
cusps on the lateral parts. The posterior linguodental
plates lie, when at rest, deep in a groove between the
tumid lateral halves of the tongue", and are only slightly
curved but otherwise of the same form as in the pre-
ceding species. Their 10 or 12 retral teeth are very small.

The orifice of the nasal duct undergoes almost the
same alterations as in the preceding species. In the
Pride it is pear-shaped, anteriorly pointed, with the
dermal margin more elevated and expanded behind. In
the Lampern it is more uniformly round, with the mar-
gin more or less elevated. In consequence of the some-
what shorter snout, the distance between this orifice and
the tip of the former is also less than in the Sea Lam-
prey. In the Pride this distance is rather less than 2/s
(about 37 7g %)■> in the Lampern rather less than 3/5
(about 53 — 59 %h ), of the length of the head to the first
gill-opening. The series of gill-openings is somewhat

longer than in the preceding species, varying, however,
between about 9 1,/2 and 11 Vs % of the length of the
body; and the several gill-openings may be closed from
the anterior margin by a continuous, even flap of skin.
In the Lampern too their posterior margin is verruculose.

In the form and position of the fins we can indeed
distinguish between two varieties of Lampern, but only
with far from reliable characters; and the variability in
these relations is such that we cannot even state a per-
fectly trustworthy distinction between the Lampern and
the Sea Lamprey. The rule, however, is that the limit
between the caudal fin and the second dorsal is less
distinct in the Lampern; and as the two dorsal fins are
sometimes contiguous, and a fin-like, though rayless,

est

Fig. 353. Oral disk of a male Lampern ( Petrornyzon fluviatilis , var.),
3 1 1/2 cm. long from Archangel, X 2. Signification of the letters
explained in the preceding figure.

dermal ridge is usually perceptible between the caudal
fin and the anus, the Lampern may be regarded as a
lower form, with more distinct vestiges of the larval
condition. As regards the difference between the two
assumed varieties of Lampern, it has been asserted, first
and foremost, that the exclusively fresh-water form
Petrornyzon brancliialis or Planeri in general has higher
and closely contiguous dorsal fins. According to K ro-
ver the following relations hold good:

o o

Petrornyzon

Petrornyzon

Planeri.

fluviatilis .

Length of the body expressed in millimetres - -

144—261

333—392

Greatest height of the first dorsal fin in % of the total length of the head and branchial region

11.1—14.8

o>

1

00

to

,, ,, „ ,. ,. ,, ,, ,, „ „ „ distance between the first dorsal and the tip of the snout

5.i — 6.6

3.2 — 3.3

,, ., ,, second „ „ „ „ „ total length of the head and branchial region

20—32.4

17.6—18.2

„ ,. ,. ,, „ ,, „ „ ,, distance between the first dorsal and the tip of the snout

9.2—14.6

6.9 — 7.7

a Our figure (PI. LIII, fig. 2) of the oral disk, as it appeared through the glass wall of an aquarium, where the Lampern had at-
tached itself, shows in the middle these lateral halves, separated by a V-shaped groove under the anterior dental plate of the mouth. Under
them the anterior linguodental plate, which is also white, may be seen; hut the dental cusps are too minute to be visible in the figure.
h In the preceding species about 63 — 68 %.

Scandinavian Fishes.

150

1192

SCANDINAVIAN FISHES.

However patent these differences may appear, inter-
mediate forms give rise to very great uncertainty. As
examples we will here adduce the corresponding measure-

ments in four Lamperns of the form usually called Petro-
myzon Planer i, two males and two females 278 — 309 mm.
long, taken together in Lake Ifo (Scania):

Minimum.

Maximum.

Greatest height of the first dorsal fin in % of the total length of the head and branchial region

9.i

14.2

„ „ „ „ „ „ ., ,, ,, „ „ distance between the first dorsal and the tin of the snout

4.3

6.5

„ „ „ „ second „ ,, „ „ „ „ total length of the head and branchial region

16.8

21.7

„ „ „ „ ,, „ „ „ ,, „ „ distance between the first dorsal and the tip of the snout

7.8

9.9

and in four females of the form Petromyzon fluviatilis, 255 — 288 mm. long, taken together at Lulea:

Minimum.

Maximum.

Greatest height of the first dorsal fin in % of the total length of the head and branchial region

7.4

9.9

„ „ „ „ „ ,, ,, ,, ,, ,, ,, distance between the first dorsal and the tip of the snout

2.8

3.7

„ „ ,, „ second „ „ ,, „ „ „ total length of the head and branchial region...

14.4

18.7

„ „ „ „ ,, „ „ ,, „ ,, distance between the first dorsal and the tip of the snout

5.6

7.7

As may be easily seen, these percentages partly
overlap one another, partly approach one another in
very close proximity.

Equally variable is the character drawn from the
relative position of the two dorsal fins. The length of
the space between them varies in Petromyzon fluviatilis,
according to Lonnberg’s measurements of 142 speci-
mens 245 — 375 mm. long, between 1* *8 and 7'8 % of
the length of the body; and on examining the results
of these measurements we find that in the 12 smallest
(245 — 255 mm. long) the average percentage was 4*5,
in the next 65 specimens (260 — 295 mm. long) 4*3,
in the next 52 (300 — 335 mm. long) 4*1, and in the
13 largest specimens (340 — 375 mm. long) 4*7. During
the period of growth examined by Lonnberg it thus
appeared that at first Petromyzon fluviatilis approached
nearer and nearer to the character of Petr. Planeri,
but eventually receded all the more abruptly therefrom.
Often, however, we find a dermal flap extending for-
ward from the beginning of the second dorsal fin in
the median dorsal line towards the first dorsal, so that
the limit of the former fin is difficult to fix. And
Schneider"1, WaygeiA, and BeneckfA have totally re-
jected the specific distinction between the two forms.

The relation between them, as Lonnberg has remarked,
is very like that between the large Trout ( grdlax )
and the Salmon ( blanklax ); and in the preceding pages
we have made the same observation in many other
parts of the history of the Scandinavian fishes. Fresh-
water life has had greater influence on Petromyzon
fluviatilis than on Petr, marinus; and two varieties have
thus arisen, which in their typical forms are indeed
easy of distinction, with the guidance of the above-
mentioned characters, but are linked together by inter-
mediate forms. “Among 7 specimens of Petromyzon
Planeri from the same locality (Bieberbach, near Gies-
sen) 3 had the two dorsal fins separated by an interval,
while in 4 the fins were contiguous” (Schneider).

The first dorsal fin begins in the Pride and usually
too in Petromyzon Planeri somewhat in front of, in the
typical Petromyzon fluviatilis somewhat behind, the
middle of the body. The second dorsal begins in the
former two in front off, in the last-mentioned form
behind, the beginning of the posterior third of the body.
The caudal fin is of the same symmetrical form as in
the preceding species. The boundary between it and
the second dorsal, marked by the more distinct rays of
the latter fin, is more prominent in Petromyzon fluvia-

a Beitr. Vergl. Anat., Entwickl. Wirbellh., p. 35, § 1.

* Verb. K. K. Zool.-Bot. Ges. Wien, XXXIII (1884), p. 311.
c Handb. Fiscliz., Fischer. (Max v. d. Borne), p. 193.
d Exceptions not seldom occur in old specimens of Petromyzon Planeri.

LAMPERN.

1193

tills than in Petr. Planeri. Its length varies here too
between about 10 and 8 % of that of the body. The
anal aperture also occupies the same position as in t he
Sea Lamprey but the urogenital papilla is generally
more developed, especially in Petromyson Planeri , and
longest and most pointed (projecting far back), as
usual, in the males. Besides tins sexual character may
be observed during the breeding season, especially in
the females, an hydropical tumidity of the skin (fig.
349), in particular round the anus and along the anal
fin-ridge, which is sometimes considerably elevated at
this season, as well as along the base and in front of
the second dorsal fin.

The coloration differs from that of the preceding
species in its monotony, being without sharply defined
spots, and as a rule on the back steel-blue or darker,
on the belly white, with a silvery lustre on the sides
of the body. Yet it varies with the age of the fish,
the season of the year, and the locality. In fresh water
and with age the back becomes greener, and the belly
assumes a muddier tone. The fins are either transparent,
grayish white, or, in old specimens, brownish. The
Prides are entirely destitute of silvery lustre and some-
times greenish yellow, sometimes bluish green. Our
figures of Prides, as well as of the Lampern, are painted
from living specimens taken by Professor Retzius at
Elfkarleby early in October.

The Lampern is dispersed throughout Europe,
Northern Asia, and Japan, as well as the northern re-
gions of North America together with Greenland. Theel
and Trybom brought home Prides of this species from
the Yenisei; Gunther referred the Petromyson japoni-
cus described by Martens" to the same species as our
Lampern; Petromyson pkmbeus ( cibarius ) and Petr, au-
reus from the west coast of North America have ap-
parently no greater claim, judging by the descriptions,
to a specific rank; and from Southern Greenland, ac-
cording to Lutken, the Museum of Copenhagen has re-
ceived two specimens of Petromyson fluviatilis. Through-
out Sweden this species occurs in both varieties, but
seems to be less common in the west than in the east,
besides which it has long been known that the more
typical form is commonest and largest in the Baltic and
the rivers of Northern Sweden. In Norway the last-

a Wiegtn. Arch. XXXIV, p. 3.

b Not adduced from Greece, however, by Apostolides.

c Ofvers. Vet. Akad. Fork. 1876, No. 3, p. 140.

d Nord. Aarsskr. Fisk. 1883, p. 307.

mentioned form, according to Collett, is very rare and
has been met with only in Christiania Fjord, but Petro-
myson Planeri is said to be common enough south of
the Dovrefjeld, where it attains, according to Rasch, a
length of at least 3 dm. North of the Dovrefjeld the
species has not been found; but Mela adduces it from
Varanger Fjord, and it has long been known from the
White Sea and the rivers of Northern Russia. In Den-
mark it has been met with both on the islands and in
Jutland. In the Sound it is no rarity. On Bornholm,
according to Feddersen, only the smaller form has been
found. In Russia, Germany, the Netherlands, Southern
Europe6, and the British Islands it is as common as in
Scandinavia, in some places perhaps commoner. Ac-
cording to Fatio it is unable to pass the great falls of
the Rhine and Rhone, being wanting both in Lake Con-
stance and the Lake of Geneva. In Scandinavia, on
the other hand, each of the forms occurs both in Lake
Wener, above Trollhattan, and in Lake Wetter. Nearest
the sources of the brooks and in small collections of
water the smaller form has its home; the larger keeps
to the lower courses of the rivers and, in the lakes,
to the outlets of the brooks. How high the Lampern
ascends in the mountain regions of Scandinavia, is
scarcely known as yet. Olsson0 says that both Lam-
perns and Prides are common in the southern part of
Stroms Vattudal (Jemtland), though they do not ascend
the streams flowing thither. According to Trybom'6

o

Prides 11 — 13 cm. long have been found at Asele
(Angerman Elf, 64° 10' N. lat.; 300 m. above the sea-
level), on flax placed in the river to steep; and similar
specimens are sometimes taken, it is stated, in Lake
Storlogcle (Logde Elf). Lilljeborg found Petromyson
Planeri at Karesuando (Muonio Elf, 68° 25' N. lat.; 320
m. above the sea-level). Fatio sets the limit to the
ascent of the Lampern in Switzerland at about 600 to
700 in. above the level of the sea.

Tenacious of life and greedy of prey, the Lampern
has the same habits as the preceding species. Its man-
ner of life in the sea is hardly known, for it is met
with too seldom in salt water. Baltic specimens have
been found to have the intestinal canal (stomach and
spiral intestine) filled to distension with a pulpy mass,
in which we could distinguish in one specimen morsels

1194

SCANDINAVIAN FISHES.

of bone and fin-rays, probably remains of some Goby.
From the island-belt of Lulea Widegren writes" that
during summer Lamperns follow the shoals of Baltic
Herrings and fasten themselves to the fish, gradually
devouring them. Collett received from Christiania
Fjord a Lampern that had attached itself to a bit of
Herring with which a long-line had been baited. From
Lough Neagh a Lampern was sent to Hyndman* 6 that
had fastened itself to a large Trout. Otherwise the
Lampern lives on minute creatures, such as worms,
crustaceans, and the larvee of insects. The Prides, which
are toothless and without closed sucking-disk, must
naturally confine themselves to the smallest prey and
the decomposing matter they may find in the mud or
sand. The Lamperns that have ascended the rivers
to spawn, with their greatly shrunken intestinal canal,
cease to take any nourishment.

The Lampern displays activity as it wriggles along
in the streams like an Eel or a snake, norv and then
taking a rest by attaching itself. At such times the
branchial region may be observed continually expand-
ing and contracting, opening and shutting the gill-
openings at a breath, about 200 times a minute when
the fish is most lively. Or it creeps up on jetties,
dams, or projecting stones, with head and branchial
region above the water, the air expressed from its gills
causing a faint bubbling sound. So tenacious is the
bite of the Lampern on these occasions that even if
the body be cut off the head sometimes retains its
position. The Lampern and its larva can live for hours
or days out of the water, and may thus be kept for
transmission or consumption as required; but in thunder-
storms it often happens that most of the captives die.

In autumn, from September till November, the
Lampern ascends the rivers from the sea. It is then
taken in great quantities, especially near the mouths of
the Norrland rivers. At this season the generative or-
gans are of fairly advanced development, with eggs
measuring 2/s mm. or less; but not until the spring or
early in the summer, when the ova have attained a
diameter of 1 mm., does the spawning take place, after
the same fashion as we have seen above in the case of
the Sea Lamprey. The Lamperns that constantly in-
habit fresh water do not repair to the spawning-places

a Landtbr. Akad. Handl. 18:de delen (1858), p. 200.

6 Thomps., Nat. Hist. Irel., vol. IV, p. 265.

Lillj., 1. c., p. 717.

d Cf. too Benecke, Fisch., Fischer ., Fisckz. 0., W. Preass., p

e This manuscript, which was rediscovered by Siebold in the 1

until shortly before the commencement of the opera-
tion, in Scandinavia usually during April or May. The
ova of the small Lamperns are then about equal in size
to those of their larger fellows. Hence the number of the
eggs is highly variable, from a thousand or so in small
individuals to several tens of thousands in large fish.

Aurell,: observed the behaviour of small Lamperns
at their spawning-places in the brooks of Lake Wetter.
“They occurred in very shallow water and where there
was a brisk current, sometimes 2 to 10 or 20 in com-
pany. When alone, they were timid and usually shot
off with almost lightning speed down the current if
disturbed. But this was not often the case, for they
sought one another with passionate eagerness. The
males were more slender and cylindrical than the fe-
males, which, seen from the sides, were broader. When
the males were touched, the milt spirted out through
the genital papilla to a distance of a yard or two, and
on being touched the female emitted her roe. Their co-
lour showed great variability, and some, both male and
female, were mottled.” Else, according to A. Muller d,
the male attaches himself to the neck of the female
and coils his body round her in a half spiral. The
spawning-place is chosen preferably on a pebbly bottom,
where the fish hollow out a cavity by wriggling the
body, or construct, as we have mentioned of the Sea
Lamprey, a shelter for the eggs. After the lapse of
about three weeks the larvae are excluded.

The development, first, to Pride and then to Lam-
pern, was known even to the Strasburg fisherman Bald-
ner, who in the 17th century wrote a Fischbucli e of the
Middle Rhine; but his remarks were forgotten until at-
tention was drawn to them by Siebold, after A. Muller
had succeeded (1856) in watching the spawning of Petro-
myzon Planeri in the Panke, a brook that falls into the
Spree -at Berlin, and in seeing Prides develop from the
deposited eggs. According to Muller the Pride lives
three or four years before its transformation into a
Lampern. It buries itself meanwhile in mud or sand,
or dwells in brooks or by the banks of rivers among
fallen and decayed leaves. At times the fish raises the
forepart of the body and swings it to and fro in the
water, or wriggles itself entirely free but hastens again
into concealment. It is so gregarious that in many

d. 38.

Museum of Strasburg, is also quoted in several passages by Willughby.

MYXINOIDS.

1195

places it may lie dug up in numbers, after the stream
has been dammed above the spot. Its Swedish name
of lindl (Flax Eel) is derived from its habit of creep-
ing among flax laid in the water to rot. In August
the metamorphosis begins, and by the New Year or
soon afterwards it is completed. The organs of genera-
tion become distinct even earlier than this.

After depositing its eggs the Lampern dies, ac-
cording to Muller; and Trybomq has remarked, as
evidence of this, that all the eggs in its ovaries are
of the same size, so that no undeveloped ova, to be
matured for another year’s spawning, can be detected.
Petromyzon Planeri, according to their assumption, only
lives a few months in its adult state. Of Petromyzon
fluviatilis, on the other hand, Benecke assumes that
the larvae migrate to the sea while undergoing meta-
morphosis; and their life in salt water is unknown.
But the fact is that sexually mature individuals of both
varieties are found differing so widely in size that it
is almost inconceivable that they can be of the same
age. It would thus appear that some attain maturity
the very first year after the metamorphosis, others not
until several years have elapsed. For the present, as

Fam. M Y ]

(Described by

Outer branchial apertures either single or several (6‘ —

Nasal duct open behint

This remarkable family in its essential characters
comes near to the preceding one, and lias therefore
been coupled therewith to form the Cyclostomous order;
but in other respects it exhibits such important dif-
ferences from the Pctromyzonidce that at present, the
links which would seem to have once connected them
having apparently disappeared, the investigation of their
mutual relationship and collocation in the phylogenetic
system is no easy task.

Perhaps they are both more or less retrogressive
forms; this is probable at least with respect to the
Myxinoids. In several respects, however, the latter
display obviously lower characters than the family of
the Lampreys.

The time may possibly come where the ontogenetic
evolution of the Myxinoids may in some degree supply

a Forh. Alim. Sv. Fisk. Konf. Goteborg 1891, p. 88.

6 According to Girard sometimes 14.

Fatio has also observed, we must consequently regard
it as an open question, whether most of the spent Lam-
perns may not survive, but conceal themselves some-
where beyond our ken, and possibly reappear at the
spawning of the following year.

The Lampern is a useful fish. For a good di-
gestion it is excellent eating. Smoked and broiled it
is a delicious whet before dinner; but for this purpose
it is lit only during the cold season; in summer it is
lean and dry. It is always a very valuable bait. It is
taken in Jcassar (closely woven baskets, preferably of
osier) and ndttingstockar (small square boxes made of
thin boards perforated with small holes), which are
placed between the stones where the current is of a
suitable strength. The fishery succeeds best on dark
nights, for the Lampern, like the Eel, is most active
between sunset and dawn.

The Lampern lias several Swedish names. Nejon-
bga is originally a German word ( Neunauge ). In Hal-
land it is known as stensugare (Stone-sucker). In Ble-
kinge it is said to share with the preceding species the
name of sillapipare. The smaller variety is called by
Swartz igel-neinogon (Leech-Lamprey).

I N 1 D JE.

USTAF RETZIUS.)

b) on each side of the forepart of the body ( the throat).

, penetrating the palate.

an answer to these questions, so important to a con-
ception of the origin from which the vertebrates have
sprung; but as long as the said evolution of the Myxi-
noids, in spite of the pains devoted to its study, re-
mains all but unknown, the proper course is to refrain
from drawing any conclusions that are not borne out
to the full by ascertained facts.

The Myxinoid family is represented in the modern
fauna by ordy two genera, Bdellostoma and Myxine. The
former, of which three species, inhabitants of the southern
seas, have been distinguished, is characterized principally
by the presence on each side of the body of six or more
outer branchial apertures, each composing the opening
duct of a gill-sac. In the genus Myxine, on the other
hand, each side of the body is furnished with only one
branchial aperture, the common orifice of the six gill-sacs.

1196

SCANDINAVIAN FISHES.

Of the latter genus too three species have been
distinguished, respectively from the North Atlantic, the
Pacific Ocean (Sea of Japan), and the extreme south
of South America.

As the Scandinavian fauna, however, contains only

one representative of the Myxinoid family, namely
Myxine glutinosa, our principal attention shall be con-
fined below to this species, which has besides in re-
cent times been the most frequent subject of investi-
gation, and whose structure is therefore best known.

Genus MYXINE.

On each side of the forepart of the body ( the throat)
by the six gill-sacs of each half of the body , i

In Myxine , as in Petr omy son, the cylindrical noto-
chord, continuous, unsegmented, persists throughout
life, anteriorly terminating in a conical tip at the basis

a single branchial aperture , at which the ducts given out
if ter uniting into a common duct , find an outlet.

a dorsal and a ventral lamella, which are confluent
behind and form round both sides of the termination
of the spinal cord a more or less narrow, short canal.

a , ventral plate of the caudal cartilage; b , ventral cartilaginous rods (fin-rays) issuing therefrom; c, right sac of the caudal heart; d , button-
shaped process of the ventral plate; e, notochord; /, spinal cord; g , medullispinal canal; h, free dorsal and i, free ventral cartilaginous rods

of the caudal fin; k, dermal muciferous glands.

cranii, under the brain, posteriorly extending close to
the tip of the tail and terminating there after gradual
contraction but somewhat obtusely, before the posterior
extremity of the spinal cord (fig. 354, e). This noto-
chord, composed of large cells filled with a clear fluid
and cemented together by their investing membranes,
is enveloped in two closely fitting sheaths, which have
been somewhat differently interpreted by morphologists.
These are surrounded by fibrous connective tissue,
which does not differentiate, however, into any true
vertebral skeleton. Only at the head and tail are
cartilages developed, but in Myxine they never assume
any vertebrally segmented character. Thus in the tail
(fig. 354, a) we find a median cartilaginous plate with

The outer edges of this cartilaginous plate send out
upwards, backwards, and downwards a great number
of slender cartilaginous rods ib), pointed at the tip,
which compose the supporting rays of the caudal fin,
and not without reason have been compared — or at
least regarded as analogous — to the spinous processes
of higher vertebrates. Even in front of the said carti-
laginous plate there appears in the caudal fin, above
and below, a series of similar rods, the proximal ends
of which terminate in a button-like form within the
septum of connective tissue and are unjoined by cartila-
ginous tissue (fig. 354, h, i). In Myxine fhere is not
a sign, as in Petromyzon , of lateral cartilaginous arches,
to represent vertebral apophyses.

HAG-FISHES.

1197

In the head of Myxine the cartilaginous skeleton
is comparatively far more developed and complex. Here
we meet with two kinds of cartilaginous tissue, one
rather hard, of a yellowish colour, the other soft, colour-
less. This cartilage forms the floor of the cranium, as
well as a number of processes and arches originating
therefrom and composing a singular framework or
“basket-work” within the head. No segmentation into
vertebral parts can be observed, as we have already
stated ; but by means of a comparison with Petromy zon
and its larval form and, above all, with the phenomena
during the development of amphibian larva?, it has
been attempted, not without success, to interpret in
detail the cartilaginous framework of the skull in
My xine. Following Parker’s explanation we here give
a brief description (fig. 355).

The anterior conical termination of the notochord
is surrounded by a sheath of hard cartilage (the para-
chordal cartilage or investing mass, fig. 355, h), which
in the median line exhibits a dorsal fissure and on
the sides passes into the rounded kidney-shaped audi-
tory capsules (c), also formed of hard cartilage, which
have their longitudinal axes directed from within and
behind outwards and forwards, and present on their
inner superior side the foramina for the branches of
the auditory nerve. In front the parachordal cartilage
passes into two slender cartilaginous rods, the so-called
cranial trabeculce (d), which enclose an elongate oval
hole (the basicranial fontanelle, e ) composing the floor
of the cerebral cavity and in great part filled by fibrous
connective tissue, but containing behind and in the
middle a spoon-shaped cartilage (the posterior intertra-
becula, /). Anteriorly the trabecula? pass on each side
into the palatine cartilages (g), which, together with
the ethmoid cartilage ( h ) joining them to each other
in front, form the anterior limit of the fontanelle. From
the anterior ends of the palatine cartilages projects on
each side a cartilaginous rod ( cornu prcepalatinum, i ),
curved outwards and forwards and pointed at the tip.
True cornua trabecularmn are wanting. In the median
line lies a long cartilaginous rod (the anterior inter-
trabecula, k) originating from the dorsal side of the
anterior termination of the fontanelle and from the
ethmoid region. This rod supports the long tube,
formed of transversal cartilaginous rings, that com-
poses the nasal duct ( l ) and leads behind partly to the
olfactory organ, partly to the palate, into the cavity
of which it has a free opening. On each side of the

fore end of the anterior intertrabecula and the anterior
opening of the nasal duct are two pairs of short, some-
what curved, anteriorly pointed rods of cartilage (m),
which are the supports of four — two on each side of
the nasal aperture — tentacles or papilla? (“barbels”). On
the outer side each of the two trabeculae is bounded by
an elongate oval hole ( fenestra suborbitalis, n) and on the
outside of the latter is another cartilage ( pterygoideum ,
o), which here forms the outer edge of the cartilaginous
framework of the skull and behind borders on another,
larger hole ( fenestra lateralis anterior, p), also of an
elongate oval shape. On the inner side of this fenestra

Fig. 355. Skull of Myxine glutinosa, seen from below, X 5.

After Parker.

a, notochord; b, parachordal cartilage; e, auditory capsules; il , trabe-
culae ; e, basicranial fontanelle; f, posterior intertrabecula; g , palatine
cartilage; h, ethmoid cartilage; lit, ethmoidal tooth; i, cornua prae-
palatina; k, anterior intertrabecula; l, nasal duct (in outline); m, carti-
laginous rods of the four circumnasal tentacles; n, suborbital fenestra;
o, pterygoideum; p, anterior lateral fenestra; q, rnandibulo-hyoid
fenestra; r , quadrature ; s, symplectic region; t, interhyoideum ; u, epi-
hyoideum; v, ceratohyoideum, x, first epibranchial ; y, posterior lateral
fenestra; z , first pharyngobranchial; a, second pharyngobranchial.

1198

SCANDINAVIAN FISHES.

the pterygoid cartilage passes into a narrow cartilage
which has been named the mandibulohyoid, and on the
inner side of the last-mentioned cartilage, close to and
below the outside of the auditory capsule, lies a small,
narrow, oval hole ( fenestra mandibulo-Jiyoidea, q). The
small part (Parker’s “pedicle”) in front of this hole
answers to the upper end of the suspensorium ; and
the part ( hyomandibidare ) that bounds the hole behind

Fig. 356. Cartilaginous framework of the tongue in Myxine gluti-
nosa, with the teeth in position, seen from above, X 5.

After Parker.

a, b, basihyal cartilages; c, supralingual cartilage; d , supralingual
teeth; e, basibranchial cartilage.

to the head of the hyoid arch. On the outside of the
anterior lateral fenestra there lie, behind the pterygoi-
deum a broad plate of hard cartilage, answering to the
quadrate region ( quadratum , r), and behind this again
a soft cartilage ( regio symplectica, s ), which forms the

posterior limit of the anterior lateral fenestra and passes
inwards, behind the hard mandibulo-hyoideum, into
another soft cartilage ( interhyoideum , t) and its conti-
nuation ( epihyoideum , u). These last-mentioned carti-
lages emit in an outward direction two soft cartilagi-
nous arches, the anterior (whose upper part is named
the ceratohyoideum (v), the extero-inferior part the
hypohyoideum ) bending outwards and downwards and
attaching itself to a hard cartilage ( basihyoidewn ), the
posterior ( epibranchiale primum , x) curving downwards
and forwards and attaching itself annularly to the sym-
plectic region, together with which it surrounds a large
hole ( fenestra lateralis posterior, y). On the inside of,
but not directly continuous with, the interhyoideum,
epihyoideum, and symplectic region, lies a cartilage
which answers to the first pharyngobranchial bone 0),
and which with its hard, clavate, anterior end, close
to the inner margin of the quadratum, occupies the
posterior part of the anterior lateral cavity and with
its soft, tapering, posterior end reaches a little way
behind the epibranchial cartilage. The second branchial
arch ( pharyngo-branchiale secundum, d) is represented
only by its pharyngeal, upper part, which is here com-
posed of a few medially connected, singularly formed,
branched rods of soft cartilage, embedded in that fold
of the palatine mucous membrane which lies behind
the pharyngeal aperture of the nasal duct.

The lingual apparatus of Myxine displays an espe-
cially powerful development and is furnished with large
cartilages both hard and soft (fig. 356). Thus there
are four hard basihyal cartilages {a, b) and on their
upper surface a part consisting of several cartilages
(the supralingual cartilages , c ), which bear the power-
fully developed supralingual teeth {d). Backwards from
the basihyal cartilage runs a dorsally grooved rod of
fibrocartilage (the basibranchial cartilage, e ), which ex-
tends back to the region of the gill-sacs.

The muscular system of Myxine exhibits about
the same developmental type as that of Petromyzon
and even, in the main, as that of the other piscine
orders, i. e. on each side a “great lateral muscle ” di-
vided into very numerous parts {myomeres) by septa
of connective tissue ( myocommata ), and thus showing a
segmental arrangement, though the axial skeleton is
not segmented, and two ventral muscles. There being
no extremities, or lateral fins, the muscles belonging
to them are also wanting; but the head, on the other
hand, contains rather numerous singular muscles, which

IIAG-FISHE8.

1199

partly enter into the composition of the tongue, partly
are intended for the locomotion in different directions
of the said organ, the mouth and nasal duct, and the
palate and branchial apparatus. These muscles consist
of striated fibre.

As for the blood-vascular system , the heart (fig. 358)
is composed, as in other fishes, of an atrium, a ven-
tricle, and a truncus arteriosus. Of the two Cuvierian
ducts only the left attains full development, and of the
two cardinal veins the right is feebly developed and
joins the left. The blood coming from the hind part
of the body accordingly flows into the left Cuvierian
duct; the hepatic veins are discharged by two trunks
directly into the sinus venosus. The portal vein, which
also receives the numerous veins coming from the gene-
rative organs (mesovarium or mesorchium), dilates into
a wide sac, the “portal heart” discovered by A. Retzius,
before distributing itself to the two lobes of the liver.

The lymphatic and serous system is strongly deve-
loped in Myxine. Under the greater portion of the
cutis there lies, throughout the dorsal region, from the
snout to the tip of the tail, on both sides a large (sub-
cutaneous) lymph- space, which dorsally extends quite to
the median line, where a thin lamina of connective
tissue, provided with nerves and bloodvessels, divides
the two lateral spaces, except in the cephalic region,
where they are in open communication with each other.
Ventrally the space extends close to the row of muci-
ferous glands that runs on each side of the body. But-
on the ventral side" there is no similar, large and con-
tinuous lymph-space, only a- fairly abundant and dense
subcutaneous connective tissue. In the head, however,
especially around the tongue and pharynx, there lie more
profound lymph-spaces, which, according to Klinckow-
Strom0, are in open communication with the large sub-
cutaneous space; even the gill-sacs are furnished, accord-
ing to the same author, with lymph-spaces. Between
these parts of the lymphatic system and the portal
heart, according to him, there apparently exists a- com-
munication. He also describes a direct communication
between the vessels metamerically radiating in the tail
from the lateral (and “ventral”) lymph-spaces and a
vessel running at the margin of the caudal cartilage
and “opening into the paired caudal heart discovered
by Retzius”. This last-mentioned organ (fig. 354, c),
which has a certain resemblance to the pulsating organ

long since observed in the Eel, appears on each side
of the ventral lamella of the caudal cartilage, is fur-
nished with special muscles, and exhibits quick, rhyth-
mical contractions; it is sanguiniferous and discharges
its contents directly into the vena caudalis6. Of interest
is the circumstance made known by Klinckowstrom
that the subcutaneous lymph-spaces as a rule carry a
red fluid with a strong admixture of blood; the ex-
planation of this may well lie supplied in part by the
above-mentioned communications between the blood-
vascular system and the lymphatic system, but the
circumstances require further elucidation, for the burst-
ing of bloodvessels in consequence of the usually
rapid transference of the animal from a great depth
may contribute to the said appearance.

The skin, which owing to the large lymph-spaces
on the greater part of the body is free and moveable,
consists of a rather thick, fibrous cutis and a super-
incumbent epidermis, which is composed of several
layers of polygonal nucleated cells, towards the surface
of the body as a rule somewhat shallower, more flat-
tened. No scale-growths can be traced in the skin.
Of glandular growths in the same only the muciferous
glands so characteristic of the Myxinoicls are known.
These are set in a single row on each side of the belly,
to a number of more than a hundred; the foremost
(fig. 363, g) are situated behind the head (2'5 — 3 cm.
from the tip of the snout); the hindmost extend, in oppo-
sition to what is sometimes stated, a good way beyond
the cloaca, back to the anterior end of the ventral la-
mella of the caudal cartilage (fig. 354, k ), though with
a break behind the cloaca (fig. 363, h). In form they
are oval, and consist of sacs about as large as hemp-
seed, lying under the skin and opening on its surface
each at a small fine pore-like aperture, which apertures
lie in a line at a distance of 2 or 3 mm. from one
another. These glands copiously secrete an extremely
tough, whitish gray mucus, which proves on microscopical
examination to consist of fine, strong threads, which
have lain rolled up in the secretory cells of the glands
and been formed of the protoplasm within those cells.

True fins there are none, but in the median line
the skin forms folds, partly in the caudal region, where
the fold contains the rays of the caudal cartilage and
thus to a certain degree represents a caudal fin, and
partly on the belly, from the cloaca forward to a point

“ Axel Klinckowstrom, Biol. Foreningens Forh., Bd IV, Nos. 1 — -2, 1891.

b G. Retzius, Ein s. g. Caudalherz bei Myxine glutinosa, Biolog. Untersuchungen, N. F. I, 1890.

Scandinavian Fishes.

151

1200

SCANDINAVIAN FISHES.

about 1 cm. behind the branchial apertures, where
there appears a fold triangular in section, thus with
broad base and as a rule with the free flap measuring
only about 2 — 3 mm. in height, which fold may be
regarded as representing a ventral fin.

No external limit is visible between the body and
the head. The tip of the snout terminates dorsally in
a median triangular dermal flap; underneath this is the
slit-like oval nostril (fig. 363, /), flanked on both sides
by two (thus altogether four) conical, pointed tentacles
(papillae) about 3 mm. long and directed forwards and
somewhat upwards. These contain in their axial part
the above-mentioned small cartilaginous rods (fig. 355, m )
and are covered with soft skin. Under the tip of the
snout the anterior surface of the head slopes backwards
and downwards, and 7 — 8 mm. behind the tip of the
snout opens the oral aperture (fig. 363, m), its appear-
ance varying with its functional position. In its su-

k o

Fig. 357. Sagittal section of the cephalic end of Myxine glutinosa, X 4.
a, month aperture; b , nostril; o, nasal duct; d, connecting canal be-
tween the nasal duct and the pharynx; e, pharynx; f, nasal tentacles;
cj, oral tentacles; li , olfactory organ; i, olfactory lobe; k, forebrain;
Z, ganglion habenulae; m, midbrain; n, hindbrain (little brain) ; o, cen-
tral ventricle ; p , afterbrain (medulla oblongata) ; q, myelon ; r, dorsal
muscular layer; s, medullispinal canal; u , notochord.

perior margin may usually be seen, when it is retracted,
two large and two small folds of skin and in its inferior
margin several small folds of a somewhat variable na-
ture. Suctorial mouth or sucking-disk, like those of
Petromyzon , there is none; nor are there any true lips.
On the other hand, both sides of the oral aperture are
furnished in Myxine with two, thus altogether four ten-
tacles (papillae) of a similar kind to those situated round
the nostril. Of these tentacles the inner pair (situated
nearer the mouth) are small and broad; the other pair,
set outside and somewhat above the former, are longer,
narrower, and more pointed. The irregularly slit-like
(“stelliform”) mouth, open only when the animal is

eating, leads into a long buccal cavity, dilating behind,
in the roof of which, about 1 cm. behind the tip of
the snout, is a somewhat crooked horny tooth with
retral tip. This tooth, surrounded by folds of mucous
membrane, is attached to the ethmoid cartilage and is
therefore called the ethmoidal tooth (fig. 355, lit). A
few millimetres further back (13 — 14 mm. behind the
tip of the snout), viz. when the tongue is in its normal
position of rest, lie the supralingual teeth (fig. 356, d),
attached to the upper surface of the tongue (to the
supralingual cartilage) and arranged in anteriorly con-
vex, approximately concentric curves; these are also
horny, and their tips are directed backwards and in-
wards. When the tongue is protruded, this dental
apparatus is visible at the oral aperture. Above and
behind the same opens the above-mentioned tubiform
nasal duct (fig. 357, c ) through a canal (d) sloping
downwards and backwards in the palate, and further
back appears a fold of mucous membrane at the tran-
sition to the pharyngeal tube, which ( e ) runs straight
back above the long tongue-apparatus to pass behind
the branchial apparatus, in the form of an also com-
paratively narrow and long “oesophagus”, into the in-
testinal canal. This (fig. 363, e, f) has the form of
a straight, thick-walled, broad and capacious tube,
furnished internally with shallow, longitudinal, zigzag
folds, but without “spiral coils”, and runs to a point
distant about 3 — 4 cm. from the tip of the tail, where,
after bending downwards (ventrally), it opens into the
cloaca , situated in the median line and nearly 1 cm.
long, which terminates in a sagittal slit, the outer
cloacal aperture (i), flanked by two dermal flaps or lips.

The gall-bladder duct enters the foremost part of
the intestine. The gall-bladder lies between the two
widely separated lobes of the liver (fig. 363, d) and
receives a duct from each of them. Of pancreas and
spleen not a trace has yet been found. The kidney of
Myxine exhibits, as Johannes Muller, in his famous
work on the Myxinoids, first showed, a very remarkable,
extremely low development (“pronephron”); on each side
of the notochord runs a duct opening into the cloaca
and receiving during its course in the abdominal cavity
a number of lateral branches, whose capsule-like dilated
ends each contain a plexus of vessels, a glomerulus
composing a branch of the aorta.

In consequence of the powerful development of the
tongue the branchial apparatus (fig. 358) is removed
far back. It is not furnished with any branchial basket

HAG-FISHES.

1 201

proper, but only with the above-mentioned cartilaginous
rods, situated in its foremost part. The six gill-sacs
(br) on each side communicate inwards by means of a
short transverse duct with the oesophagus; outwards
each of them emits a longer duct (hr), curving down-
wards and outwards. These latter ducts coalesce on
each side into a single canal, thus finding an orifice
on the two outer sides of the body at an opening
situated far back, the two outer branchial apertures
(fig. 358, s; fig. 363, k).

Nervous system. The brain of Myxine exhibits a
rudimentary structure with decidedly retrogressive cha-
racters (figs. 357 and 359). As mentioned above, the
investing cerebral capsule consists of cartilage under-

Fig. 358. Branchial apparatus, heart, branchial artery, and oesophagus
of Myxine glutinosa, after Johannes Muller. Nat. size.

o, oesophagus; i, inner branchial ducts; hr , gill-sacs; br\ outer bran-
chial ducts, which coalesce and have a common opening at s, the outer
branchial aperture; a, atrium of the heart; v, its ventricle; ab, branchial
artery, which sends a division to each sac; d, flaps of skin, folded back.

neath alone; its sides and roof are of a fibrous texture.
The brain is first enveloped in a thin, blood-vascular,
connective tissue, a kind of pi a mater , which descends
into and fills the cerebral sulci. Seen from above (fig.
359) the naked brain is triangular, with the base of
the triangle directed forwards. By a dorsal longitu-
dinal sulcus in the median line a bisection into two
lateral halves is indicated. In each lateral half five
divisions, bounded by transverse sulci, may be traced,
namely, beginning from in front, the olfactory brain,
forebrain , midbrain, hindbrain, and afterbrain. Of the

' tweenbrain there appears dorsally, in the angle between
the forebrain and midbrain, a small, rounded or oval
protuberance, the ganglion habenulae, usually consisting
of a right and a left lobe; but of the epiphysis itself
not a trace can else be detected; on the ventral side
is a small infundibulum with a lamellar appendage,
the hypophysis. The afterbrain (medulla oblongata) is
strongly developed; its anteriorly dilated central canal
advances through the midbrain to a ciecal termination,
somewhat before the posterior half thereof. The third
ventricle is obliterated, with the exception of two small
vestiges. Of lateral ventricles there is no trace. No
ependymal pallium, like that of the Teleosts and Petro-

9

Fig. 359. Brain of Myxine glutinosa, seen from above, X 7.

a, olfactory lobe; b, forebrain; c, midbrain; d, hindbrain; e, after-
brain; f, ganglion habenula;; g, spinal cord; h , nasal duct with carti-
laginous rings; i , olfactory organ with cartilaginous basket; k , eye;
l, optic nerve; m, trigeminus; n, facialis (?); o, trigeminus (?); p,
acoustic nerve ; q, vagus ; r, sensor}- roots of spinal nerves.

myzon larvte, is to be found. It is hence impossible
for the present to determine with certainty in the adult
state whether the forebrain of Myxine merely corre-
sponds to the corpora striata, or whether the cerebral
pallium also enters into its structure.

The nerves issuing from the brain on each side
are: olfactorius, opticus, trigeminus, facialis, acusticus,
and vagus', of the others no trace can be seen.

1202

SCANDINAVIAN FISHES.

The spinal cord consists of a flattened band, of
uniform breadth and somewhat triangular in section,
which in a fairly spacious medullispinal canal, lying on
the notochord, follows the latter to its posterior termi-
nation, and there extends in a contracted ependymal
filum terminate somewhat behind the same (fig. 354,/).
By means of improved colouring methods the form, dis-
tribution, and processes of the ependymal as well as
the neurogliar and ganglion cells has been elucidated on
a fairly extensive scale in the spinal cord and partly
too in the brain.

The nerves originating in pairs, but not quite sym-
metrically, from the spinal cord (spinal nerves) each
have a ventral (motory) and a dorsal (sensory) root,
and the latter is furnished with a ganglion. As in
Petromy zon, all the nerve-fibres are “pale”, being with-
out myeline sheath. The motory nerve-fibres end their
terminal ramifications on the muscular fibres in free

D- /’ c

— 0

A

*

... /XA,

4 # 1

h |

.r" -if<

f\V 0

flAt 1

. J ' ...'a?---

v'

i g d

Fig. 360. Auditory organ of the right side of Myxine glutinosa ,
seen from above and within, X 18.
a, internal division with e, the macula acustica; b, external division
with an ampulla at each end (c, d) and a crista acustica (/, g ) in
each ampulla; i, i, branches of the acoustic nerve; h, endolymphatic duct.

“end-trees” within more or less marked end-hillocks
(end-plates). The sensory fibres terminate within the
different organs also in free intercellular ramifications.
Such is the case too with the nerves in the epithelium
of the skin and mucous membranes. No special sensory
organs (end-cells, end-buds, sensory corpuscles, etc.)
have been demonstrated. Even in the tentacles sur-
rounding the oral and nasal apertures only free terminal
ramifications of the sensory nerves in the epidermis
have been traced, but here there is a slight sign of
end-buds, round whose cells the nerve-ends ramify.

No lateral line with appertaining end-buds appears either
on the head or the body. In the subcutaneous nerve-
plexus, on the other hand, especially in the head, nu-
merous ganglion cells have been found embedded. Pos-
sibly they belong to the sympathetic nervous system ,
whereof else no trace has hitherto been discovered.
These ganglion cells further remind us by their po-
sition in some degree of the cells in the rostral region
of the Lancelet, close to the foremost nerve-pair.

Of the organs of sense proper the olfactory organ
is obviously the most powerfully developed. It forms
(fig. 359, i ) a conical organ situated close before the
brain in a cartilaginous basket and consisting of nearly
vertical and dense folds of mucous membrane, their
surface covered with olfactory epithelium, i. e. olfactory
cells with supporting cells arranged between them. The
filamentous, central processes of the olfactory cells enter
the olfactory bulb, and- terminate there in numerous
ramifications and with free ends in the olfactory glome-
ruli, in which other nerve-fibres, from the ganglion
cells (mitral cells) of the bulb, running towards the
periphery, also terminate in free ramifications. The ol-
factory organ of Myxine thus exhibits in the main the
same structure as this organ retains throughout the
whole of the vertebrates, even including man. Here
there is no trace of “olfactory end-buds”, as Blaue
has endeavoured to demonstrate in the Teleosts. Neither
in the latter nor the Cyclostomes, nor indeed in any of
the vertebrates, do such “end-buds” exist as Blaue
meant; and his theory of the genetic connexion between
the olfactory organ and the sense-organs of the skin,
especially the lateral line, is therefore without ground
in the structure of the said organs.

Special organs of taste have not yet been demon-
strated in Myxine, whereas Petromy zon and its larva dis-
play within the mouth cavity bulbous or gemmiform
organs that may with reason be regarded as represent-
ing the gustatory organs of the higher vertebrates®.

The organ of sight in Myxine 6 is very rudimentary.
It is composed of the two very small, oblong eyes dis-
covered by Johannes Muller, which, covered by the
skin and a layer of muscles, lie on each side of the
fore end of the brain (olfactory brain) and close to the
posterior outer angle of the cartilaginous basket invest-
ing the olfactory organ (fig. 359, h). The two optic

° Cf. G. Retzius, Biol. Unters., N. Folge, Bd. V, p. 69, taf. XXVII.
b Cf. „ , 1. c., p. 64, taf. XXVI.

HAG-FISHES.

1203

nerves (l), which, after crossing, emerge from the vent-
ral side of the midbrain and afterwards turn first out-
wards and then forwards, penetrate the hind extremity
of either eye, and their fibres may be traced some way
in within its central parts. The eyes, surrounded by
a capsule of connective tissue, do not show the regular
structure common in the vertebrates; cornea, iris, and
lens are entirely wanting, and of vitreous body there is
only a rudiment; the retina does not present the nor-
mal arrangement, it fills nearly the whole orbit, its
stratification is very irregular and difficult of elucida-
tion, instead of rods and cones there is only an epi-
thelial layer of obscure nature. By means of the mo-
dern colouring methods the presence of numerous cells
of neuroglia-like appearance may be demonstrated, but
also of some other cells that may possibly be of a true
nervous description. Hence it appears that the eye of

each side of the anterior end of the notochord at the
bottom of the cranium (fig. 355, c ). In the annular
tube may be distinguished an inner division (tig. 360, a),
furnished with a long nerve-ending, a macula acustica
(e), and an outer division (5), furnished at each end
with an ampullaceous dilatation (c, d) and a nerve-
ending, thus with two cristas acustica? (/, g). This
outer division may reasonably be interpreted as repre-
senting two semicircular canals, each with an ampulla.
The inner division gives out a ceecally terminating en-
dolymphatic duct ill).

It is else not easy to institute any close comparison
between the so little differentiated parts of the auditory
organ in Myxine and those of the corresponding organ
in other fishes and the higher vertebrates. Unfortuna-
tely this applies also to the same organ in Petromyzon ,
where it has many peculiarities and forms no true link

c b

e' d

Fig. 361. Auditory organ (right) of a Lampern ( Petromyzon fluviatilis ), X 18.

A, seen from below; B, seen from above and without: a, a1, vestibule; 6, anterior semicircular canal; c, c\ anterior ampulla; d, posterior
semicircular canal; e, e1, posterior ampulla; /, saccate appendage; f1, its nerve-ending; fa, commissure; g , ganglion of the acoustic nerve; h , acoustic
nerve; i, cerebral part from which the acoustic nerve originates; k, anterior branch of the acoustic nerve; l , posterior branch of the acoustic nerve.

Myxine is not only undeveloped, but has probably also
undergone retrogressive development. That no true sight
exists, is evident from the structure of the eye and its
concealment below the surface of the body. Yet phy-
siological experiments might well be made to investigate
whether, in spite of this, the fish may not be capable
of appreciating with these rudimentary eyes certain
intense lights.

The auditory organ of Myxine (fig. 360) is struc-
turally the simplest known within the vertebrate king-
dom, if we except Branchiostoma , in which no trace of
such an organ has been discovered. It consists of a
paired, annular, membranous organ, which lies enclosed
in the above-described cartilaginous capsule situated on

between that of Myxine and of the other vertebrates.
In Petromyzon (fig. 361) may be distinguished a large
common chamber (the vestibule), a saccate appendage,
and two semicircular canals each with an ampulla and
appertaining crista acustica. In the vestibule is a large
nerve-ending (macula acustica) perhaps composed of se-
veral, and a smaller one appears in the saccate appendage.

It is first in the osseous and cartilaginous fishes
(fig. 362, A, B, C , B) that the membranous auditory or-
gan is developed to that fundamental type which it
afterwards retains throughout its evolution within the
vertebrate kingdom, even in the human race. It parts
into two divisions, a, superior and an inferior. The
superior consists of a fairly wide, horizontally and sagit-

1204

SCANDINAVIAN FISHES.

tally directed tube, utriculus (u), which dilates anteriorly
into a vesicular bulb, recessus utriculi ( rec ), in whose
anterior and outer sides open two vesicular tubes, the
anterior (aa) and the external {ae) ampullce, continued
each by a narrower, tubiform duct, the anterior ( ca )
and the external {ce) semicircular canals, which bend

round and at their other end again open into the utri-
culus, the former at the upper end of an upward in-
vagination thereof, sinus utriculi superior {ss), the latter
into the lower part of it. At the posterior termination
of the utriculus, sinus utriculi posterior ( sp ), opens a
third vesicular tube, the posterior ampulla ( ap ), which

Fig. 362. Auditory organ of the right side: A, in a Perch ( Perea fiuviatilis ; X 3, seen from within); B, in a Sturgeon ( Acipenser sturio;
X 37a, seen from without); C, in a Chimcera ■ (Chimcera rnonstrosa; X 3 1 / 4 , seen from within); D, in a Rough Hound ( Scylliorhwus cani-
cula; X 3l/3), and E, in a Mud-eel ( Siren lacertina ; X 1 1 1/4), both the last seen from within.

aa, ampulla anterior

ае, ,. externa

ар, ,, posterior

ас, nervus acusticus

ade, apertura externa ductus endolympliatici
adp, ,, ductus perilymphatici
ass, apex sinus superioris

с, cutis

ca, canalis anterior

ce, ,, externus

cp, ,, posterior

cr, crista acustica ampullarum

ers, canalis recessu-saccularis

cus, canalis utriculo-saccularis
cle, ductus endolymphaticus
dep, ductus canalis posterioris
dp, „ perilymphaticus

l, lagena cochleae
mn, macula acustica neglecta

ms, ,, ,, sacculi

mu, ,, ,, recessus utriculi

o, otolites (in the recessus utriculi, sacculus
and lagena)

pi, papilla acustica lagenae
raa, ramulus ampullae anterioris
rae, ,, ,, externa?,

rap, ramulus ampullae posterioris
rl, ,, lageuae

rn, ,, neglectus

rs, ,, sacculi

ru , ,, recessus utriculi

rec, recessus utriculi
s, sacculus

se, saccus endolymphaticus

sp, sinus utriculi posterior

ss, ,, ,, superior

sq, squamae cutis
u, utriculus.

AUDITORY ORGANS.

1205

passes behind into a third semicircular canal , canalis
posterior ( cp ), that bends upwards and afterwards for-
wards and inwards, opening into the upper end of the
superior sinus (.ss).

The superior division of the organ usually com-
municates by means of a small opening or a short tube,
canalis utriculo-saccularis (cus), in the bottom of the
utriculus, with the inferior division. The latter consists
of a saccate part, sacculus (s), which in most fishes
possesses behind a pouch-like appendage, lagena coch-
leae (l), the first rudiment of the cochlea so richly deve-
loped in the higher animals and, above all, in man,
together with its highest development (the organ of
Corti). From the inner superior side of the sacculus
there issues besides a narrow canal, ductus endolympha-
ticus, which as a rule lays itself on the membranes of
the brain and terminates csecally, but sometimes, as in
the Elasmobranchs, ascends to the upper surface of the
head, where it has a free orifice.

Of nerve-enclings the superior division contains at
least 4, namely a large one in the bottom of the re-
cessus utriculi ( macula ac. recessus utriculi , mu) and
one in each of the three ampullae ( cristce ac. ampullce
anterioris, externce, and posterioris, cr), besides which
there appears in most fishes at the bottom of the utri-
culus itself, in the neighbourhood of the opening into
the sacculus, a small nerve-endiug ( macula ac. neglecta ,
mn). In the inferior division may be found, within
the sacculus, a large ending ( macula ac. sacculi , ms)
and in most cases another, posterior one ( papilla ac.
lagence , pi), which is situated on the wall of the pouch-
like cochlear appendage, as soon as this is developed.
Thus in some fishes the number of nerve-endings is
only 6, in others 7. These endings are everywhere of
the same structure, the epithelium which lines them
consisting of numerous cells (auditory or hair cells)
furnished with hair-like processes (auditory hairs) pro-
jecting freely into the endolymphatic fluid, and of
supporting cells set between the former and isolating
them.

The acoustic nerve divides into several branches,
which supply all these nerve-endings each with a bundle
of nerve-fibres. These fibres, each of which issues
from a bipolar ganglion cell embedded in the acoustic
nerve and its branches, penetrate the epithelium of
the nerve- endings and ramify around the auditory cells,
over Avhich they spin free terminal ramifications, with-
out being directly continuous therewith.

Here we give some figures (fig. 362) to illustrate
the various development of the auditory organ in carti-
laginous and osseous fishes, namely Cliimcera, Acipen-
ser, and Perea, adding a figure of Scylliorliinus to show
further how the organ has developed a special type in
the Sharks and Rays. The auditory organ of the Lung-
fishes and amphibians is most closely linked, among
the forms adduced here, to the Ganoid type, as shown
by fig. 362, E , which represents the appearance of this
organ in Siren lacertina. The present, however, is no
suitable opportunity for a, closer consideration of these
questions.

From the above it appears that, as we have al-
ready pointed out, the auditory organ of Myxine may
justly be regarded as the lowest known form of the
development of this organ among the vertebrates, but
that, the intermediate links being absent, we can only
imperfectly sketch its phylogenesis. In Myxine a di-
vision into a superior and an inferior part has pre-
sumably not yet taken place, and the inner part of the
annular tube probably represents both utriculus and
sacculus, which is also indicated by the emission from
this part of the endolymphatic duct. The outer part
of the annular tube probably answers to the anterior
(or the outer) and the posterior semicircular canals with
their respective ampullse. Possibly in the future, when
the development of Myxine has been brought to light,
some knowledge may be gained both of this question
and so many others, touching the systematic position
of this remarkable creature and its relation to the rest
of the vertebrates.

In the anatomical structure of Myxine we have
still to consider the organs of generation and in con-
nexion therewith the present knowledge of the im-
pregnation and development.

The structure of the ovary and eggs has long been
known. In the abdominal cavity of most specimens is
found along the right side of the intestine a somewhat
lobate, longitudinal mesentery, within which are en-
closed, as it were suspended, rather numerous ova of
varying dimensions. In small individuals these ova are
generally of insignificant size and rounded form, some
hardly visible to the naked eye, other somewhat larger,
up to the size of a pin’s head. In more full-grown
individuals many similar eggs are also found, but there
besides appear, as a rule, a number (10 — 15 or more)
of larger ova, which are always oblong, fusiform, mea-
suring in some cases only 3 — 10 mm., in others 11 — 12

1206

SCANDINAVIAN FISHES.

mm. These eggs are enveloped in a firm membrane
and yellowish in colour; the greater part of their con-
tents consists of yolk. In its hindmost part the said
ovarian mesentery (mesovarium) presents a totally dif-
ferent appearance; no eggs are visible here, and the
mesentery assumes a still more lobate, tuberculate or
granular form. On microscopical examination no eggs
are found here, but only vesicles or capsules, contain-
ing a great number of large cells, many of which are
in process of division. I. T. Cunningham" was the
first to express the opinion, based on his investigations,
that this hindmost part of the mesentery is a mesor-
chium and contains the male organ. Nay, he even
considered that he had here found spermatozoa at dif-
ferent stages of development. Fridtjof Nansen2, fol-
lowed up these researches and adopted the theory of
hermaphroditism, adding that he had observed that, in
young individuals which had not yet attained a length
of 320 — 330 mm., the male organ was, as a rule, the
more developed and contained almost ripe spermatozoa,
whereas the female organ was more or less undeve-
loped. Larger (older) individuals he found, on the
other hand, to contain more developed female organs
and also large eggs, while the male organ was more
rudimentary. Hence he concluded that in the younger
stages of its development Myxine is masculine (prot-
andric), but afterwards in the older stages feminine.
But there besides occur, according to Nansen, indivi-
duals that are males alone. The same investigator also
gives figures of spermatozoa he had observed in Myxine ,
and asserts that those described by Cunningham are
not of a true nature. The latter writer has subsequently
given a new description of the spermatozoa observed
by him, and insisted upon his priority with respect to
the theory of hermaphroditism. As I myself too, during
a series of years (since 1877), have made repeated in-
vestigations into the structure of Myxine and its mys-
terious development, I will state that, so far as I can
find, no distinctly developed male individual thereof
has hitherto been observed with certainty, and that
there is much 'to be said for the Cunningham-Nansen
theory of its hermaphroditism. Considering that I have
macroscopically examined a great number of Hag-fishes
and have microscopically prosecuted these investigations
in many instances that seemed especially remarkable as

regards the point in question, and this at every season
of the year, it may appear strange that I have never suc-
ceeded in finding within the cell-filled vesicles of the so-
called mesorchium fully developed spermatozoa, but only
lower and somewhat dubious stages thereof. Yet from
preparations kindly sent me by Nansen it would seem
that the structures found by him presented in a high
degree the appearance of spermatozoa. Phenomena of
movement, however, have not yet been observed in
them, and even their development is not fully known®.

As for the season of impregnation and oviposition,
these do not appear to be restricted to any fixed time
of year. All the investigators who have made any
minute researches into this question are apparently
agreed that ripening eggs are met with in these crea-
tures all the year round, but that at the same time in
other individuals ova may be found at the most different
stages, from cjuite small, colourless globules, hardly
distinguishable to the naked eye, to the opaque, yellow-
ish eggs 10 — 12 mm. long, of an elongated oval form,
pointed at the ends, enveloped in a horny shell, and
filled for the greater part by a large yolk. Of such
ripening, but not yet fully developed eggs a number
of 10 — 15, up to 26, are as a rule found in the same
individual, all then exhibiting the same size and ap-
pearance. These ova, however, as mentioned above,
are not quite ripe for deposition. There are a few in-
stances in which specimens have been secured of fully
developed eggs. In 1859 Allen Thomson figured and
described in Todd’s Cyclopaedia of Anatomy and Phy-
siology ova of Myxine that measured about 25 mm. in
length, had an oval form, and were furnished with a
firm, horny shell and at each end with a number of
horny threads, each having hooks (barbs) at the end.
Some years afterwards J. Steenstrup (Oversigt af d.
K. Danske Vidensk. Selsk. Forhandl. f. 1863) described
and figured eggs of similar appearance that had been
found in a Glutinous Hag. A. W. Malm (Goteb., Bohusl.
Fauna) also secured ripe eggs 19 mm. long that were
met with in the stomach of a Cod (thus presumably
devoured after their deposition); a couple of these were
described in 1875 by W. Muller in the “Jenaische
Zeitschrift f. Naturwissenschaft”. Furthermore I. T.
Cunningham has described in the Quart. Journ. of Micr.
Sc., N. S., Y. 27, 1887 an egg preserved in the Edin-

a I. T. Cunningham, Quart. Journal of Microsc. Science, N. S., V. 27, 1887.
h Fridtjof Nansen, Bergens Museums Aarsberetning f. 1887 (ed. 1888).
c See further G. Retzius, Biolog. Foreningens Forhandl., Bd 2, No. 8, 1889 — 90.

HAG-FISHES.

1207

burgh Anatomical Museum (possibly the same specimen
as formerly described by Thomson). Nansen also states
that he observed in the Museum of Bergen a ripe egg
which had been dredged up by I). C. Danielssen at
Molde. In July, 1888, while examining a large num-
ber of Glutinous Hags from Gullmar Fjord, I found in
an individual 30 cm. long, containing 12 large eggs, a
ripe egg furnished with threads at the ends (G. Retzius,
Biol. Foren. Forhand., Bd. 1, Okt. 1888). This egg
was still attached in the mesovarium, the third from
behind in the chain of ripening ova. It was larger
than the others, measuring 14 mm., had a yellower
colour, was blunter at the ends, and harder on the

ripe as those described by Steenstrup and the other
above-mentioned writers. Since that occasion I have
opened and examined several thousand Glutinous Hags,
but I never succeeded in finding another egg so far
advanced in development.

On the 1st of January last (1895), however, Baron
Axel Klinckowstrom (Ofvers. K. Vet. Akad. FOrh. 1895,
No. 1) found, in a Glutinous Hag taken together with a

o

number of others in Gullmar Fjord by Fisherman Aback
a day or two before, an egg at about the same stage
of development as that described by me. This egg,
which was of a bright reddish yellow, almost orange-
coloured, lay entirely free in the abdominal cavity and

Fig. 363. A Glutinous Hag, nat. size, seen obliquely from below and from the left. Abdominal cavity cut open to show its contents.
a , lobes of the so-called mesorchium (male organ); b, a large, nearly ripe egg with thick, horny shell and horny appendages at both ends,
of which the anterior was still attached to the mesovarium; c, less ripe eggs in the mesovarium; d , liver; e, /, intestinal canal; g, apertures
of the foremost muciferous glands of the skin; h, apertures of the hindmost group of muciferous glands; i, cloacal aperture; h , outer branchial

apertures; l, nasal aperture; in, oral aperture.

surface, which evidently consisted of a strong, horny
shell. The ends were each furnished with a horny
process 4 mm. long, which proved to consist of a bunch
of dense threads, all of them tipped with a button -
shaped thickening armed with four barbs. One (the
anterior) of these processes was attached to the con-
nective tissue of the mesovarium; otherwise the egg
hung free in the abdominal cavity. Fig. 363 repre-
sents this specimen of Myxine with the abdominal ca-
vity cut open and the ripened egg (b) suspended from
the mesovarium. Probably this egg was not quite so

dropped out on the latter being opened. Its total length
was 20‘5 mm., the egg itself measuring 16 mm., and
the two terminal bunches of barbed threads respectively
3 and 1’5 mm. This find is of special interest as con-
firming the opinion that the spawning of the Glutinous
Hag is not restricted to an}- particular season.

This is in effect our whole knowledge of this
question. Manifold endeavours have been made during
the last two decennia by several investigators (W. Muller,
Cunningham, Nansen, myself, Tiieel, Tullberg, Klinc-
kowstrom) to trace the development of the Glutinous Hag;

Scandinavian Fishes.

152

1208

SCANDINAVIAN FISHES.

but hitherto they have proved all but fruitless. It has
been attempted, for instances, with dredges of a special
construction to collect in the clayey bottoms where the
Glutinous Hag has its home deposited ova and fry in
course of development. Further experiments have been
made by keeping a number of Hags in aquaria, or by
sinking them in corves to the depths where they live
in a state of nature and examining at intervals the
contents of the corves; but all to no purpose. Yet it
is quite possible that the question may be solved by
still more systematical and comprehensive investigations,
attention being paid to the biotic conditions of the Glu-
tinous Hag (the temperature of the water, the pressure,
etc.). Or perhaps, a circumstance of which the history
of science affords more than one example, a happy ac-
cident may throw light upon the mystery. In any case
each new contribution to this result, each new sugges-
tion is of value. It is not yet known with certainty
whether the eggs are impregnated after or before their
deposition. W. Muller and Cunningham have described
a micropyle in the ripe eggs. An impregnation of the
ova after their exclusion from the parent-fish is ac-

cordingly not impossible. Nor is it known how and
where the eggs are laid, whether this is done on the
clayey bottom, as is most probable, or perhaps among
algae on a rocky ground. It has been stated that the
Glutinous Hag constructs with its slime a kind of
nest in the clay for the reception of its ova; but
nothing positive is known on this head. It is not
known whether the embryos live in the clay or free
in the sea- water. Their appearance is utterly un-
known. Perhaps they are larval forms of quite a pe-
culiar type, as the Ammoccetes form of Petromyzon.
The smallest recorded specimens of the Glutinous
Hag, taken alone now and then in pots baited with
dead fish, measure no less than 9 cm., and they
already conform in structure to the developed animal.
It consequently remains for the investigator to secure
them at earlier stages and, above all, to find new-laid,
impregnated eggs, as well as to watch the earliest
development of the embryos. This question acquires
greater and greater biological importance year by year,
and it is to be hoped that the solution thereof will
not baftie research too long.

THE GLUTINOUS HAG OR MYXINE (sw. piralen or pilalen).

MYXINE GLUT1NOSA, Linn.

Plate LIII, fig. 5.

Sup rating ual teeth larger in the anterior outer than in the posterior inner row , and numbering in the former 7
or 8, in the latter 8 — 10, the two foremost being besides confluent at the base.

Syn. Pihral 1. Ingeris Pi It, Kalm, Peso N. Amer ., tom. I, p.

100. Hviid-Aal , Strom, Sondm. Beskr., pt. I, p. 287.
Steensuer, Olav., Skag. Beskr., p. 156, tab. 3, figg. 3 et 4.
Myxina glutinosa , Lin., Mus. Ad. Frid ., tom. I, p. 91 (inter
Vermes), tab. VIII, fig. 4; Syst. Nat., ed. X, tom. I, p.
650 ( Myxine ); Gunnerus {Sleep- Marken), Trondhj. Selsk.
Skr., vol. II (1763), p. 250, tab. Ill; Penn. {Glutinous Hag),
Brit. Zool., vol. IV (ed. 1777), p. 33, tab. XX, No. 15;
A. J. Retz., Vet. Akad. Handl. 1790, p. 110, tab. IV (sim.
cum Petromyzonibus inter Vermes et Pisces interm.); Fare.,
Fna Groenl., p. 344 (inter Vermes ); A. J. Retz., Fna Suec.
Lin., p. 301 (inter Pisces)', A. Retz., Vet. Akad. Handl.
1822, p. 233, tab. Ill; 1824, p. 408, tab. VI; Nilss., Prodr.
Ichthyol. Scand., p. 123; J. MWll., Vergl. Anat. Myxin., Abh.
Akad. Wiss. Berk, Pliys. Cl., 1834. 1837, 1838; Sund.,
v. Wr., Skand. Fisk., ed. 1, p. 121, tab. 28; Kr., Damn.
Fisk., vol. Ill, p. 1068; Nilss., Skand. Fna, Fisk., p. 750;
Strp, Overs. D. Vid. Sels. Fork. 1864, p. 233; Couch {Borer),
Fish. Brit. Isl., vol. IV, p. 408, tab. CCXLVIII, fig. 3;
Gthr, Cat. Brit. Mus., Fish., vol. VIII, p. 510; Putn., Proc.
Bost. Soc. Nat. Hist. 1873, p. 135; Coll., Forh. Vid. Sels.
Clirnia 1874, Tillasgsh., p. 220; Malm, Gbgs, Boh. Fna,

p. 637 ; Winth., Naturli. Tidskr. Kbhvn, ser. 3, vol. XII, p.
62; Mela, Vert. Fenn ., p. 372, tab. X; Storm, Vid. Sels.
Skrift. Trondhj. 1883, p. 48; Day, Fish. Gt. Brit., Irel.,
vol. II, p. 364, tab. CLXXIX, fig. 3; Cunningh., 1. supr. c.;
Nansen, 1. supr. c. ; G. Retz., Biol. For. Forh. Sthlm, Bd
I, p. 22, tab. Ill; Bd II, p. 80, tab. Ill; Lillj., Sv., Norg.
Fisk., vol. Ill, p. 730.

Gastrobranchus coecus, Bl., Ausl. Fisch., pt. IX, p. 67, tab.
CCCCXIII.

My, vine limosa, Gir., Proc. Acad. Nat. Sc. Philad. 1858, p. 223.

The Flag that lives on the west coast of Swe-
den and in Norwegian waters has an average length
of 25 — 30 cm. Sometimes, however, individuals mea-
suring 45 cm. or a little more are met with.

The body is of almost uniform thickness, for the
most part terete, Eel-like; behind the middle it shows
some lateral compression, and this is increased towards
the tail, which is thin, flattened, and somewhat dia-

GLUTINOUS HAG.

1209

phanous. The coloration is of a, reddish gray, more
or less strongly suffused on the back and the upper
part of the sides with blackish gray, owing to the pre-
sence of subcutaneous pigment, a vague bluish violet
tint being thus produced in those regions. Sometimes
the pigment is unevenly distributed, giving the body
a piebald appearance. On the belly the colour is of
a somewhat lighter gray.

The head is not delimited from the body, being of
about the same thickness and terete form, but it tapers
in a forward direction towards the tip of the snout.

In its appearance and structure the species be-
longing to the Scandinavian fauna exhibits all the pe-
culiarities enumerated above as characteristic of the
genus Myxine. and no further description is therefore
necessary.

The Glutinous Hag occurs throughout the west coast
of Sweden, from the Sound upwards, and along the whole
Norwegian coast up to Finnmark. In the Sound, how-
ever, it appears to be very rare. From Mount Kullen
upwards, along the west coast of Sweden, and all the
coasts of Norway, it is common in such localities as
are congenial to its manner of life, viz. where the bot-
tom consists of soft clay and mud, the water is of a
suitable depth, about 20“ to 50 fathoms, and preferably
where there is some ground current. The Glutinous
Hag is quite at home, however, in deeper water (100
— 200 fathoms), as has been observed off the northern
most coasts of Norway. It seems besides to occur in
the Arctic seas, off Greenland (Fabricius) and the east
coast of North America, though it is apparently not
common there. On the coasts of Scotland and Northern
England it is also met with, but is less numerous, to
judge by the statements on record, than in Scandi-
navian waters.

As regards its manner of life, it seems as a rule
to lie embedded in the clayey bottom. This is its
habit too, when kept in aquaria with a bottom of clay
or mud, and supplied with running water, an experi-
ment which I have repeatedly made. It soon burrows
into the clay and remains lying there, with only the
tip of the snout exposed to view. But it does not live
long in captivity. As a rule the. prisoners die in the

and sunk in salt water at a depth of some fathoms.
At first they show great activity in their movements,
wriggling like Eels, generally forwards, but sometimes
backwards, and swaying the head now to one side, now
to the other. In these actions they display fairly great
strength, and they secrete meanwhile large quantities
of slime, in which they sometimes entangle themselves.
They often project the tongue and draw it in again,
thus getting portions of the slimy secretion into their
mouth, and the dead Hags are consequently found in
many cases to have clots of slime adhering far back
in the mouth cavity.

Of their habits little or nothing more is known.
That they are true parasites, as was formerly asserted
and is still stated in some foreign manuals, is in the
highest degree improbable. At least there is no direct
evidence of this. They apparently live on dead ani-
mals, principally fish. This appears distinctly from
their not seldom attacking netted fish, or creeping into
pots containing dead fish or fish oft’al and sunk in
places frequented by this species. The most enticing
bait seems to be Haddock or Whiting, and by this
method great numbers of Glutinous Hags may be taken.
In one single pot a hundred or two may sometimes be
caught. On examination the intestinal canal of the
captured Hags is found to be full of recently de-
voured morsels of fish. They apparently do not with-
draw to any distance from the sea-bottom, and accord-
ing to the statements of experienced fishermen they
fasten upon netted or hooked fish only when this is
on or close to the ground.

That the Glutinous Hag also attacks the corpses of
drowned human beings, is a fact known on the west
coast of Sweden.

The Glutinous Hag would thus appear strictly to
feed on dead creatures. It may be regarded as a sca-
venger of the depths. But, guided bjT its powerfully
developed organ of smell, it also assails hooked fish
hanging on the line, and must thus be stigmatized as
in some degree injurious to the fishery.

Itself it possesses no economical value. Its flesh
is not good to eat and is nowhere used for food. Pos-
sibly it might be smoked for consumption, like the
Lampern. ' (G. Retzius.)

course of a few hours, even if they be placed in a corf

a The least depth at which I succeeded in obtaining it in Gullinar Fjord was 16 — 17 fathoms, but, according to statements given by
fishermen, the animal lives during the winter at a still less depth (10 — 11 fathoms).

1210

SCANDINAVIAN FISHES.

PISCES LEPTOCARDII.

Fishes whose unaltered notochord extends further forward than the spinal cord, which is
of uniform thickness and without true brain. No head proper; no true heart. Respiratory

apparatus a large, retiform branchial basket.

Fig. 364. Lancelet ( Branchiqstoma lanceolatum ), seen from the left, with the skin and muscles removed from this side. About 3 times

nat. size. After Vogt and Yong.

a , dorsal, b, ventral fin-ray growths; c, organs of generation; d, branchial basket; e, anal aperture; /, notochord; g , spinal cord; h , intestinal
appendage (hepatic caecum), situated on the right side, seen through the branchial basket; i, intestine; k, mouth; l, great lateral muscle;

m, rostral fin; n, caudal fin; o, abdominal pore.

Here it may be questioned even more than in the
preceding order whether these creatures are vertebrates
or invertebrates. It is not only the low rank occupied
by this order in comparison with the preceding ones;
we shall besides find here an asymmetry that strongly
reminds us of certain invertebrates.

Johannes Muller interpreted this order as a se-
parate subclass among fishes — but so he also regarded
the Cyclostomes and Elasmobranchs — and gave it the
name of Leptocardii11. Under the name of Cirrostomi
the order was coupled by Owen* 6 together with the
Cyclostomes in a subclass, Dermopteri (with fins in the
form of dermal folds). Others have gone still further
and regarded the order as a separate class among the
vertebrates. So the Leptocardii , under the same name,
were conceived by Gillc; and E. Heckel called them*6
Vertebrata Acrania. Others e have indeed considered

these creatures to be the lowest vertebrates, but have
assumed that they attained this position by a retro-
grade development (degeneration) from better equipped
vertebrates, which have now disappeared. Others again
have ranged them beside the Tunicata, and regarded
them as one of the types for the predecessors of the
protovertebrates (the supposed ancestors of the verteb-
rates). Thus Balfour called them7, together with the
Tunicates, Proto chord at a (primordial forms among the
animals furnished with permanent notochord), and con-
ferred upon them the special name of Cephalochorda
(with notochord in the head), a reference to the for-
ward extension of the notochord in that part which
answers to the head of the vertebrates, as distinguished
from the Tunicates ( Urochorda ), which have a notochord
only in the tails of the larvae. The significance of
these divergent opinions is best appreciated after we

a Ub. d. Bau u. d. Qrenzen der Ganoiden, Abh. Akad. Wiss. Berk, 1844, Physik. Abh., p. 204, Xsmog, thin and yMgdla, heart.

6 Comp. Anat ., Phys. Vertebr., vol. I, p. 7.
c Arr. Fam. Fish., Smiths. Misc. Coll., No. 247, p. IX.
d Gener. Morphol. Organ., Bd 2, p. CXIX.
e Dohrn, Ursp. Wirbelth., Leipz. 1875, pp. 51 — 55.
f Comp. Embry ol., vol. II, p. 271.

LEPTOCARDIANS.

1211

have gained an insight into the structure and develop- The few forms of the order belong to a single

ment of the Lept'ocardians. ' family, the

Fam. AIPHI0XI1)

whose characters thus coincide with those of the order.
Nor have reasons been discovered for the establishment

within this family of more than one genus.

Genus BRANCHIOSTOMA.

Body more or less elongated , lanceolate , pointed and laterally compressed at loth ends , intermediately triangular
in section. Mouth ventral, elliptical, fringed with tentacles, hut jawless. A dermal fold edges the body from
the mouth round the tip of the snout, along the lack and round the tip of the tail, and, usually b, forward along
the ventral margin to the abdominal pore, whence two lateral folds extend to the mouth. Anus situated far lack,

beside the ventral margin.

This genus is probably dispersed throughout the
temperate and tropical coasts of all the oceans, but
sporadically. It is known from Europe, Australia, Peru,
Brazil, and the east coast of North America. Where
it occurs, it is generally met with in numbers. Usu-
ally it lives on sandbanks, where it has shelter and
plenty of food. Its diet appears to consist exclusively
of Infusoria and similar minute creatures or the lowest
vegetable forms. If a Branchiostoma (PI. LIII, fig. 6)
be placed in a vessel of seawater with a little sand on
the bottom, it will perhaps lie still on its side for a
long while, as if dead; but on being touched it leaps
up and tosses itself to and fro with swaying move-
ments, until it eventually lies down to rest again on
the sand or buries itself in the bottom. This is done
with great rapidity, in a moment; and afterwards it
only sticks the oral end up into the water, opens its
wreath of tentacles, and commences in its mouth cavity
the ciliary motion that conducts water and food into the
respiratory cavity, which also serves as an oesophagus.

That which first gives this animal, in contradis-
tinction to the rest of the vertebrates, a singular posi-
tion in the system, is, as mentioned above, its almost
entire lack of that we call head. With the same justice
as we call a mussel a headless mollusk, we may also
describe Branchiostoma as a headless vertebrate.

Fig. 365. Anterior end of the body of the Lancelet, seen from the

right, magnified.

a, site of the olfactory organ (situated on the left side); b, site of
the eye (pigment spot); c, second pair of nerves, counting from in
front; d, spinal nerves; e, homologues of the upper spinous processes
of vertebras ; /, cartilaginous ring, supporting the external mouth aper-
ture; g, cirri; My, spinal cord ( myelon ); Ch, notochord (chorda dor-
salis). After Quatrefages and Huxley.

Throughout the length of the body — excepting
the extreme tips of the snout and tail — there run in
Branchiostoma (fig. 365) a spinal cord (My) and a
notochord (Ch), the latter representing a structure pre-
sent in the embryos of all other vertebrates. But in
them the notochord terminates anteriorly under the
swelling of the medullispinal canal, the rudimentary

a Amphioxini, J. Muller, 1. c. Afterwards the family was called Branchiostomidce in Bonaparte, Cat. Met. Pesci Europ. (1846),
pp. 9 and 92.

6 In the Australian Branchiostoma ( Epigonichthys ) cidtellus described by Peters this part of the dermal fold is said to be rudimentary,
the anus of this species being accordingly situated in the median ventral line.

1212

SCANDINAVIAN FISHES.

brain. Here in Branchiostoma there is hardly any such
swelling: — the spinal cord is a string of almost uni-
form thickness, gradually tapering backwards, and only
slightly dilated at its anterior end (fig. 366, a) so as

Fig. 366. Anterior end of the spinal cord and the nerves thereof in
Branchiostoma lanc-eolatum , seen from above and magnified. After
Vogt and Yung.

a. medullispinal swelling and its cavity; b, first, c, second pair of
nerves; d, pigment Spot ( eye spot?); e, third and fourth pairs of
upper (posterior, sensory) nerves; /, spinal cord.

Fig. 367. Caudal end of Branchiostoma lanceolatum, seen from the
left and magnified. After Vogt and Yung.
a, spinal cord; b, notochord; c, upper ramifications of the sensory
nerves; d, lower ramifications of the same; e , hindmost myocomma
(boundary of the hindmost myomere).

to form a kind of cerebral vesicle. And the notochord
tapers to a pointed termination at each end of the body.
A peculiarity never seen in any vertebrate higher than

Branchiostoma, is that the notochord (fig. 365, Ch)
projects forward beyond the anterior end of the spinal
cord {My) and in front of the mouth aperture (/ and g).
At the posterior end of the body the case is different.
There (fig. 367) we find an upcurved dilatation of
the medullispinal canal (a), reminding us of Myxinc
(fig. 354).

The mouth is of a curious structure for a verteb-
rate. It is neither a prehensory nor a suctorial mouth;
it is, so to say, merely a deglutitory mouth, edged
with a ring of small cartilages (fig. 365, /), in texture
resembling the notochord. Each one of these cartilages
gives out a process (g) to form a support in one of
the so-called cirri which surround the oral aperture

Fig. 368. Posterior wall of the mouth cavity (postoral sphincter or
velum ) with its twelve centripetally meeting cirri, which are furnished
with the sensory bodies first described by Langerhans, bundles of
sense-hairs on thickened parts of the epithelium. After Ray Lankester.

outside, but are folded inwards when the mouth is
closed. These cirri are almost exactly similar to the
sense-filaments or tentacles possessed by many of the
lower animals. At their base they are united to a
membrane coursed by muscular fibres; and their sur-
face is clothed with a cylindrical epithelium, among
whose cells some are ciliated, others prolongated out-
wards into stiff setae, which are collected in verruciform
bundles, and connected inwards with nerve fibrillae,
whence they have been interpreted as organs of taste.

Within the mouth is a buccal (pharyngeal) cavity,
the inner surface of which displays active ciliary mo-
tion, especially along the margins of a number of

L EPTOC A RDIANS .

1213

digitiform (process-like) thickenings (tig. 365, vertically
above /); and a constant current of water is thus kept
up into the branchial sac (respiratory cavity, tig. 364, d ).
The mouth cavity is delimited from this sac by a
circlet of cirri (tig. 368), similar to those of the oral
margin.

The walls of the respiratory cavity, which also
serves, however, as a pharynx or rather perhaps an
(Esophagus, are laterally constructed of a network of
cartilaginous rods (tig. 369), closely resembling the
homologous structure in the Ascidians; but above and
below the respiratory cavity has a continuous groove,
the upper known as the epibranchial, the lower as the
hypobranchial groove. The obliquely vertical rods set
transversely in the body are rib-like growths on both
sides, connected above with the outer sheath of the noto-
chord (the so-called skeletogenous layer), the most pri-
mitive rudiment of an axial skeleton. Below (towards
the ventral side of the animal) these rods are free, some
(the so-called primary bars) bifurcated, others (the so-
called secondary bars or tongue-bars) single. The con-
necting rods between them stand in the longitudinal
direction of the body. Thus a network is formed, and
this is faced everywhere, as the two grooves are also
lined, with an epithelium of ciliated cells, exactly si-
milar to those we have just seen in the mouth cavity.
But the meshes of the net are minute, hardly visible
even under a high magnifying power and under ordi-
nary circumstances do not transmit anything but the
clearest, purest water. In the adult state of the animal
this water, which has now served its respiratory pur-
pose, passes out on the sides of the respiratory cavity
and into a. special chamber, a secondary compartment
of the abdominal cavity. This chamber, which has been
named by Ray Lankester" the atrium , surrounding
the respiratory cavity on the sides and underneath and
extending some way behind it, has a special aperture
(the abdominal or atrial pore) behind on the ventral
side (tig. 364, o) for the discharge of the water. Every-
thing else that has accompanied the water into the re-
spiratory cavity, and especially the food — Infusoria
and other minute creatures — finds its way through the
oesophagus proper, a tubiform prolongation of the epi-
branchial groove, into a kind of stomach (fig. 364, just
in front of i) with a long csecal appendage (/<), and all
the useless matter now passes out through the straight

intestine ( i ) and the anal aperture (e). The ctecal ap-
pendage projects forward on the right side into the
atrium, pressed close to the respiratory cavity. The
whole digestive canal, the stomach and intestine, is lined
with an epithelium of ciliated cells and exhibits a vibra-
tile motion, just as the intestinal canal of the worms.

The blood of Branchiostoma is colourless; and the
circulation, of which little is yet known with certainty,
may indeed be referred to the piscine type, for the vascular
system is divided into certain regions which apparently
lend themselves to a comparison with the heart and the
great bloodvessels of fishes; but here too we are re-
minded of the worms. Under the respiratory cavity
runs a long contractile canal with a contractile vesicle
to each ascending rod in the branchial network. This

Fig. 369. A portion of the right side of the branchial basket in
Branchiostoma lanceolatum, seen from without and powerfully magni-
fied. After Vogt and Yung.
a , secondary rods, so-called tongue-bars; h, primary rods.

canal has been compared to the venous heart of fishes.
Another contractile tube, anteriorly double (one on the
right side, one on the left), posteriorly single, runs along
the dorsal margin of the abdominal cavity, under the
notochord. This canal has been compared to the aorta
of fishes. Another similar canal, according to J. Mul-
ler, connects on each side these two canals in front,
and this has been compared to the so-called ductus ar-
teriosus Botalli of the lower vertebrates and of the
higher vertebrates in their embryonic stages. Along the
intestinal canal run two ducts of a venous nature. The

Quart. Journ. Micr. Sc., n. ser., No. LIX (July, 1875), p. 267; No. CXVI (Apr., 1889), p. 365.

1214

SCANDINAVIAN FISHES.

one, which is said to have its roots in the capillaries of
the intestinal canal, runs under the stomach and caecum,
there ramifying into a capillary network between the
green cells in the wall of the caecum, which is said to
functionate as a kind of liver. Here the blood, it is
stated, is again collected, on the upper side of the cae-
cum, in the other, superior canal, whose contractions
run backwards in the opposite direction, to the exit of
the caecum from the stomach, and convey the blood to
the heart: — and we should thus have found here a
portal vein (the inferior canal) and an efferent hepatic
vein (the superior canal). Descriptions have besides
been given of a separate, but wall-less lymphatic system,
tilling all the cavities of the body and ramifying into
the smallest lacume between the organs. But in all the
said contractile canals there appear distinct traits of the
typical arrangement of the vessels in a worm. In those
paired connecting vessels between the upper vessel and

Fig. 370. Anterior end of a Branchiostoma lanceolatum , seen from
the right and magnified. After Vogt and Yung.
a , olfactory depression; b, first, c, second pair of nerves; d , eye spot;
e, peripheral ganglion cells in the skin; /, notochord; g , spinal cord;
h , fin-ray growths (spinous processes?); i , myocomma.

the lower which are most plentiful in the forepart of
the worms ( Annulata ), we may well seek a parallel to
the respiratory system of Branchiostoma. And the two
contractile vessels that run bulging in the latter along
the upper and lower sides of the intestinal canal also
functionate in the worm for the first sucking up of the
alimentary juice and its conversion into nutritive fluid.

The vertebral segmentation of the body is indeed
subindicated even in Branchiostoma by a transversal
division of the membrano-cartilaginous sheath that en-
velops the notochord, expands both around the neural

0 Monatsber. Akad. Wiss. Berlin, 1839, p. 198.
b Mull. Arch. Anat., Physiol., 1843, p. 32.
c Ann. Sc. Nat., ser. 3, Zool., tome IV (1845), p. 226.

canal and the h genial cavity, and breaks up at the base
of the vertical dermal fin-fold along the dorsal and the
posterior part of the ventral margin into pieces resem-
bling fin-rays (fig. 364, a and h ; fig. 370, h) — though
far more numerous than the muscle flakes. But the
muscular and nervous systems are the organs that here
most distinctly call to mind the vertebrate type.

That portion of the muscular system which answers
to the great lateral muscles of fishes is divided into
transverse bands ( myomeres ), which are angular, with
the angle directed forwards; and each myomere is com-
posed of bundles of muscle fibre extended between ten-
dinous expansions of the notochordal sheath ( myocom -
mata), just in the manner of the said piscine muscles.
Along' the ventral side of the abdominal region, between
the mouth and the atrial pore, extends an horizontal
muscular plate, composed of transversal muscle fibres.
Its function, consisting of rhythmical contractions, is
mainly subservient to the respiration, and its posterior
part, at the atrial pore, swells into the so-called abdo-
minal papilla, coursed by muscle fibres in all directions.

The nervous system shows the peculiarity that the
spinal nerves (except the first two pairs) do not. issue
in pairs, exactly opposite each other, but alternately,
the nerves of one side in front of those of the other
(fig. 366, e), and that only the basal parts of the dorsal
branches (sensory nerves) are entire, those of the ven-
tral (motory) nerves being broken up into fibres. In
the spinal cord there appear, as in the preceding order,
besides the ordinary ganglion cells, large so-called co-
lossal cells. Among the organs of sense A. Retzius was
the first" to discover a black pigment spot (fig. 370, d)
on the anterior end of the spinal cord. This spot is
interpreted as an eye of the most rudimentary descrip-
tion. It is comparatively larger in young specimens
than in old, which suggests a still advancing reduction;
and its visual function is somewhat dubious, for similar
spots also occur in a row on each side along the greater
part of the spinal cord. Above the anterior end of the
spinal cord, but on the left side of the body, Kolliker'
and, after him, Quatkefages0 found a ciliated depres-
sion (fig. 370, a), the simplest form of an olfactory
apparatus. The organs of taste we have already re-
marked on the cirri of the mouth aperture and the
postoral velum; and a similar cell structure also appears

L A N C ELET-FISHES .

1215

in the hypobranchial groove. Langerhans described0
as sensory cells a number of cylindrical epithelial cells
scattered in the skin and most numerous on the fore-
most part of the body, furnished distally with a long,
stiff hair, proximally with a process to which the peri-
pheral end of a nerve fibrilla attaches itself. In the
vertical dermal fin around the anterior end of the body
Vogt and Yung remarked6 small, round or oval, trans-
parent ganglion cells, situated oftenest in the fork be-
tween .two nerve twigs (fig. 370, e ).

These organs of sense have indeed scarcely been
submitted as yet to adequate investigation; but so much
is known, that the rudimentary eye has the same struc-
ture as in the lower Polychtetous worms, and the ol-
factory depression also finds its nearest anatomical homo-
logue in Nemertines, Turbellarians, and some other
worms. There is this difference, however, that in
Branchiostoma both these organs are single, and that
the olfactory organ is asymmetrical, situated on the left
side of the body. The only further reason which might
be adduced foi* calling the anterior end of Branchiostoma
a rudimentary head, is that the spinal cord, as men-
tioned above, dilates in front, though but slightly, into
a vesicle from which there originate two fairly sym-
metrical pairs of nerves, the second pair larger and
more numerously ramified than the other spinal nerves.
It is therefore not surprising that some have proposed
to separate tins animal into a distinct class, and have
said: it is neither a vertebrate nor an invertebrate, it
is something between the two.

The organs of generation (fig. 364, c) are a row
of saccate vesicles, free from each other and usually
numbering 26 — 28 on each side in full-grown indivi-
duals, between the abdominal wall and the atrium. The
male and female can be distinguished only by the con-
tents of the vesicles0; and the sexual products are pro-
bably emitted, as a rule, through the atrial pore. Ivo-
walewsky07 and Hatschek0, however, saw these pro-
ducts discharged through the mouth aperture; and one
of the females received by us from Bohuslan had several
ripe eggs in the mouth cavity, where they may easily

arrive, as Hatschek has pointed out, after first being
liberated, by the bursting of the generative sacs, in the
atrium and then passing, through the clefts of the
branchial sac, into this cavity and out at its anterior
(sphincterial) aperture.

As excretory organs in the full-grown Branchio-
stoma Weiss has adduced' a large amount of the epi-
thelial lining of the atrium, especially on two tubiform
prolongations of the said cavity which project into the
abdominal cavity proper or coelom (atrio-coelomic fun-
nels). In this connection he also draws attention to his
own discovery of small, tubular glands (kidneys?) si-
tuated at the top of the branchial tongue-bars (secon-
dary cartilaginous rods) and opening on the outer side
thereof into the atrium.

The character of the genus and its relation to other
animals are best explained, however, by the history of
evolution87. Branchiostoma deposits its ova (fig. 371, A),

Fig. 371. A , an unimpregnated egg, with the so-called polar body
(Richtungskorper, R), a small body liberated from the egg and marking
that pole thereof at which the segmentation is most active. B, an
impregnated egg, with two cleavage-cells not yet quite divided from
each other. C , an egg segmented into 8 parts. D , an egg with 8 large
cells round the inferior pole and 5 sixteen-celled circles higher up. E ,
optical section at the Blastula stage (simple sacculation), with the in-
ferior pole flattened, the commencement of its upward invagination
towards the inside of the wall of the upper pole. F, optical section
of an embryo in the Gastrula stage, with the above-mentioned in-
vagination completed. Each cell of the ectoderm (EM) is furnished
with a flagellum. Ent, entoderm, answering to the lower part of the
Blastula wall in the preceding stage. The Gastrula mouth ( Gm ) is
the downward opening. After Hatschek. X 140.

0 Arch. Mikrosk. Anat., Bd. XII (1875—76), p. 303.
b Lehrb. prakt. verrjl. Anat., Bd. 2, p. 361.

c And also, according to Vogt and Yung (1. c., pp. 375, 376) by the inner epithelial investment of the transverse ventral muscle,
this being developed into a kind of support for the organs of generation (deferent grooves?), different in males and females.
d Mem. Acad. Sc. Petersb., ser. VII, tome XI, No. 4, p. 1.

e Sturl. iib. Entw. d. Amphioxus , Arb. Zool. Inst. Univ. Wien, tom. IV, H. 1, p. 14.
f Quart. Journ. Micr. Sc., n. ser., No. CXXIV (Nov. 1890), p. 489.

^ Kowalewsky, 1. c., and Arch. Mikr. Anat., Bd. XIII (1876 — 77), p. 181; Hatschek, 1. c., Ray Lankester and Willey, Quart.

Journ. Micr. Sc., n. ser., No. CXXIII (Aug. 1890), p. 445.

Scandinavian Fishes.

153

1216

SCANDINAVIAN FISHES.

enveloped in a transparent membrane, free in the water,
where they are impregnated. When one of these eggs
has undergone its total segmentation, it hollows itself
out — it becomes a vesicle or rather an entirely closed
sac (tig. 371, E), the cavity of which answers to the
segmentation cavity (under the blastoderm) in the ova
of the higher vertebrates and, like the said cavity, is
principally obliterated, the sac being flattened ( E ), so
that the lower wall comes nearer to the upper and at
last lays itself close up to the same ( F ). The original
sac has now become a bowl; and the rim hereof is
contracted into a narrower and narrower aperture ( Gm ).
Meanwhile the external layer of the bowl ( Ekt ) has
developed small mobile bristles (flagella) on its cells,
one to each cell (Germ. Geisselcell ): — the egg has at-
tained a stage of development common to most of the
invertebrates and called by Haeckel the gastrula. Now
it dances briskly about within its egg-shell (vitelline

Fig. 372. A', an embryo, prolongated and with the rudiments of the
first proto vertebra} (one on each side of the intestinal canal) seen from
the dorsal side. After Hatschek. X 77. A", transverse section of a
similar embryo. After Kowalewsky. A"’, optical longitudinal section
through the middle of A'. After Hatschek. X 140. B’ , a larva with
two pairs of protovertebrm, seen from the dorsal side. After Hat-
schek. X 77. B ", optical longitudinal section through the middle
of a similar larva. After Hatschek. X 140.

At the stage A (after Hatschek) the embryo quits the egg by
bursting the membrane thereof. The gastrula mouth is still present, but
is overgrown from below (see B") by the ectoderm, which has besides
raised itself on the sides of the back, thus forming a wide groove (n),
the edges of which (dorsal ridges, r) grow more and more together to
form a canal (the future central canal of the spinal cord; cf. nc in fig.
380). In A the first protovertebra ( 1 — on each side a vesicular dilatation
of the entoderm) has begun to differentiate itself, after the appearance
between the first two germinal layers (ectoderm, Ekt and entoderm,
Eiit) of the middle (third) germinal layer (mesoderm, fig. A", con-
sisting of the animal (a) and the vegetative ( v ) lamina). According
to Hatschek the mesoderm is developed from the entoderm, in the
hindmost part of which two cells, larger than the rest, indicate the
posterior limit of the mesodermal growth and have therefore been
named the polar cells of the mesoderm (mp). Simultaneously with
the formation of the mesoderm and its division into an outer (animal)

and an inner (vegetative) lamina, a body-cavity (coelom, cl) has arisen
round the intestinal canal. The lumen (internal space) of the latter
is indicated by i. In B the rudiments of two pairs of protovertebree
(/ and 2) have appeared.

membrane). The two layers of which the body of this
gastrula is composed correspond to the first two ger-
minal layers of the higher vertebrates. The structure
is the same, though the manner of life is widely dif-
ferent, for this gastrula has no vitellus on which to
rest, or from which to derive its nourishment. On the
other hand, it has a stomach, which it has acquired by
the invagination of the lower (eventually inner) part of
its wall ( Ent ). In the embryos of the highest verteb-

rates the intestinal canal indeed has quite a different
appearance — though fundanentally constructed on the
same principle — and is not formed until after a con-
siderable alteration of the original germ; but in the
batrachians the development of the said canal calls to
mind the above-mentioned invagination, although the

Fig. 373. Two optical longitudinal sections of a larva with 9 proto-
vertebrge (somites, 1 — 9 ), seen from the left {A) and from the dorsal
side ( B ). After Hatschek. X 140. The cerebrospinal canal (n) has
now been formed and is separated from the ectoderm, but in front
(at hy) open and behind (through the original gastrula mouth, Gin )
continuous with the intestinal canal (i). This has dilated anteriorly
to the right ( dvd ) and the left ( dvs ). Under the cerebrospinal canal
the notochord ( ch ) has appeared, and in the protovertebrm muscle cells
begin to develop. In B it appears that the protovertebrae are set
obliquely opposite each other, the right a little further back than the
left; but the anterior lateral dilatations of the intestinal canal are
still of equal size (symmetrical).

batrachian embryo, like the mammalian, has been sup-
plied by the transformations of the egg with a kind
of nutritive yolk under it.

The gastrula of the future Branchiostoma elongates
its form of body (fig. 372), thus acquiring an anterior
and a posterior end. In the latter lies its mouth. Soon
it acquires a dorsal and a ventral side, for on the
former a primitive groove appears, which becomes a
cerebrospinal canal, following the same manner of deve-

LANCELET-FISHES.

1217

(gl) on the right side of the intestinal canal becomes
a gland* * 6 which also disappears at the close of the larval
period, and whose outer efferent duct bends round be-
low to the left side of the body (fig. 375). But in the
median ventral line the intestinal canal coalesces at one
point after another, backwards from in front, with the
wall of the body, becomes thickened there (fig. 374, at I),
and eventually opens into gill-slits (fig. 375), which are
first moved up to the right, but afterwards migrate to
the left side of the body. In the meantime, however,
new gill-slits (fig. 378) have been similarly formed in

Fig. 374. Optical section of a larva with its protovertebrse, seen from the right. After Hatschek. X 280.

Here the lateral dilatations of the foremost part of the intestinal canal have diverged widely; the right ( dvd ) is elongate and thin-walled,
the left (dvs, which from that side shines through in the figure, at about the middle of the former) has thicker walls (deeper cells) and is
round. Both separate by constriction from the intestinal canal, the anterior end of which is thereby thrust further back. A glandular growth
(the so-called club-shaped gland of Hatschek, gl) has besides begun to develop on the inside of the intestinal canal by means of a trans-
versal constriction, first canaliculate and afterwards, owing to the coalescence of the edges of the groove, tubiform, of the right side of the in-
testinal wall behind the said lateral dilatation. Behind this gland the wall of the intestinal canal has been thickened in the inferior (ventral)
median line of the larva and has coalesced with the ectoderm at a. point (I) where the first gill-opening afterwards appears. As yet there

is no true mouth; and the intestinal canal is continuous behind with the cerebrospinal.

ch oc cil m

Fig. 375. A larva l1/2 mm. long, seen from the left. After Ray Lankestek and Willey.

Of the four gill-slits, which are situated on the right side (I — IV), the foremost three are rather large, the fourth is rudimentary. The
first is visible through the posterior part of the semioval mouth aperture, which belongs to the left side of the body. Through the anterior
part of this aperture are visible, also from the right side of the body, both the club-shaped gland (gl), the outer orifice of which may be
seen below, on the wall of the body below the mouth aperture, and, just in front of the said gland, a transversal, but curved thickening (end),
bisected along the middle by a groove, the future endostyle. Before the upper edge of the mouth aperture appears the opening of the funnel-
shaped, ciliated organ (cil). Above this and from the anterior to the posterior end of the body (in front of the caudal fin) lies the notochord
(ch), divided into its numerous transverse disks. Above the notochord, and posteriorly round the termination thereof, the spinal cord (m) is
extended, with the large pigment spot (ocular rudiment, oc) at its anterior end and some smaller sparse pigment spots on its sides. In the
lanceolate caudal fin a sparse collection of pigment spots is also visible. The number of the myomeres is 36; the first myocomma runs
obliquely across the anterior edge of the ocular rudiment and down along the subjacent part of the notochord. In the posterior half of the
intestinal canal (below the 15th and 16th myomeres) the wall of this canal is thickened (v), an indication of the rudimentary stomach.

The anal aperture (an) lies on the left side of the body.

lopment as in the higher vertebrates". Between the
first two germinal layers an intermediate layer (the
mesoderm. , fig. 372, A') is formed; and now that the
intestinal canal and an incipient abdominal cavity have
begun to differentiate themselves, there develop a noto-
chord and, by means of bulgings and constrictions of
the intestinal canal, primitive vertebrae ( protovertebrce ),
which increase their number backwards from in front.
The foremost bulges (figs. 373 and 374) meet with a
singular fate. The right (dvd) dwindles into the hol-
low at the tip of the snout. The left, on the other

hand, develops into a funnel-shaped, ciliated organ,
probably an organ of smell, which disappears, however,
at the termination of the larval period. A special bulge

a row higher up on the right side, and these become
the permanent gill-slits on that side. This only ap-
plies, however, to the anterior part of the branchial

a Branchiostoma differs, however, from the higher vertebrates in that the rudiment of the spinal cord is separated from the circum-

jacent cells of the ectoderm before the canal is closed above (Hatschek).

6 See Willey, Quart. Journ. Micr. Sc., n. ser., vol. XXXII, pt 2 (No. CXXVI, March 1891), p. 209.

1218

SCANDINAVIAN FISHES.

wards and forwards over the tops of the branchial slits.
The latter {end, with its supporting rods, the so-called
endostyle , with a right and a left part), on the other
hand, follows, according to Willey, an opposite direction
of development, backwards from in front, originating in
a thickening, at first transversal (figs. 375 — 379, end),
of the intestinal wall beside the club-shaped gland.
The first rudiment of the atrium appears (fig 380, A,
sar ) where the atriopore afterwards has its place, and
presents the appearance of two horizontal ridges, grow-

m

VII

Fig. 376. A larva 31/., mm. long, taken swimming freely in the sea, between 1 and 4 fathoms below the surface, at Kristineberg (Bohus-
lan), August 17, 1894. Seen from the left. Letters and numerals as in the preceding figure; besides: am, the ciliated mouth of the cerebro-
spinal canal; ep, the forward growth on the inside of the intestine, a fold originating from the inferior wall of the intestinal canal. Number
of the myomeres here 61. Gill-slits and their rudiments visible here to a number of 13 (7 — XIII).

basket. Some of the first-formed gill-openings close
again; and the posterior part of the branchial basket
has its slits symmetrically arranged after the termina-
tion of the larval period. The mouth appears on the
left side, first as a fine slit, afterwards as a large, el-
liptical aperture with tumid margin (fig. 375). A true
anus opens, also, as a rule, on the left side, and the
original gastrula mouth grows together, the connexion
between the intestinal canal and the cerebrospinal canal
being thus cut oft’. As yet the larva moves only with

ch

imt !

the aid of the dermal cilia; but it has begun to bend
its body, since the rostral and caudal fins have become
sharpened. Over the gill -slits there grows from above
(fig. 377) a longitudinal fold from the walls of the
body (somatopleures, the outer parts of which grow
into the so-called metapleures or abdominal fins, met,

Fig. 377. Forepart of a larva 3 1 /2 mm. loug, seen from the right.
After Ray Lankester and Willey. Here it appears how the wall of
the body from the ninth gill-slit forward lies like a leaf outside the
upper parts of the slits, and further forward is continued in a sharp,
free edge, forming an upward curve on the right side of the head.

Letters and numerals as in the preceding figure.

figs. 379 and 380) on each side, first on the right;
and between these folds the atrium is formed, first only
on the ventral side and growing forwards from behind.
The construction of the branchial basket is continued
(figs. 376, 378, and 379) by the addition of epibran-
chial and hypobranchial grooves. The former {ep) ori-
ginates, so far as we have been able to see, as a fold-
like incurvature, advancing in its development forward
from behind along the lower wall of the intestinal canal,
which at the hindmost rudimentary gill-slit grows up-

ing to meet each other (fig. 380, B), on the inside of
the above-mentioned dermal folds {met), whose inferior
margin subsequently becomes on each side of the body
a longitudinal lateral ridge (abdominal fin or meta-
pleure). Afterwards the atrium expands upwards, en-
croaching upon the region of the abdominal cavity on

fr VII- !'■ ’■ cil fr

Fig. 378. Anterior end of a somewhat larger larva, seen from the
right. After Willey. Here a series of seven secondary gill-slits
(/- — VII') has appeared on the right side, above the inferior ends
of the primary slits (7 — XIII), which have been translated for the
most part to the left side, and among which the thirteenth (hindmost)
has been obliterated, and the first has almost suffered the same fate.
The endostyle (end) has grown backwards, and its posterior extremity
extends behind the club-shaped gland (gl). Further forward the velum
of the mouth cavity ( vel ) has begun to develop. In the skeletogenous
layer (chordal sheath) a series of so-called fin-rays (fr) has appeared.
On the atrial floor two so-called renal organs (r) may be seen. Letters
and numerals otherwise as in the preceding figure.

the sides of the branchial basket, and continued within
the abdominal cavity even behind the atriopore, beside
the intestine.

The details of this development should be easy of
comprehension with the aid of the figures which we

LAN CELET-FI SUES .

1219

give here and their explanations. In the last stages
the external form of the body is almost complete. But
it still remains for the month to assume its ventral
position and its wreath of tentacles instead of the long
vibratile hairs (not included in our figures) which are
set during the larval period on a verruciform thicken-
ing just in front of the mouth-aperture.

Br and \io stoma besides teaches us the appearance
presented by the locomotive organs of the vertebrates
in their simplest form. The locomotive organ proper
is the hind part of the body, edged above and below
by a kind of caudal fin or rather a compound of rudi-
ments of the posterior part of a, dorsal fin, a caudal
bn, and an anal bn. Before the anus this bn-growth

Fig. 379. A larva 5 mm. long, taken together with the original of fig. 376. From a preparation briefiy treated with a weak solution of
osmium bichromate by Prof. G. Retzios and preserved in glycerine. A, from the right; B, from the left. Here the above-mentioned fold (ep)
originating in the inferior wall of the intestinal canal to form the epibranchial (hyperbranchial) groove has divided into two parts, the one
(ep') following the tops of the branchial arches, the other (ep") taking its course up towards the roof of the abdominal cavity. In B the
larval mouth shows the commencement of its alteration and translation from the left side of the body to the ventral margin. A derma] fold
(or) has laid itself over the upper part thereof; and the superior margin shows signs of dissolution, atrp, atriopore; metcl , edge of right,
mets, of left metapleure; n), first pair of nerves, with a few ganglion cells (cf. fig. 370, e) visible; other letters as in the preceding figure.

Fig. 380. Diagrammatical transverse section, A, through a larva of Branchiostoma lanceolatum , with 11 or 12 primary gill-slits, and B,
through a full-grown specimen of the same species. Enlarged. After Ray Lankester and Willey.
dm, myoccel of the dorsal fin (that part of the body-cavity (coelom) which belongs to the dorsal fin-lobe); fr, series of fin-rays; m , great
lateral muscles; my, muscle coelom, myocoel (that part of the body-cavity which belongs to the lateral muscles); nc, spinal cord, with its
central canal; nch, notochord; d. a., dorsal aorta; v. a., ventral vessel; hit, intestine; sp, splanchnoeoel (intestinal part of the abdominal ca-
vity), which (in B) is divided by the ingrowth of the atrial chamber into an inner (sp') and an outer (sp") cavity; sp'" (in B), perigonadial
dilatation of the splanchnoeoel; d. 1. m ., double-layered membrane separating the myocoel from the splanchnoeoel; at, atrium; met , metapleure, one on
each side of the body, its cavity, according to Ray Lankester, not in open communication with the coelom; s. a. r., floor of the atrial chamber.

1220

SCANDINAVIAN FISHES.

advances on the ventral margin forward to the atrio-
pore, but there it has a double series of supporting
cartilages (fin-rays), instead of the single row in the
other parts of the vertical fin. In front of the atrio-
pore are the two lateral ridges (metapleures) answering
to the first rudiment of or at least foreshadowing la-
teral extremities, whether they represent both extremital
pairs of the other vertebrates, being not yet differen-
tiated into an anterior and a posterior pair, or correspond
to the pectoral fins alone, in which case the double
series of rays in the atrio-anal fin would appear to have
some homology with ventral fins. But as yet they are
merely steering organs. This is a function, however,
which is for the most part retained by the unpaired fins
of the back and belly, and indeed by the lateral fins, even
in the highest evolutional grades of the piscine type.

The structure and development of Branchiostoma
have now shown us a creature with the vertebrate di-
vision of the body into a neural and a haemal canal,
with a notochord and the membranous elements of an
axial skeleton between and around those canals, and

with the vertebrate type expressed in musculature,
spinal nerves, respiratory, excretory, and generative
organs. But in other respects we have seen an asym-
metry, partly persistent, partly temporary, which is
distinctly foreign to the vertebrates, but common among
the mollusks; the respiratory organs present a striking
resemblance to those of the Ascidians; and the most
important organs of nutrition and circulation, though
comparable with the vertebrate type, retain much of
their structure in the worms. Hence the great morpho-
logical interest attached to this genus, which in other
respects is not especially remarkable. Its forms are so
similar that in his famous Catalogue Gunther recog-
nised only a single species. Sundevall, however, had
already (1852 — 53) distinguished between three species,
with characters derived from the form of body and the
number of myomeres. At a more recent date Gunther
has decided, basing his conclusions in the main on the
same characters, that six different species deserve re-
cognition within the genus. Only one of these belongs
to the seacoasts of Europe.

THE LANCELET (sw. lansettfisken).
BRANCHIOSTOMA LANCEOLATUM.
Plate L1II, fig. 6.

Greatest- depth of the body about 9 — 10ll2 %a, greatest breadth ( thickness ) thereof about 6 — 8 %b , of the length
of the same. Distance from, the tip of the snout to the posterior limit of the mouth aperture about 8 %e, to the
atriopore about 70 %d , to the vent about 88 %\ of the length of the body. Number of myomeres 60 — 62,

11 — 13 situated behind the vent.

Syn. Limax lanceolatus, Pall., Spicil. Zool., fasc. X, p. 19, tab.

I, fig. 11; ( lanceolaris ), Naturg. Merkw. Th., Samml. X,
p. 24, tab. I, fig. 11; Yakr. (Amphioxus lanceolatus ), Brit.
Fish., ed. 1, vol. II, p. 468; Fr., Vet. Akad. Handl. 1838,
p. 336, tab. 4, fig. 3; Sundev., F., Fork. Skand. Naturf.
Kbhvn 1840, p. 280; Sundev., C. J., Arsber. Zool. Framst.
1840 — 42, Vertebr., p. 287; (Avers. Vet. Akad. Fork. 1852,
p. 148; 1853 ( Branchiostoma ), p. 12; Quatref., Ann. Sc.
Nat., ser. 3, Zool., torn. IV, p. 197, tabb. 10 — 13; Kr.,
Damn. Fisk., vol. Ill, p. 1087; Nilss,, Skand. Fna, Fisk.,
p. 753; Couch ( Amphioxus ), Fish. Brit. Isl., vol. IV, p.
415, tab. CCXLVIII, fig. 4; Gthr ( Branchiostoma ), Cat.
Brit. Mus., Fish., vol. VIII, p. 513; Rep. Zool. Coll. Alert
(Brit. Mus. Nat. Hist. 1884), p. 32; Coll., Fork. Vid. Sels.

Ckrnia 1874, Tillsegsk., p. 222; N. Mag. Naturv., Bd 29
(1884), p. 123; Malm, Gbgs, Boh. Fna, p. 641; Winth.,
Naturli. Tidskr. Kblivn, ser. 3, vol. XII, p. 62; Mor., Hist.
Nat. Poiss. Fr., torn. Ill, p. 618; Storm, N. Vid. Sels.
Skr. Trondhj. 1883, p. 48; Day, Fish. Gt. Brit., Irel., vol.
II, p. 366, tab. CLXXIX, fig. 4; Ptrn, Vid. Meddel. Naturli.
For. Kbhvn 1884—86, p. 160; Retz., G. ( Amphioxus ),
Biol. Unters., N. F., vol. II, p. 29, tabb. XI— XIV; Car.
( Branchiostoma ), Prodr. Fnce Medit., vol. II, p. 498.
Branchiostoma luhricum, Costa, Cenni Zool. (1834), p. 49;
Fna Regn. Nap., Pesci, part. II, tav. XXX; J. MOll., Abk.
Akad. Wiss. Berk 1842, Pkysik. Abh., p. 79.
Gasterobranchus glutinosusf, Rasch, Mag. Naturv. Chrnia, Bd
12 (1836), p. 325.

" 8'9 — 10‘6 % according to our measurements of specimens 43 — 48 mm. long.

6 5'8 — 8‘3 % according to our measurements.

7 ' 5 8’8 % ,, ,, ,, ,,

67 —71 % „ „ „

' 86-5—90 % „ „ „

11

LANOELET.

1221

The Lancelet attains in Scandinavia a length of
about V2 dm. Lilljeborg’s largest specimens were 52
mm. long. According to Carus it sometimes mea-
sures at least 70 or even 100 mm.a During life the
body is of a jelly-like transparency; through the skin
and muscles may be perceived the light green notochord,
the somewhat darker c tecum, and the stomach with
intestine, the last-mentioned organs more or less dark
according to the nature of their contents. In a certain
light the skin appears somewhat iridescent, and a hand-
some appearance, as of a row of white, transparent
rings, is presented by the plate- like rudimentary fin-
rays. When the seminal or ovarial sacs are full, they
are perceivable on each side as a series of white or
green globules, the green egg-sacs each with a dark dot.

The habits of the Lancelet have been sketched
above. Its breeding-time lasts throughout the summer;
but in wet and chilly weather it does not spawn.
Theel obtained at the beginning of July a large number
of specimens which were placed in the aquaria of Kris-
tineberg and laid eggs that developed normally. At
Haugesund in Norway Lilljeborg received at the be-
ginning of August specimens not yet quite ripe. On
the west coast of France and in the Mediterranean the
Lancelet commences spawning in March. According to
Hatschek, as well as Ray Lankester and Willey,
who made their observations in the well-known lagoon
of Pontano, near Messina, the spawning always takes
place in the evening, segmentation commencing between
7 and 8 p. m. At 11, say the latter writers, the
gastrula begins to develop, and at 1 a. m. it is com-
plete. At 3 a. m. it begins to revolve by cilia within
the egg-membrane, and at 5 a. m. it already has two
pairs of myocoelomic pouches, when it bursts the egg-
membrane and becomes free-swimming. Thirty-six
hours after the commencement of segmentation the
embryo has acquired a mouth and a rudimentary gill-
slit. Soon afterwards the anus opens, and the larval
period now begins. The larva; swim both at the sur-
face, where Malm found them off Gasd Island (Bohus-
]&n), and deeper down, at a depth of 15 — 20 fthrns.,
according to Ray Lankester and Willey. Here they
are found in countless myriads together with other
minute marine creatures ( Noctilucce , Beroe, Sagittce,

Calani ) and their larvae, amongst that mass (Hensen’s
plankton ) which the Norwegians call Aatb. The deve-
lopment of the larva progresses much more slowly than
that of the embryo. A fortnight is said to elapse be-
fore the formation of the second gill-cleft; and the
remainder of the development, until the atrium is
complete, and the Lancelet begins to bury itself in
sand or mud, is said to take months (Hatschek). The
course of the development is besides very irregular,
and it may be so protracted, at least in the North,
that Dr. C. W. Aurivillius found larvae 5 mm. long
swimming freely at a depth of 15 — 30 m. below the
surface of Gullmar Fjord, in the middle of November.

In its hiding-place at the bottom the Lancelet
leads the life of the worms, and to escape danger it
buries itself deeper down, for Malm remarks that, un-
less the dredge — the instrument most generally em-
ployed in the capture of marine animals at the bottom
— takes a sufficient hold in the gravel, sand, or mud,
there is little hope, even in places known to be fre-
quented by the Lancelet, of securing a single specimen.
Few fishes are so tenacious of life; and even if the
body be cut into pieces, the ciliary motion does not
cease at once in the severed parts. In salt water
changed from time to time, it may be kept alive for
months; and Lilljeborg relates that a number of Lan-
celets which he had dredged up at Haugesund on the
2nd of August, in a quantity of shellsand, and thrown
on the dry rock, were alive three days afterwards,
though the moisture of the sand was not very great,
and though rain had fallen in the interval. In spite
of the sluggish life it usually leads, the Lancelet is
quick of movement when disturbed, wriggling about
in the aquarium like a young Eel, but with either end
foremost. Its endurance is not great, however; if the
observer should desire to induce the captive to lie still
for examination, it may easily be tired out, and it
will then suffer itself without more ado to be conveyed
where a closer view may be obtained, provided only
that it has no soft bottom in which to hide itself. In
the day-time it lies motionless. At night it moves
about; but if a light be brought near the aquarium,
it hastens to conceal itself. As human food it is worth-
less; but it has enemies, no doubt, in plenty.

“ Carus, however, combines this species with the Pacific Branchiostoma , and his account of Lancelots 1 dm. long may perhaps de-
pend on that fact.

b See above, pp. 970, 971.

c Bih. Vet. Akad. Hand!., Bd 20, Afd. IV, No. 3, p. 11.

1222

SCANDINAVIAN FISHES.

The Lancelet was first described in 1774 by Pallas,
who had received a specimen from the Cornish coast.
He took the ventral side between the metapleures to
be a creeping-disk, and therefore referred the animal
to the genus Umax. His figure was copied in PI. 80,
among the vers mollusques , in the Encyclopedic Metlio-
dique; but the Lancelet remained otherwise unknown
for nearly sixty years. The species was then redis-
covered at no great intervals of time in four localities,
and was shown to be a fish. Couch found it in 1831
on the shore near Polperro, Rasch in 1833 on the
Norwegian coast, Costa in 1834 at Naples, and Fr.
Sundevall and S. Loven in the same year on the
Weather Islands (Bohuslan). In 1838 our figure was
painted by v. Wright for Fries, with a view to a
description which death prevented the latter from com-
pleting. The specimens collected by Fries, however,
came into the hands of C. J. Sundevall, his successor
at the Royal Museum, and Anders Retzius, and sug-
gested the first scientific examination of the fish, an
undertaking which was carried out on the coast of
Bohuslan in Retzius’s company by Johannes Muller.
In later times the most important contributions to our
knowledge of the Lancelet have come from Messina
and the Zoological Station at Naples. On the west
coast of France and the English coast the species is
also common enough in suitable localities, on a bottom
of sand and gravel, preferably shell-sand, from the
tide-mark to a depth of some tens of fathoms. In
Scandinavian waters it is dispersed from Trondhjem

Fjord (Storm) and along Southern Norway into the
Cattegat, where it is commonest, according to Wintiier,
in 5 — 9 fathoms of water, but has also been found at
a depth of 1 7 1/s fathoms. The southern limit of its
known range in Scandinavia is the north of the Sound,
at Hellebiek in Zealand (Lutken) and on both sides
of Samso, between Zealand and Jutland. On the coast
of Bohuslan it is plentiful, according to Malm and
Theel, in the tine shell-sand of the eastern harbour
on Storo, one of the Weather Islands, where it was
found first by Loven and Sundevall. Tiieel found
numerous specimens of the Lancelet on the skerry of
Bouden, and Malm a few on Flatholm, several on Gaso
(all three localities at the entrances of Gullmar Fjord),
and a few on Paternoster Skerry, north of Marstrand.
The specimens collected on Bonden by Theel at the
end of June, 1894, were kept for some days at Kris-
tineberg in a glass bowl with tine sand at the bottom,
before being sent alive to the Royal Museum. In the
meantime (till the 30th of June) they had deposited
quantities of eggs, which had already attained or passed
the gastrula stage. Some days afterwards larvae 1 1/s
mm. long were taken in the aquarium, their stage of
development being that shown above in tig. 375. After-
wards numerous larvae 372 — 5 mm. long were caught,
at the stages represented in our figures 376 and 379,
swimming freely in the sea at Kristineberg, close up
to the jetties on the shore, at a depth of 1 — 4 fthms.
below the surface. These larvae too were forwarded
without difficulty alive to the Royal Museum.

INDEX.

Aalekone, Aalekuse, Aalemoder, Aalmutter, 607.
Aat, 970.

Abborre, 26.

abbreviatus (Cyprinus), 733.

Abdominales, Acanthopterygii, 24.
Abdominales, Malacopterygii, 689.
Abramidinae, 720, 790.

Abramidopsis, 816, 817.

Abramis, 720, 790, 796, 802, 812.
abramo-rutilus (Abramis, Bliccopsis), 808, 809.
abraptns (Leucaspius), 787.
abyssoruin (Gadus, Lota, Molva), 521.
acadianus (Glyptocephalus), 379.

Acanthias, 1158.

acanthias (Acanthorhimis, Canis, Galeus, Spinax,
Squalus), 983, 1079, 1157, 1158.
acanthias (Galeus, seir spinax fuscus), 1163.
Acantkocottus, 168, 169.

Acanthocybium, 90, 116.

Acantholabrus, 4, 5.

Acanthopsides, 703.

Acauthopsis, 706.
acanthopteri, Heterocormi, 24.

Acanthopterygii, 2.

Acanthopterygii centrisciformes, 635.
Acanthopterygii gasterosteiformes 635.
Acanthorhinus, 1156, 1166, 11(57.
Acanthuridae, 620, 631.

Acerina, 25, 40.

Acipenser, 1054, 1056.

Acipenseridse, 1044.

Acrania, Yertebrata, 1210.

Acrochilus, 716.

acronius (Coregonus), 899, 902.

aculeata (Cobites), 706.

Aculeatus, 638.
acnleatus (Balistes), 621.

aculeatus (Clinus, Leptoclinus, Stichaeus, Lum-
penus), 228.

aculeatus (Etmopterus), 1163.
aculeatus (Gasteraeanthus, Gasterosteus), 636,
644, 645, 647, 648, 660.
acirleatus (Orthagoriscus), 626.
aculeatus (Rhombus), 398, 434.
acuminatus (Cyprinus), 724.

Acus, 347, 680, 686.
acus (Beloue), 343, 347.
acus (Fierasfer), 598.

acus (Siphonostoma, Siphostoma, Syngnathus),
664, 666, 668, 672, 674, 675.
acuta (Perea), 27.

acutirostris (Anguilla), 1023, 1024.
acutus (Alburnus), 792.
acutus (Exocoetus) 355.

Adelviscb, 909.

^Eglefinus, 466.

aeglefinus (Gadus, Melanogrammus, Merlangus,
Morrhua), 465, 466, 467, 484.
segyptiaca (Anguilla), 1023.
iEliani (Thymallus), 884.
sequoreus (Acus, Entelurus, Nerophis, Syn-
gnathus), 666, 680, 681.
affinis (Beryx), 68.
affinis (Cottus), 170.
affinis (Thynnus), 93.

Agassizii (Brama), 81.

Agassizii (Callionymus), 271.
agilis (Gadus) 484.

Agon 984.

Agonidee, 126, 202.

Agonopsis, 204.

Agonostoma, 328.

Agonus, 147, 204.

Aigle, 468.

Aiglefin, 476, 468.

Aigrefin, 466.

Ailiaeformes, 692.

Akajei (Trygon), 1099.
alalonga (Thynnus), 91.

Alandblecke, 798.
alascanus (Ammodytes), 577.

Alausa, 984.

alba (Clupea, Rogenia), 954.
alba (Raja) 1117.
albicora (Thynnus) 91.
albidus (Gadus), 540.
albiensis (Leuciscus), 770.

Albicore, 97.

Albula, 792, 898.

albula (Argyrosomus, Coregonus, Salmo), 891,
893, 895, 896.

alburniformis (Bliccopsis), 796.
alburniformis (Scardiniopsis), 795.
alburnoides (Leuciscus), 792.

Alburnus, 720, 786, 790, 791.
alburnus (Abramis, Aspius, Cyprinus, Leucis-
cus), 792.

Alburnus lucidus, 33.

alburnus (Menticirrus, Umbrina), 120.

albus (Coregonus), 899.

albus (Gobius, Latrunculus), 266.

albus (Merlangus), 511.

albus (Squalius), 770.

alepidotus (Cyprinus carpio), 724.

Alepocephalus 997.

algeriensis (Batrachus), 135.

algeriensis (Gasterosteus), 648.

alipes (Salmo), 843, 844.

Alkufva. Alkula, 757.

Alkutta, 757.

Allice Shad, 985.

allitteratus (Euthynnus, Scomber), 91, 93.
Alopecias, 1137.

Alopias, 1136.

Alosa, 952, 978.

Alosa vera auctorum, 985.

alosa (Clupea), 952, 983—985, 987.

Alose, 984.

alpinus (Salmo), 830, 831, 835, 842, 851,
867

Alse, 952.

altirostris (Anguilla) 1023.
altivelis (Trachypterus), 314.
amarus (Rhodeus), 715, 720.
americana (Morone, Roccus), 44.
americana (Morrhua), 473.
americana (Perea), 27.

americanum (Amphiprion, Polyprion), 47, 48.
americanus (Acanthias), 1158.
americanus (Ammodytes), 577.
americanus (Brosmius), 562.
americanus (Limanda, Pleuronectes), 362, 363,
390.

americanus (Petromyzon), 1183.
amethystino-punctatus (Maurolicus), 932 — 935.
Amia, 718, 946, 1012.

Ammocoetes, 1172.

Ammodytes, 461, 569.

Ammodytes Anglorum verus, 569.
Ammodytidae, 462, 557, 567.

Ammodytini, 567.
amphibius (Salarias), 212.

Amphioxidas, 1211.

Ainphioxus, 1220.

Amphiprion, 48.

Anacanthini, 371, 461, 567.

Anacanthini gadoidei, 461.
anadromus (Capito), 799.

Anarrhichadidse , 126, 231.

Anarrhichas, 137, 21 1, 231, 602
Anchois, 876.

Anchovus, 876.

Anchovy, European, 990, 992.
ancidda (Anguilla), 1023.

Ancylodon, 51.

Angelkrok, 1007.

Anglepiga, 390.

Angler, 138.

anglorum (Lumpus), 294.

Angmarset, 876.

Anguilla 1022.

anguilla (Murasna, Ophichtlius), 1023.
anguillaris (Enchelyopus), 605.

Anguillidse, 1022.

anguineus (Chlamydoselache), 1063.

Scandinavian Fishes.

154

1224

anguineus (Entelurus, Nerophis, Syngnathus),
680, 681.

angulosus (Balistes), 634.
anhypopteriens, Squales, 1156.

Ankarvad, 1035.
annectus (Coregonus), 899.
annulatum (Scyllium, Squalus), 1149.

Anomali, 24.

anomalopteri, Acanthopterygii, 24, 126.

Anon, 466.

Antaceus, 1054, 1055.
antarctica (Sciaena), 51.

Antennarii, 136, 144.

Antennarius, 144.
antennatus (Diodon), 620.

Anticitharus, 426.

Antimora, 539, 581.
antiquorum (Hippocampus), 662.

Apeltes, 649.

Aper (Labrus), 7.

Apfisk, 1079.

Aphya, 242, 264, 787, 976.

Apliya cobites, 262, 264.

aphya (Cyprinus, Phoxinus), 33, 720, 721, 754.

aphya (Gobius), 251, 262.

Aphyonus, 462.

Apodes, 461, 567, 595.

appendix (Petromyzon), 1183, 1187.

aquila (Cheilodipterus, Sciaena), 50, 51.

Aquila marina, 1095.

aquila (Myliobatis, Raja), 1094, 1095.

aquila (Scifena), 975.

araneus (Trachinus), 128.

Archagonus, 206.
arctica (ChimaBra), 1080.
arctica (Liparis), 287.
arcticus (Chironectes), 146.
arcticus (Galeocerdo, Squalus), 1129.
arcticus (Gymnetrus, Gynmogaster, Trachy-
pterus), 315.

arcticus (Mallotus, Osmerus, Salmo), 876.
arcticus (Thymallus), 883.
arctique (Chimere), 1079.
argentatissimus (Gasterosteus), 648.
argentatus (Merlucius), 514, 515.
argentea (Chimtera), 1079.
argentea (Mursena), 1023.
argenteolirs (Gadus, Motella, Couchia), 550,
553, 554, 556.
argenteus (Cyprinus), 733.
argenteus (Fario), 851.
argenteus (Gadiculus, Gadus), 464.
argenteus (Leuciscus), 760.
argenteus (Petromyzon), 1189.

Argentina, 827, 828, 889, 912, 917.
argentinum (Goniosoma), 918.
argenti-vittatus (Thynnus), 91.
argyreus (Abramis), 812.
argyroleuca (Blicca), 803.

Argyropelecus, 924.
argyropomus (Gasterosteus) 648.

Argyrosomus, 892.

Aristotelis (Canicula), 1154.

Arius, 692.

armatus (Aspidophorus), 208.
armatus (Caranx), 85.

Arnoglossus, 363, 426, 427, 428.
arnoglossus (Pleuronectes, Rhombus), 428,
429.

Artedi (Pristiurus, Scyllium), 1149.

Ascanii (Bletmiops, Blennius, Carelophus), 218.
Ascanii (Gymnetrus), 322.

Ascanii (Silus), 914.

Asellus, 472, 518.

Asp, 720, 783.

asper (Limauda, Pleuronectes), 390.
aspera (Dasybatis), 1104.
aspera (Lepidotrigla), 195.
aspera (Raja), 1115.

Asperi (Coregonus), 899.

Aspicottus, 187.

Aspidophoroides, 203.

Aspidophorus, 204.

Aspius, 720, 752, 782, 786.
aspius (Abramis, Aspius, Cyprinus, Leuciscus),
783.

aspius (Coregonus), 891, 905, 907.

Asp redo, 692.

Asterospondy li , 1128.

Astronesthes, 921.

Atheresthes, 408.

Atherina, 265.

Atherinidse, 327.

Atkinsii (Gasterosteus), 648.
atlanticus (Macrurus), 586.
atlanticus (Sudis), 827, 920, 923, 946.
atrovirens (Cyprinus), 724.

Attilus, 1056.
aula (Leuciscus), 777.

Aulopyge, 715.

Aurata, 41.
aurata (Sparus), 53.
aurata (Tinea chrysitis), 748.
auratus (Carassius, Cyprinopsis, Cyprinus), 723,
733.

auratus (Mugil), 333, 337.
aurea, var. (Perea fluviatilis), 30.
aureus (Ammocoetes, Petromyzon), 1189, 1193.
aurita (Clupea), 952, 978.

Ausonii (Salmo), 851.
australe (Amphiprion), 48.
australis (Brama), 77.
australis (Maurolicus), 932, 935.
australis (Zeus), 306, 309.
autumnalis (Clupea harengus), 959.

Auxis, 99, 107.

Ayresii (Petromyzon), 1189.

Backeljauue, Backeljo, 476.

Badfisk, 776.

Bagrus, 691.

bahiensis (Exocoetus), 358.
bahusica (Clupea harengus), 959.
baicalensis (Thymallus Grubei), 882.

Bailloni (Gasterosteus), 648.

Baingyl, Bainkyl, 657.

Bairdii (Bathymyzon), 1182.

Baldneri (Leuciscus), 797.

Balistes, 2, 137, 632, 633.

Balistidae, 631.

Balistina, 622, 631.

Ballan (Wrasse), 7.

Ballerus, 802, 804.

ballerus (Abramis, Cyprinus, Leuciscus), 720,
802, 819, 820.
balleus (Coregonus), 899.
baltheatus (Thynnus), 91.
baltica, var. (Pleuronectes), 393.
balticus (Hemiramphus), 345.
balticus (Lycostomus), 992.

Banksii (Gymnetrus, Regalecus), 323.
barbatula (Cobitis, Nemachilus), 704, 705, 711,
714.

barbatus (Batrachus), 135.
barbatus (Gadus), 473, 479, 493.
barbatus (Liparis), 287.
barbatus (Lopbius), 139.
barbatus (Mullus), 62.
barbatus (Rhombus), 441.

Barbel, Sea, 6 1 .

Barbus, 540, 558, 722, 742.

Barracuda, 327.

Bartel, 540.

Bartelfisk, 539.

Basking Shark, 1144.

Bathymyzon, 1182.

Bathyonus, 596.

batis (Dasybatus, Laeviraja, Raja), 1087, 1103,

1120.

Batrachidee, 126, 133, 1012.

Batracocephalus, 558.

Batrachus, 134.

Baudroies, 136.

Bdellostoma, 1195.

Beanii (Limanda, Pleuronectes), 390.

Beard, Great Forked, 540.
bearnensis (Squalius), 760.

Beaumaris Shark, 1138.

Becard, 855.

Behnii (Hemiramphus), 345.

Beinhaakall, 1147.

belone (Belone, Esox, Rhamphistoma), 347.
Belonidfe, 343.

Beloninae, 344.

Belonini, 342.
benacensis (Gobio), 743.

Benloja, 794.

Benunge, 660.

Bergabborre, 151.

Bergflundra, 385.

Berggalt, 7, 10.

Berggylta (Labrus), 4, 6, 7, 10.

Berghvarf, 456.

Berglax, 590.

Bergnabba, 14.

Bergsimpa, 173.

Bergskadda, 383, 385.

Bergsnultra, 10.

Bergstubb, 257.

Bergtorsk, 477.

Bergtunga, 375.

Bergulke, 190.

Beryeidae, 25, 66.

Beryx, 66.

Bethaj, 1132.

Bezola, 898.

biaculeatus (Gasterosteus), 648.

Bib, 493, 498.

Bibroni (Anguilla), 1023.

Bichir, 1044.

bicolor (Ammocoetes), 1183, 1186, 1189.

bicornis (Cottus), 165, 166.

bifurcus (Gadus), 540.

bilinearis (Merlucius), 120, 516.

bilineatus (Orcynus), 89.

bimaculata (Acara, Labrus, Scitena), 11.

Bimaculated Sucker, 302.

bimaculatus (Cyclopterus, Lepadogaster), 302.
bipunctatus (Abramis, Alburnus, Aspius, Cy-
prinus, Leuciscus, Spirlinus), 721, 790, 797.
bipunctatus (Gobius), 251.

Birkelanga, 521.
bison (Aspicottus), 187.
bisus (Scomber), 108.
bithynicus (Cyprinus), 724.

Bitterling, 720.

Bjelaja Rybiza, Beloribitza, 890.

bjoerkna (Abramis, Blicca, Cyprinus), 721,

’ 803, 819.

Bjalke, 772.

Bjorkare, Bjorktisk, 806.

Bjorkna, 720, 803.

Blaafinne, 806.

Blaaspol, 785.

1225

Blagarnsrocka, 1117.

Blagoma, 238.

Blainvillei (Acanthias, Squalus), 975, 1158.
Blanchardi (Gasterosteus), 658.

Blankesten, 59, 914.

Blankha, 1084.

Blanklax, 831, 851, 853, 856, 860.
Blanklodde, 882.

Blanksej, 506.

Blanksill, 989.

Blaufelchen, 909.

Blaunase, 801.

Bleak, 720, 792.

Blec-ka, 806, 822.

Bleekeria, 568.

Blege, 806.

Blekroe, 846.

Blenniidaj, 126, 212, 709.

Blenniini, 212.

blennioides (Batracocephalus), 559.

Blenniops, 217.

Blennius, 211, 212, 213, 602.

Blennoidea, 211, 212.
blennoides (Gadus, Phycis), 540.
Blennomorphi, 126, 211.

Blennus, 212.

Blenny, Butterfly, 212.

Blenny, Crested, 219.

Blenny, Viviparous, 602.

Blicca, 802, 803.

blicca (Abramis, Blicca, Cyprinus), 719, 720,
802, 803.

Bliccopsis, 796, 807, Bid.

Blicke, 806, 822.

Blinds, 493.

Blindsill, 757.

Blochii (Orthagoriscus), 626.

Blochii, (Trigla), 197.

Blomstergadda, 1005.

Blue-moulh, 155.

Blaa hafkatten, 237.

Blagunnar, 512.

Blahaj , 1130.

Blakaft, 154.

Blakaxa, 1163.

Blamag-mar , 1170.

Blamauer, 1170.

Blapanka, 806, 815.

Blaskal, 14.

Blasnultra, 10.

Blastak, 14.

Blastal, 14, 272.

Blastral, 14.

Blatobis, 574.

Blackfisk, 561.

Blotal, 1032.

Boca negra, 155.

Bogdanovii (Pleuronectes), 399, 405.

Bogmarus, 315.

bogmarus (Trachypterus), 315.

Bogini, 53.

Bola, 50.

Boleophthalmus, 240.

bolmensis (Coregonus lavaretus), 904, 905, 907.
bolniensis (Leuciscus rutilus), 777.

Bonaparti (Cybium, Pelamys), 103.

Bone Shark, 1147.

Bonit, 95.

Bonito, 95.

Bonito, Plain, 108.

boops (Ostracion), 623, 626.

Bordeliere, 806.
borealis (Batrachus), 135.
borealis (Beryx), 67.
borealis (Chiinsera), 1080.

borealis (Lsemargus, Scymnus, Squalus), 613,
1168.

borealis (Maurolicus, Scopelus), 932.
borealis (Pleuronectes), 392, 396.
borealis (Sudis), 945.

Boreogadus, 484.

Boscii (Lepidorhombus, Pleuronectes), 447.
bostoniensis (Anguilla, Muraena), 1023.
Botargo, 333.

Bothina, 371, 425.

Bothragonus, 203.

Bothus, 426, 432.

Botia, 706.

Botta, 441.

Bottengnidare, 30.

Bottenmus, 208, 211.

Bottenror, 1010.

Bottensik, 900.

bovina (Myliobatis), 1094.

Brachirus, 371.

brachycentrus (Gasterosteus), 648.

Brachy entri , 717.

Brachymystax, 882.

brachymystax (Coregonus), 899, 906, 907.
bracliypoma (Salmo), 851.
brachypterus (Thynnus), 91, 98.

Brama, 70, 72, 75, 106, 720, 812.
brama (Abramis, Cyprinus, Leuciscus), 719,
720, 812, 820.
brama (Cantharus), 54.
brama (Pterycombus), 73.
bramante (Lfeviraja, Raja), 1117.

Bramidae, 70, 309, 709.

branchialis (Ammocoetes, Petromyzon), 1189.
Branchiostegi, 619.

Branchiostoma, 1211.

Branchiostomidas, 1211.

Brandy-fish, 376.

Brandtii (Cottus), 181.
brasiliensis (Isistius), 1166.
brasiliensis (Thynnus), 93.

Braxen, 720, 812.

Braxenfiia, Braxenflira, 822.

Braxenloja, 797.

Braxenpanka, 804, 822.

Braxenstand, Bream-stand, 815.

Bream, 720, 812.

Bream, Black Sea, 54.

Bream, Common Sea, 59.

Bream, Pomeranian, 816, 817.

Bream, Rasch’s Sea, 80.

Bream, Ray’s Sea, 77.

Bream, Spanish Sea, 58.

Bream, White, 720, 803.

Breamflat, 803.

Bredtorsk, 493.

Bregmaceros, 463.
breviceps (Alburrms), 792.
breviceps (Gasterosteus), 658.
brevipinna (Lseinargus, Scymnus, Somniosus),
1168.

brevipinnis (Thynnus), 91, 93.
brevirostris (Scombresox), 352.
brevirostris (Syngnathus), 672.
brevirostris (Thymallus), 883.
brevirostrum (Acipenser), 1057.
brevis (Cephalus), 626.
brevis (Coregonus), 894.
brevis (Cyprinus), 735.
brevissimus (Batistes), 634.

Brevorti, (Brama), 81.

Brill, 441.

Brill, Turbot-like, 445.

Brismak, 562.

Brissling, 974.

Brook-Trout, 830.

Brosrne, 562.

brostne (Brosrnius, Gadus), 562.
brosme (Centronotus), 218.
brosmiana (Lota), 532.

Brosmiinae, 464.

Brosrnius 464, 562.

Brosmus, 562.

Brotula, 595.

Brotulina, 463, 595.

Brugde, 1143, 1147.

Brunsnultra, 5.

Brygd, 1144, 1147.

Bradsump, 1034.

bubalis (Cottus, Enophrys), 169, 187, 192.
bucculentus (Syngnathus), 672.
bucephalus (Cyprinus), 736.

Bucldiorn, 491.

Budd, Butt, 757.
budegassa (Lophius), 139.

Buggenhagii (Abramidopsis, Abramis, Bliccopsis,
Cyprinus, Leuciscus), 719, 808, 809, 817.
Bullerflundre, 403.

Bullhead, Alpine, 173.

Bullhead, Armed, 208.

Bullhead, River, 170.

Bundgarn, 972.

buniva (Balistes, Melichthys), 631.

Burbot, 532.

Burbot, Stone, 711.

Burbot, Tang, 711.
burdigalensis (Squalius), 760, 761.

Burgeri (Cyprinus), 733.
burgundianus (Gasterosteus), 658.

Burton Skate, 1117.

Bussei (Gasterosteus), 647.

Butirinus, 1, 947.

Butta, 432.

Butte, 432, 441.
buvosa (Leviraja), 1125.

Byrkelange (Gadus, Lota, Molva), 521.
Backro, 831.

Borting, 831, 833, 837.

Bottvad, 1035.

Cnbeljou, 479.

caucus (Gastrobranchus), 1208.

Cagnota, 213.

Calimande, grande, 451.

callarias (Gadus), 462, 464, 465, 467, 472.

callensis (Anguilla), 1023.

Callichthys, 692.

Callionymidae, 271.

Callionymini, 271.

Callionymus, 271.

Callorhynchus, 1078, 1084.

Callyodon, 6.

cambricus (Salmo), 851, 856.

Camperii (Scombresox), 353.
canadensis (Lucioperca, Stizostedium), 37.
canariensis (Anguilla), 1023.
candidissimus (Leptocephalus), 1038.

Canicula, 1152.

canicula (Scylliorhinus, Scyllium, Squalus), 1070,
1148, 1152, 1154, 1204.

Canis, 1158, 1167.
canis (Galeus), 975, 1133.

Cantharini, 53.

Cantharus, 53.
cantharus (Sparus), 54.

Cantrainii (Ramphistoma), 349.

Capelan, Capelin, Capelinus, 497, 875, 876.
capensis (Genypterus), 595.
capensis (Gymnetrus), 323.
capensis (Trigla), 202.

1226

capensis (Zens), 308.
capitatus (Cottus), 170.

Capito 764, 770, 783, 799.

capito (Mugil), 333, 334, 338, 339, 340.

capitoue (Anguilla), 1023.

Capridae, 70.
capriscus (Balistes), 634.

Carangidas, 70, 82.

Caranx, 84.

carassioides (Cyprinus), 733.

Carassius, 733.

carassius (Cyprinopsis, Cyprinus), 720, 723,
735, 736.“

carbonarius (Gadus, Merlangus, Pollachius), 500.
carbonarius (Salmo), 842.

Carcharias, 1129.

carcharias (Acanthorbinus, Canis), 1068, 1157,
1167, 1168.

Carchariidae, 1128.

Carchariin se , 1129.

Carcharodon, 1138.

cardina (Pleuronectes, Rhombus), 453.

Cardin e, 447.

Carelophus, 217.
carinatus (Cyprinus), 799.
carneus (Labrus), 10.

Carp, 720, 723, 862.

Carp, Crucian, 720, 735.

Carp, Lake, 727.

Carp, Leather, 727.

Carp, Mirror, 727.

Carp, Pond, 727.

Carp, River, 727.

Carpenteri (Onos), 544.

Carpio, 723, 731.
carpio (Cyprinus), 720, 723, 724.
carpio (Salmo), 851.
carribseus (Macrurus), 586.

Carter, 447.

Cartilaginous Fishes, 829.
caspia (Cobitis), 706.
casurus (Pleuronectes), 428.
cataphracta (Trigla), 195.
cataphractum (Peristedium), 195.
cataphractus (Agonus, Aspidophorus, Cottus),
204, 207, 208.

cataphractus (Gasterosteus), 648.

Catostomidas, 702.

Catulus, 1152.

catulus (Pristiurus, Scylliorhinus, Scyllium, Squa-
lus), 1148, 1149, 1152, 1154, 1168.
caudacuta (Motella, Rhinonemus), 545.
caudispinosus (Scopelus), 937.

Cavedanus, 770.
cavedanus (Leuciscus), 770.

Caviare, 1062.

Cefalo, 331.

Celerins, 979.

Centridermichthys, 157, 162.

Centrina, 1157.

Centriscidas, 637.

Centrisciformes, Acanthopterygii, 635.
Centriscus, 638.

centrodontus (Pagellus, Pagrus, Sparus), 56,
57, 59.

Centrolabrus, 4.

Centrophoru s , 1157.

Centroscyllium, 1157.

Cephalacanthus, 24.

Cephalaspis, 1044.

Cephalochorda, 1210.

Cephaloptera, 1094.

Cephalus, 626, 770.

ceplialus (Cyprinus, Leuciscus, Squalius), 719,
759, 769, 775.

cephalus (Mugil), 333, 334, 340.

Cepolidae, 211.

Ceratocottus, 187.

Ceratodus, 1066.

Ceratopterinae, 1094.
cerniuin (Polyprion), 48.
cernua (Acerina, Perea), 41.

Cestraciontidae, 1085.

Cestrteus, 328.

Cetorhinus, 1142.

Chsenomugil, 328, 333.

Chastodon, 24.

Chaetodontidae, 71.
chagrinea (Raja), 1115, 1125.

Chalcis, 952.

chalybaeus (Squalius), 760.

Characinidae, 66, 702.

Charr, 829, 830, 841.

Charr, Black, 846.

Charr, Light, 846.

Charr, Northern, 842.

Charr, Wetter, 842.

Chatoessinm, 952.

Chauliodontidae, 930, 936.

Chela, 823.

Chelidon, 356.

chelo (Mugil), 328, 333, 334, 340.

Chelon, 331, 334.
chilensis (Brama), 77.
chilensis (Sarda), 102, 104.

Chilomycterus, 620.

Chimsera, 1043, 1063, 1079.

Chimaeridas, 1078.

chinensis (Albula, Salmo), 827.

chinensis (Cyprinus), 720.

Chiridae, 147, 211.

Chirolophis, 213, 217.

Chironectes, 145.

Chlamydoselache, 1063, 1068.

Chondropterygii, 1, 1043, 1063.

Chondrostei, 1, 1043.

Chondrostoma, 716.

Chorinemus, 82.

Chorisochismus, 300.

Christmas-fish, 406.

Chromidas, 1131.
chrysitis (Tinea), 748.
chrysochloris (Clupea), 987, 988.

Chrysophrys, 56.
chrysops (Roccus), 44, 46.
cibarius (Ammocoetes, Petromyzon), 1189,
1193.

cicatricosus (Pleuronectes), 404, 422, 429.
cicerelus (Ammodytes), 569, 573.
cii (Leuciscus). 770, 772.

Ciliata, 544.

cimbrica (Clupea hareugus), 959.
cinibrius, (cimbricus, Enchelyopus, Gadus, Mo-
tella, Onos), 544.
cinerea (Raja), 1120.
circularis (Raja), 1112.

Cirrostomes, 1 .

Cirrostomi, 1210.
cithara (Callionymus), 279.

Citharichthys, 426.

Citharus, 371, 426.

Civelle, 1032.

Clarias, 690, 691.
clathratus (Squalius), 770.
clavata (Dasybatis, Raja), 1064, 1074, 1087,
1103, 1104, 1108.

Clavatee, Rajas, 1104.
claviger (Cottus), 188, 192.

Climbing-fish, 240.

Clinus, 219.

Clupea, 952, 953.

Clupea harengus, 149.

Clupeidae, 827, 946.
clupeiformis (Coregonus), 899.
clupeoides (Coregonus), 890, 894, 899.
Clypeocottus, 187.

Coalfish, 465, 499, 500.
cobites (Aphya), 262, 265.

Cobitidse, 702, 703.

Cobitis, 703, 705.

Coccia, 930, 934.

Cocciinfe, 923, 930.
coccineus (Lycodes), 612.

Cod, 465.

Cod, Bearded, 493.

Cod, Broad, 493.

Cod, Common, 472.

Cod, Green, 500.

Cod, Polar, 463, 484.

Cod, Poor or Power, 495, 498.

Cod, Tang, 477.

Cod, Three Bearded, 550.

Coelorhynchus, 584.

coelorhynchus (Lepidoleprus, Macrurus), 581,
584, 585.

coerulea (Pelamis), 93.
coeruleus (Carassius), 733.
coeruleus (Carcharias), 1130.
coeruleus (Cyelopterus), 294.
coeruleus (Labrus), 10.

Coilia, 990.

Coltish, 500.

colias (Scomber), 90, 110.

Colin, 500.
comber (Labrus), 7.

Commersonii (Scomber), 93.

Common Trout, 856.
communis (Alosa), 985.
communis (Lota), 532.
communis (Merlangus), 511.
compressus (Cyprinus), 779.
compressus (Gadus), 532.

Concadita, 331.

concinnus (Gasterosteus), 658.

Conger, 1036, 1037.

conger (Anguilla, Leptocephalus , Muraana,
Ophichthus), 1037.

Congrogadus, 595, 603.
conirostris (Cyprinus), 724.

Connor, 18.

conorhyncbus (Coregonus), 899.
conspersus (Pleuronectes), 429, 430.
conspicillum (Batracbus), 135.

Corax, 200.
corax (Trigla), 201.

Coregonus, 828, 884, 891, 892.
coretta (Thynnus), 91, 98.
coriaceus (Cyprinus), 724.

Coris, 21.

cornubicus (Isurus, Lamna), 1069, 1129, 1136,
1138.

cornubiensis (Labrus), 7, 18.
cornubiensis (Barbus minor), 558.
cornuta (Myliobatis), 1095.
coronatus (Cyelopterus), 294.
corrugatus (Mugil), 334.

Corvina, 51.

Coryphaena, 96, 309, 567, 588.

Corypliaenidaa, 70.

Coryphaenoides, 583.

Cossyphus, 5.

Costas Prickfisk, 937.

Cottida?, 126, 156.

Cottinae, 156.

Cottomorphi, 126, 146.

1227

Cotto-scombriformes, 70.

Cottunculus, 157.

Cottunculus, Small-eyed, 158.

Cottus, 146, 157, 168, 272, 273.

Cottus, Bearded, 208.

Cottus, Branch-spined, 160.

Cottus, Four-horned, 175.

Cottus, Short-spined, 180.

Couehia, 544, 567.

Couchii (Acantholabrus), 5.

Coucliii (Crenilabrus), 18.

Crangon, 258.

crassiceps (Macrurus), 581.

crassus (Leuciscus rutilus), 777.

Crenilabrus, 4, 8.
cristata (Chimtera), 1070.
cri status (Pleuronectes), 441.

Crossorbinus, 1147.

Crucian Carp, Lake, 736.

Crucian Carp, Pond, 738.
cruentatus (Gobius), 248.

Crystallogobius, 242, 268.

Ctenodon, 224.

Ctenolabrus, 4, 16.
cubana (Anguilla), 1023.
cuculus (Trigla), 195, 197.
cultratus (Abramis, Cyprinus, Leuciscus, Pele-
cus), 721, 823.
curtus (Mugil), 334.

Cuvieri (Anguilla), 1023.

Cuvieri (Argentina), 917.

Cuvieri (Cyprinus), 733.

Cuvieri (Trachurus), 88.

Cuvierii (Alosa), 985.

Cybium, 89, 91, 99.

Cyclogaster. 282, 283.

Cyclopteridse, 126, 147, 282.

Cyclopterus, 282, 293.

Cyclospondyli, 1128, 1156.

Cyclostomes, 1, 1043.

Cyclostomi, 1063, 1172, 1195.

Cynicoglossus, 386.

Cynoglossus, 371.

cynoglossus (Glyptocephalus, Platessa, Pleuro-
nectes), 364, 378, 383, 417.

Cyprinidas, 66, 702, 714.

Cyprininfe, 720, 722.

Cyprinodontidse, 342, 997.

Cyprinodontime, 702.
cyprinoides (Coregonus), 891.

Cyprinornorphi, 690, 702.

Cyprinopsis, 733, 735.

Cyprinus, 719, 722, 723.

Cypselurus, 356.

Cypsilurus, 356.

Cyttidfe, 304, 305.

Cyttomorphi, 126, 304.

Czernayi (Owsianka), 787.

Dab, Common, 386.

Dab, Lemon, 383.

Dab, Rough, 421.

Dab, Smear, 383.

Dace, 720, 759, 760.

dactyloptera (Scorpsena, Sebastes), 153, 154,
522.

Dactylopterus, 147, 193.

Dajaus, 328.

Dammruda, 738.

Deal fish, 315.

decadactylus (Beryx), 67, 429.
decagon (Argentina), 918.
decagonus (Agonus, Archagonus, Aspidophorus,
Cottus, Podothecus), 204, 205, 206.

decipiens (Leuciscus), 773.
de Filippi (Vexillifer), 600.

Dekayi (Gasterosteus), 658.
delicatulus (Spratelloides), 952.
delineatus (Leucaspius, Squalius), 721,754,787.
dentatus (Echiodon, Fierasfer, Ophidium), 599,
601.

dentatus (Paralichthys, Pseudorhombus), 366.
dentex (Cliorisochismus), 300.
dentex (Osmerus), 867, 870, 875.
denticulatus (Anarrhichas), 237.
dergle (Scardinius), 7 79.

Dermopteri, 1210.

Desfontainii (Lepadogaster, Mirbelia), 303.
Devil-fish, 1094.
diaphanus (Pleuronectes), 428.
diceraus (Ceratocottus), 187.

Dicerobatis, 1094.

Dick-kopp, 772.

didactylus (Batrachus), 134.

dimidiatus (Gasterosteus), 648.

Diodon, 620.

Diplanchias, 626.

Diplodus, 53.

Diplophus, 930.

dipterygia (Molua), 521, 525, 531.
Discoboles, 282.

Discoboli, 282.
discolor (Carassius), 733.
dispar (Labrus), 10.
dispersus (Coregonus), 899.

Djuproding, 847.

dobula (Cyprinus, Leuciscus, Squalius), 721,
760, 770.

dobuloides (Alburnus), 795.

Dog-Fish, Black-mouthed, 1149.

Dog-Fish, Picked, 1158.

Dogge, 476.

Dogs’-fish, 997.

dolabratus (Alburnus), 719, 795.

Donovani (Crenilabrus), 18.

Donovani (Labrus), 7.

Doras, 188, 691.

Dorosomatinae, 952.
dorsatus (Petromyzon), 1187.

Dory, 306.

draco (Trachinus), 128, 272.
draculus (Callionymus), 273, 279.
Dracunculus, 272, 279.

dracunculus (Callionymus, Uranoscopus), 273.
Drag, 1010.

Dragonet, Gemmeous, 273.

Dragonet, Lesser, 279.

Dragskadda, 391.

Dragspan, 1010.

Drepanopsetta, 408, 420.

Drift-net, 971.

Drifvad, 1035.

Drummondii (Echiodon), 601.
dubius (Mullus), 63.

Diimerilii (Syugnathus), 672, 673.
Dussumierinae, 952.

Dvergsimpa, 191.

Dvergulk, 190.
dvinensis (Osmerus), 867.
dvinensis (Platessa) 405, 406.
dypterigius (Gadus), 521.

Echelus, 1037.

Echeneis, 89, 91.

Ecbinorhini, 1157.

Echinorhinidaa, 1166.

Echinorhinus, 1169.

Echiodon, 601.

Edwardii (Couehia), 545.

Eel, Common, 1023.

Eel, Electric, 1086.

Eel-basket, 1034.

Eel-box, 1034.

Eel-fare, 1031.

Eel -fishes, 1011.

Eel-line, 1035.

Eelpout, 603.

Eel-seine, 1035.

Eel-spear, 1035.

Eel-torsk, Lycodes, 607.

Eel-torsk, Eel-shaped, 617.

Eel-torsk, Plain, 610.

Eel-torsk, Reticulated, 611.

Eel-torsk, Sarsian, 616.

Eel-torsk, Vahl’s, 613.

Eel-trunk, 1034.

Eel-weir, 1035.

Eels, Freshwater, 1022.

Eggerti (Anarrhichas), 236.

Ekstromii (Gobius), 262.

Ekstromii (Liparis), 284.

Ekstrom’s Topknot, 453.

Elacate, 89, 91.

Elasmobranchii, 1043, 1063.
elatus (Cyprinus), 724.
elatus (Leuciscus rutilus), 777.

Elbbutt, 443.

Elbe caviare, 1062.

El but, Elbutte, 757.

Ehlstodjning, 1010.
elegans (Gasterosteus), 648.

Eleginus, 482.

Eleotriformes, 240.

Eleuterognathi, 2.
elongata (Argentina), 912, 918.
elongata (Clupea), 954.
elongata (Cobitis), 706.

elongata (Glyptocephalus, Platessa, Pleuro-
nectes), 379.

[ elongata (Lotta), 521.
elongatum (Myctophum, Scopelus), 923,937, 993.
elongatus (Abramis, Leucabramis), 798.
elongatus (Leuciscus rutilus), 7 77.

Elops, 1056.

Elritza, 720, 754, 757.

Emd, 767.

Encheliophis, 462, 601.

Enchelymorphi, 690, 1011.

Enchelyopus, 211, 537, 570, 603, 1030.
Encrasicholus, 992.

encrasicholus (Clupea, Engraulis, Stolephorus),
952, 975, 992.

[ Engelsmau, 425.

Engraulis, 952, 991.
j Engyschistae, Muramidas, 1021.

Enkrasicolus, Enkraulos, 952.

Enophrys, 181, 187.

Entelurus, 680.

Epelan, 869.

Eperlan de Seine, 798.
eperlano-marinus (Salmo), 869.

Eperlanus, 869.

eperlanus (Osmerus, Salmo), 867, 869, 875,
876, 878.

Epigonichthys, 1211.

| Epinephelus, 48.
j equirostrum (Scombresox), 353.

Eriksmass-lax, 832.
eriox (Salmo), 832, 850, 851.
erythraeus (Salmo), 842.
erytbrinus (Sparus), 56, 57, 58.
erythro-bjorkna (Scardo-Blicca), 807.
erythrophthalmoides (Bliccopsis), 807, 808.

1228

Ery throplitb al m us , 779.

erytlirophthalmus (Cyprinus, Leucisou's, Scar-
dinius), 720, 779.
erythrops (Cyprinus), 779.
erythropterus (Abramis), 803.
erythrostomus (Aspius), 782.
esculentus (Merlucius), 515.

Esmarkii (Gadus), 466, 467, 508.

Esmarkii (Lycodes), 613.

Esocida?, 997.

Esociformes, 690, 997.

Esox, 342, 998.

Esox brasiliensis, 345.
estor (Esox), 999.

Ethmopterus, 1162.
ethon (Syngnathus), 669.

Etmopterus, 1157, 1162, 1166.

Eucalia, 647.

Euchalarodus, 405.

Eulachon, 875.

Euprotomicrus, 1156.
europaeus (Acipenser), 1056.
europasus (Aspidophorus), 208.
europaeus (Hemirampbus), 345.
eurybrachii, Acanthopterygii, 24.
euryrliynchi (Coregoni lavareti), 902.
euryripidi, Acanthopterygii, 24.
eurypterus (Lophius), 139, 142, 143.
eurystoma (Anguilla), 1023.

Euthynnus, 91, 92.

euxinus (Gadus), 464, 487.

evolaus (Exocoetus, Halocypselus), 358.

exiguus (Coregonus), 899.

exiliens (Exocoetus), 358.

Exocoetiforines, 343.

Exocoetinae, 355.

Exocoetini, 342.

Exocoetus, 356.

exoletus (Acantholabrus, Centrolabrus, Labrus),
4, 10, 14.

exoletus (Crenilabrus), 5.
exoletus (Labrus), 941.
exsiliens (Exocoetus), 358.

1'aber (Zeus), 305, 306, 975.

Fabraei (Alburnus), 792.

Fabricii (Boreogadus, Gadus), 464, 484.
Fabricii (Liparis, Cyclogaster), 287, 288.
Fabricii (Lumpenus), 224, 225, 226.

Fabricii (Macrurus), 584, 586.

Facbse-lodde, 882.

Faisan d’eau, 440.

fallax (Clupea), 984.

falsa (Alausa), 894.

falsavela (Raja), 1089, 1103, 1112.

Fana, 822.

Faren, 720, 812, 819, 822.
farenus (Cyprinus), 812, 819.

Fario, 841, 851.

fario (Salmo), 832, 851, 855.

Fame, 822.

Farra, 898.

Father-lasher, 187
Fengommare, 73.

Fenknoten, 200.
fera (Coregonus), 899.
fernandinus (Spinax, Squalus), 1158.
ferox (Salmo), 851, 857, 858.

Ferra, 898.

ferrugineus (Limanda, Pleuronectes), 390.
festiva (Julis), 21.

Fetsik, 900.
beta (Alosa), 984.

Fierasfer, 462, 598, 600.

Fierasfinae, 597.
filamentosus (Krolinius), 581.

Filare, 631.

File-fish, 631, 717.

File-fish, Spotted, 633.

finta (Alausa, Alosa, Clupea), 984, 987.

Fire-pan, 656.

Fishing-frog, 138.

Fistulariidse, 637.

Fjord -lodde, 882.

Fjallflundra, 391.

Fjard-lax, 832.

Fjarsingen, vanliga, 128.

Fjarsingen, lilla, 131.

Fjasing, 131.

fiammeus (Cyprinus), 724.

Flatfishes, 361.

flavescens (Bodianus, Perea), 27.
flavescens (Brosmius), 562.
flavescens (Gobius), 243, 244, 250, 251, 257.
tiavipinnis (Cyprinus), 724.

Flek-Steenbit, 236.

flesus (Platessa, Pleuronectes), 378, 398, 407.
Flira, 720, 815, 819, 822.

Flire, 806.

Flodharr, 887.

Flodnejonoga, 1188.

Flotqvabba, marmorerad, 145.

Flotqvabbor, 144.

Flounder, 398.

Flounder, King’s, 396.

Fluke, Sail, 451.

Flundra, 2, 403.

Flute-mouth, 635, 637.
fluviatilis (Anguilla), 1023.
fluviatilis (Cephalus), 770.
fluviatilis (Gobio), 720, 743.
fluviatilis (Lampetra, Petromyzon), 1173, 1174,
1188, 1189, 1203.
fluviatilis (Perea), 26, 1204.
fluviatilis (Rubellus), 773.
fluviatilis (Salmo), 830.
fluviatilis (Trutta), 851, 852.

Flydra, 416.

Flygfisk, 272.

Flying-fish, 355.

Flying-fish, Great, 357.

Flackfilare, 633.

Flack pagell, 59.
fontinalis (Salmo), 830, 845.

Forel, 831, 833.

Forked-beard, Great, 540.

Forked-beard, Lesser, 558.

fossilis (Cobitis, Misgurnus), 703, 704.

Franklinii (Pleuronectes), 404, 405.

Friesii (Gobius), 244, 248.
frigidus (Leuciscus), 770.
frigidus (Lycodes), 609, 610.
fuliginosus (Gadus), 559.
fullonica (Leucoraja, Raja), 1 103, 1108, 1115,
1117.

Fundulus, 743.

furcatus (Phycis, Physcis), 540.
furciger (Icelus), 166.
fuscus (Motella, Onos), 551, 556.
fuscus (Blennius, Raniceps), 558, 559.

Farna, 769, 772.

Farsing, 131.

Gradidae, 462, 463.
Gadinae, 464.
gadoides (Blennius), 540.
Gadus, 464, 514, 693.
Gaimardi (Raja), 1121.

Galaxiidse, 829.

galeatus (Gymnacanthus), 160.

Galeorhinus, 1072, 1 129, 1132.

galerita (Blemdops, Blennius, Chirolophis), 218.

Galeus, 1 1 33.

galeus (Canis, Galeorhinus, Galeus, Squalus),
1132, 1133.

Gallichthys, 309.

Gall-id, 767.
gallivensis (Salmo), 851.

Ganoidei, 581, 718, 829, 1043.

Ganoids, 1.

Garpike, 347.

Gasteracanthus, 647, 658.

Gasterobranchus, 1220.

Gasterosteidae, 341, 637.
gasterosteiformes, Acanthopterygii, 635.
Gasterosteus, 637, 638, 644, 658.

Gastraea, 637, 638.

Gastrobranchus, 1208.

Gayi (Merlucius), 515.

Gehini (Abramis), 812.

gelatinosus (Careproctus, Cyclopterus, Liparis),
283, 287, 291.
generosus (Coregonus), 899.

Gestria, 1182.

germo (Orcynus, Thynnus), 90, 91, 97.

Gers, 41.

Gertrude’s Pike, 1005.

Ghini (Orthagoriscus), 626.
gibba, var. (Perea), 29.
gibbus (Labrus), 18.
gibbus (Liparis), 287.

gibelio (Carassius, Cyprinopsis, Cyprinus), 721,
735, 738.

gibelioides (Cyprinus), 733.

Giblichen, 735.

Giersii (Pleuronectes), 395.

Gillaroo- trout, 840.

Gilt Charr, 831.

Gilt-head, 18.

Giofredi (Julis), 21.
glabra (Piatessa), 405.

glaciale (Myctophum, Scopelus), 923, 941, 945.
glacialis (Cottus), 180.
glacialis (Gadus), 484.

glacialis (Pleuronectes), 404, 405, 422, 429.
glacialis (Squalus), 1168.

Glade, 391.

gladius (Cepola, Gymnetrus, Regalecus), 321,
323.

gladius (Xiphias), 118.
glanis (Silurus), 693.

Glanomorphi, 690, 1043.

Glansfisk, 123.

Glasbult, 269.

Glashvarf, 448.

Glasstubb, 269.

glauca (Ciliata, Couchia, Motella), 554, 556.
glauca (Licbia), 8-3.

glaucus (Carcharias, Carcharinus, Galeus, Squa-
lus), 983, 1075, 1130.
glaucus (Squalus, Haabrandunge), 1138.
glesne (Gymnetrus, Ophidium, Regalecus), 322.
Gli, Glirr, 757.

Glib, 425.

Glibskadda, 425.

globiceps (Gasterosteus), 658.

Glossgadda, 1005.

Gloveri (Salmo), 850.

glutinosa (Myxina, Myxine), 472. 1196, 1208.
glutinosus (Gasterobranchus), 1220.

Glutinous Hag, 1208.

Glyptocephalus, 378, 379.

Glysa, 497.

1229

Glyskolja, 495, 497.

Gmelini (Phycis), 543.

Gobiesocidm, 126, 300.

Gobiesociformes, 239.

Gobiidie, 126, 240.

Gobiifornies, 239, 240.

Gobiime, 240.

Gobio, 720, 722, 742.

gobio (Cottus), 157, 160, 169, 170, 173.

gobio (Cyprinus, Leuciscus), 743.

Gobioidei, 239.

Gobiomorphi, 1 26, 239.

Gobitis, 706.

Gobi us, 157, 242.

gobius (Cyclopterus, Liparis), 284.

Goby, Black, 245.

Goby, Doubly spotted, 251.

Goby, Freckled, 262.

Goby, Jeffrey’s, 261.

Goby, Spotted, 262.

Goby, White, 266.

Goedenii (Salmo), 851.

Goldfish, 733.

Gold Ide, 767, 777.

Gold Perch, 30.

Gold Roach, 777.

Goldsinny, 16.

Goniosoma, 918.

gracilis (Blennius, Clinus, Leptoblennius, Lum-
penus), 225.

gracilis (Coregonus), 898.

gracilis (Gadus, Tilesia), 465, 467, 481.

gracilis (Gobius), 248, 262.

gracilis (Ly codes), 613, 615.

gracilis (Perea), 27.

Grammiconotus, 352.

Grande Caliinande, 451.
grandoculis (Carassius), 733.
granulata (Perea), 27.
granulosus (Centrophorus), 1158.
granulosus (Spinax), 1162, 1166.

Grayling, 882, 884.

Greenfish, 580.

Grenadiers, 581.

Grentaggiga Ulken, 160.

Grillii (Gymnetrus), 322.
grisea (Lucioperca), 37.
griseus (Cantharus), 54.

grislagine (Cyprinus, Leucisgus), 720, 759, 761,
763, 775.

Grodqvabba, 144.
i grcenlandicus (Cottus), 180.

groenlandicus (Hippoglossus), 417.
grcenlandicus (Mallotus, Salmo), 876.

Groploja, 787.

Grubei (Thymallus), 882, 883.
gruncus (Echelus), 1037.

Grundforell, 838.

Grytmort, 777.

Gralax, 830, 831, 851.

Gralanning, 746.

Grasej, 499, 500, 503.

Grasik, 503.

Grasgadda, 1005.

Grassnultra, 14.

Grassill, bolnislansk, 959, 964.

Grastorsk, 476.

Grasal, 1032.

Groning, 579.

Gronling, 711.

Guaperua, 634.

Gudgeon, 720, 743.

Gudlax, Guldlax, 126.

Guiniad, Gwiniad, Gwyniad, 898, 909.
Guldmulle, 63.

Guldmulte, 337.

Guldmort, 777.

Gulha, 1084.

Gunnel, Spotted, 220.

gunnellus (Blennius, Centronotus, Murrenoides,
Ophisomus, Pholis), 220.

Gunneri (Scymnus, Squalus), 1168.

Gunneri (Spinax), 1163.

Gunnerianus (Cetorhinus, Squalus), 1144.
Gurnard, Grey, 197.

Gurnard, Red, 195.

Gurnard, Sapphirine, 200.

Gurnards, Cuirassed, 202.
gurnardus (Trigla), 194, 197.

Gussonii (Leptocephalus), 1038.
guttatus (Lampris, Zeus), 123.

Guldenstadti (Acipenser), 1059, 1060.

Glister, 806.

Gwyniad, 891, 898.

Gwyniad, Asp, 891.

Gwyniad, Beaked, 902.

Gymnetrus, 320.

Gymnocantbus, 157, 159.

Gymnodontes, 622.

Gymnogaster, 314.
gymnogaster (Thymallus), 884.
Gymnothoracidse, 1021.

Gymnothorax, 1022.
gymnothorax (Thymallus), 884.

Gymnotidae, 1086.

gymnurus (Gasterosteus aculeatus), 644, 648.
Gardal, 1035.

Gadda, 998.

Gokrocka, 1115.

Gos, 36.

Haa, 1161.

Haabrand, 1138.

Haae-Kierring, 1167.

Haae-Storje, 1056.

Haaf ur, 1158, 1161.

Haa-ising, 425.

Haakall, 1167.

Haameer, 1138.

Haavar, 1161.

Gaberdine, 479.

Haddock, 466.

Haddock, Norway, 148.

Hrnlt, 909.

haematopterus (Cyprinus), 724.

ILemulon, 52.

Hafgos, 50.

Hafgosfiskar, 49.

Hafkarp, 10.

Hafkatt, bla, 237.

Hafkatt, tigerflackad, 236.

Hafkatt, vanlig, 232.

Hafmus, nordisk, 1079, 1084.

Hafpadda, 561.

Hafr, 1161.

Hafsabborre, 45.

Hafsapa, 1084.

Haf sbarber, 6 1 .

Hafskatt, 1161.

Hafsnejonoga, 1183.

Hafsnalar, 679.

Hafsnalen, stora, 680.

Hafspadda, 144.

Hafsruda, 52.

Hafssill, 959.

Hafsal, 1037.

Hafulk, 144.

Hag, Glutinous, 1208.

Haj, 1161.

Hake, 465, 515.

Hake, Trifurcated, 559.

Ilalb Harass, 719.

Halec, 952.

Halecoides, 826.

Halecula, 992.

Halfulk, 163.

Halhos, 197.

Halibut, 409.

Halibut, Lesser, 417.

Haliophis, 595.

Halocypselus, 356.

Haloporphyrus, 539, 581.

hamatus (Centridermichthys, Icelus), 165.

hamatus (Salmo), 850, 855.

Hammerhead, 1071, 1128.

Hamntorsk, 757.

Handvad, 1035.

Hansitig, 396.

Haplochitonidse, 826.

Haploidonotus, 49.

Haplomi, 690, 997.

Hardinii (Salmo), 850.

Hareng, 954.

Harengus, 954.

harengus (Clupea), 952, 954.

Harkning, 1035.

Harr, 884.

Hasel, 760.

Haspa, 887.

Hausen, 1055, 1058.

Havkatt, 1079, 1084.

Havkrage, 1084.

Havkvabbe, 543.

Havlodde, 882.

Havmus, Havmuus, 211, 1079.

Hawkenii (Gymnetrus), 322.

Hawkinsii (Gymnetrus), 322.

Hay, 1167.

hebridicus (Argentina, Osmerus), 917.

Heckelii (Abramis), 818.

Heckelii (Leuciscus), 774, 777.

Heckelii (Nerophis), 680.

Heiligbutt, 416.

Heilag-fiske, 416.
lielena (Mursena), 1021.

Helmichthyidie, 1020.

Helmichthys, 1020.

Helmictis, 1020.
helvetica (PercaJ, 27.

Helgeflundra, 409, 416.

Plelgeflundran, lilla, 417.

Hellefiyndre, 416.

Hemibranchii, 2, 635.

hemigymnus (Argyropelecus), 305, 925, 927,
929, 930.

Hemilepidotus, 156.

Hemirhamphiformes, 344.

Hemirhamphus, 342.

Hemirhombus, 426.
heptagonus (Cyclopterus), 294.

Herring, 947, 954.

Herrings, 826, 946.

Heterobranchus, 690, 691.

Heterocerci, 1044.

Heterocormi, 24.

Heterolepidotidm, 147, 211.
heterolobus (Stolephorus), 990.
heteropteras, Siluridm, 692.

Heteropygii, 997.

lieterorhynchus (Labeo, Schismatorhynchus),

716.

Heterosomata, 3 6 1 .

Heterosomes, 361.

Heterothrissa, 990.

1230

liexacornis (Cottus), 175, 178.
hiatula (Labrus), 11.
hiemalis (Clupea harengus), 959.
hiemalis (Coregonus), 899.

Hierax, 352.

Hippocampime, 663.

Hippocampus, 662.

Hippoglossina, 371, 408.

Hippoglossoides, 420.

hippoglossoides (Hippoglossus, Platysomatich-
tliys, Pleuronectes, Reinhardtius), 408, 417.
Hippoglossus, 408, 409, 420.
hippoglossus (Pleuronectes), 409.

Hirax, 352.

hirtus (Pleuronectes, Rhombus, Zeugopterus),
458.

hirundinaceus (Carcharias), 1130.

Hirundo, 200, 356.

hirundo (Trigla), 195, 197, 200.

hispidus (Orthagoriscus), 626.

Histiophorus, 117.

histrio (Antennarius, Lophius), 136, 145, 634.
Hlyre, 236.

Holocentrini, 66.

Holocentrum, 66, 69.

Holocephali, 1072, 1078.
hololepidotus (Labrus, Sciama), 52.

Holostei, 1044.

Homalopteridse, 702.

Homianus (Cetorhinus, Squalus), 1144.

Hork, 43.

Hornfisk, 347.

Hornlisk, Hornstagg, Horntagg, 657.
Horngadda, 347.

Horrigall, 347.

Hornsimpa, 175.

Hornskalle, 185.

Iiorntunga, 425.

Hornuggla, 211.

Hornulke, 190.

Horn-al, 1035.

Horr, 43.

Horse-Mackerel, 86.

Horunge, 393, 443.
hospitus (Acipenser), 1057.

Hound, Nurse, 1076, 1152.

Hound, Rough, 1070, 1 154.

Houting, 902, 909.

Huch, 830, 831.

Huch, Danube, 830.

Huch, Siberian, 830.
bucho (Salino), 830, 851.

Hulekolja, 561.

Humboldti (Scopelus), 932.
humilis (Coregonus), 898.
humilis (Cyprinus), 736.

Hundfisk, 'll 58.

Hundgadda, 757.
hungaricus (Cyprinus), 724.
huntia (Molva), 532.
hurta (Sparus), 56.

Huso, 1054, 1055.

huso (Acipenser), 1057, 1059.

Huttning, 1035.

Hvarf, luden, 456.

Hvassbuk, 974.

Hviid-Aal, 1208.

Hvitling, 487.

Hvitlingiyra, 499, 508.

Hvitrocka, 1117.

Hvitsik, 900.

Hvit-tobis, 570.
hybiscoides (Cyprinus), 724.
hybridus (Aspius), 782.
hybridus (Rhombus), 432.

' hyperborea (Raja), 1103, 1110, 1111.
Hyponiesus, 875.

Hypotremes, 1093.

hypselorhynchi (Coregoni Javareli), 902.

Ha, 1161.

Habrand, 1138.

Hafisk, 1161.

Hagai, 1149.

Hakal, 1161.

Hakatt, 1161.

Hakaring, 1161, 1167.

Hamus, 1084.

Hamar, 1161, 1170.

Harnacka, 769.

Harump, 1162.

Haskarding, 1167.

Haskarfving, 1155.

Hastorje, 1132.

Hallefisk, 416.

Halleflundra, 416.

Hogfenig smorbult, 259, 260.

la, 859.

Ice-fish, 767.

Icelus, 162.

Ichthyodorulites, 1067, 1156.

Ichthyomyzon, 1182.

Ide, 720, 764, 768.

Ide, Gold, 767.

Idbarn, Idplugg, 767.

Idbarus (Cyprinus), 721, 764.
idiopteri, Acanthopterygii, 24.
idus (Cyprinus, Leuciscus), 720, 759, 764,
770, 775.

Igelneinoga (-nejonoga), 1195.

Iggling, 757.

Illanka (Salmo), 851.
imberbe (Ophidium), 220, 598.
imbricatus (Acantholabrus), 5.
immaculatus (Salmo), 914.
imperiale (Ramphistoma), 349.
imperialis (Sebastes), 154.
insequalis (Leptocephalus), 1021, 1038.
inconstans (Eucalia, Gasterosteus), 647.

Ingeris Pilt, 1208.

Ingminnuiset, 587, 589.
inopinatus (Gasterosteus), 648.
inornata (Lota), 532.
intermedia (Raja), 1123, 1125.
interruptus (Roccus), 44.

Ipnops, 923.

irideus (Salmo), 830.

iris (Trachypterus), 311, 312, 313, 315.

Isa, 466.

Isabborre, 30.

Ischyodon, 1078.

Isfisk, 767.

Isgalt, 1084.

Ising, 391.

Isistius, 1156.

islandicus (Boginarus, Vogmarus), 315.
islandicus (Centronotus, Gunellus, Lumpenus,
Stichseus), 225.

islandicus (Gasterosteus), 648, 649.

Ismort, 776.

isodus (Squalus), 1144.

Isospondyli, 690, 826.

Isurus, 1069, 1138.
italica (Tinea), 748.
italica (Perea), 27.

italicus (Hymenocephalus, Macrurus, Mysta-
couurus), 583.

jacksonionsis (Regalecus, Trachypterus), 314.

japonica (Anguilla), 1023.

japonica (Brama), 77.

japonica (Cobitis trenia), 706.

japonicus (Gasterosteus), 647.

japonicus (Macrurus), 583.

japonicus (Petromyzon), 1193.

japonicus (Zeus), 306, 308.

Jeffreysii (Gobius), 244, 261.

Jernlodde, 882.

Jesen, 764.

jeses (Cyprinus, Idus), 765.

John Dory, 306.
jozo (Gobius), 245.

Judepiga, 425.

jugulares, Acanthopterygii, 24.

Julidinse, 20.

julis (Coris, Labrus), 21.

Jydekjaerling, 425.

Jydetunga, 378.

Jydetuuge, 425.

Kabeljaauw, 476.

Kabiljo, 473, 476.

Kalleraglik, 418.

kanagurta (Scomber), 90, 110-

Kantnal, 674.

Karas, Harass, 735.

Karp, 720, 768.

Karpffkarass, 719.

Karpruda, 732.

Karpsten, 716.

Kartmort, 777.

Katsa, 816.

Kaidbarsch, 43.

Keila, 562.

Kelt, 837.

Kieneri (Anguilla, Zoarces), 617, 1023.

Kilcli, 909.

Kilnot, 859.

King of the Herrings, 322, 1084.

King of the Sand-Eels, 580.

Kin-ju. 733.

kitt (Pleuronectes), 383.

Kivik-sill, 959, 963.

Kjarta, 806.

Klrep-al, 1033.

Klarbult, 266.

Klarstubb, 266.

Klorocka, 1108.

Klumpfisk, 2, 625.

Kluinpkrok, 1008.

Klykskrabb, 165.

Knaggrocka, 1104.

Knerii (Polyprion), 48.

Knorrhane, 197.

Knot, 197.

Kolja, 466.

Kolje, 497.

Kollarii (Carpio, Cyprinus), 731.

Kolmule, 511.

Kolmun, 519.

Kongeral, 1038.

Koning van Klipvischen, 595.

Kordylos, 99.

Kortsa, 763.

Kortstjertgadda, 1005.

Krohnius, 581.

KrOyeri (Scopelus), 938, 941.

Kuderusk, 211.

Kulla Herring, 959, 963.

Kullbas, 43.

Kullribas, 43.

Kulmund, 519.

1231

Kummel, 515, 519.

Kumrill, 540, 543.

Rung, 579.

Kungsfisk, 148, 152.

Kungsflundra, 396.

Kussa, 607.

Kveise, 131.

Karing, 151.

Labeo, 716, 718.

Laberdan, 479.

Labordii (Euprotomicrus), 1166.

Iabradoricus (Cottus), 180.

Labrax, 45.

labrax (Perea, Roccus), 45.

Labridse, 3.

Labrina, 4.

Labroides, 2.

Labromorphi, 2.

Labrus, 2, 5.

Lacepedii (Coregonus), 899.
lacertina (Siren), 1204, 1205.

Lachsforel, 838.

laenstris (Cyprinus), 763.

lacirstris (Gadns, Lota), 532.

lacustris (Salmo, Trutta), 831, 850, 851.

Lsemargi, 1156.

Leemargus, 1167.

Lminonema, 539.

lseves, Rajas, 1114.

lasvigatus (Chironcctes), 145, 146.

lasvis (Arnoglossus), 428.

lasvis (Cataphractus, Pholis), 214.

lasvis (Gasterosteus), 658.

lasvis (Macrurus, Malacocephalus), 584, 593.

lasvis (Phoxinus), 754.

lasvis (Pleuronectes), 383.

lasvis (Pleuronectes, Rhombus), 441.

lasvis (Raja), 1117, 1120.

laevis (Trigla), 201.

Laggbuk, 622.

Lakatrubb, 174.

Lake, 532.

Lakegers, 174.

Lamna, 1138.

Lamnidae, 1128, 1135.

Lainninae, 1135.

Lampanyctus, 937.

Lampern, *1 188.

Lampetra, 1183.

lampetra (Petromyzon), 1183.

Lainpreda, 1183, 1188.

Lampreta, 1183.

lampretiformis (Blennius, Clinus, Lumpenus),
225.

Lamprey, 1183.

Lampreys, 1, 1182.

Lamprididse, 70, 121.

Lamprini, 121.

Lampris, 122, 309.

Lamproye d’eau doulce, 1188.

Lamproye de mer, 1183.

Lamproyon, 1188.
lancastriensis (Leuciscus), 761.
lancea (Ammodytes), 575.

Lancelet, 1210, 1220.
lanceolaris (Limax), 1220.
lanceolatus (Ammodytes), 568, 569, 570.
lanceolatus (Amphioxus, Branchiostoma, Limax),
1210, 1220.

lanceolatus (Gobius), 24.

Lancelet, 1, 1220.

Lancelet-fishes, 1210.

Landroding, 847.

Langsdorffii (Cyprinus), 733.

Lansettfisk, 1220.

Lantern-tisli, 428.

Lapplandsroding, 830, 831.
lapponicus (Coregonus), 898.
larvata (Cobilis), 706.
laskyr (Blicca, Cyprinus), 803.
laterna (Arnoglossus, Platophrys, Pleuronectes),
426, 428, 429.
laticeps (Gobius), 258.
latidens (Mierostomus), 383.
latifrons (Anarrhichas), 232, 237.
latifrons (Cyprinus, Leuciscus), 770.
latirostris (Acipenser), 1057.
latirostris (Anguilla, Murasna), 1023, 1024.
Latulus, 974.
latus (Cyprinus), 812.
latus (Salmo Thymallus), 898.

Launce, 569, 574.
lauta (Lampris), 124.

Lavaret, Lavaretus, 898.
lavaretus (Coregonus, Salmo), 891, 898, 900,
904, 907.

Lax, 831, 833, 849.

Laxbadd, Laxbudd, 757.

Laxesturje, 125.

Laxgard, 859.

Laxmina, 859.

Lax-ockel, 832.

Lax-roding, 848.

Lax-unge, 832.

Lax-yngel, 833.

Lax-oring, 831, 832, 860.
leachianus (Scomber), 93.

Leacliii (Clupea), 954.

Lebetus, 259.

Leiche, 1167.

leioglossa (Argentina), 912, 917.

Leister, 1010.

Leiter, 719, 720, 816.
leiurus (Gasterosteus), 648.

Lekmort, 776.

Lemani lacus (Salmo), 841.
lemanus (Fario, Salmo), 851.

Lemniscati, 1020.
leopardus (Anarrhichas), 236.

Leotardi (Pleuronectes), 428.

Lepadogaster, 301.

Lepibema, 44.

Lepidocephaliclithys, 7 02.

Lepidocephalus, 702.

Lepidoleprus, 585.

Lepidonote, 230.

Lepidopleurini, 71.

Lepidopus, 567.

Lepidorhombus, 426, 447.

Lepidosteus, 718, 829, 946, 1044, 1045.
Leptagonus, 206.

Leptoblennius, 224.

Leptocardii, 1063, 1210.

Leptocephalidas, 1020.

Leptocephalus, 1020, 1037, 1038, 1041.
leptocephalus (Gadus), 500.

Leptoclinus, 224.

Lepturus, 314.
lepturus (Caranx), 85.
lepusculus (Squalius), 7 60.

Lerbleking, 504, 506.

Lerflundra, 421.

Lerstubb, 256.

Lesueurii (Gobius), 248, 249, 250.
Leucabramis, 790, 798.

Leucaspius, 752, 786
leucichthys (Salmo), 890.

Leuciscina?, 720, 752, 759.

Leucisco-Blicca, 807.

Leuciscus, 719, 720, 723, 752, 759, 760, 778
leuciscus (Cyprinus, Squalius), 760.

Leuckartii (Abramidopsis, Abramis), 817.
leucophasus (Conger), 1038.
levenensis (Salmo), 851.

Lichia, 82, 85.

Lichtensteiuii (Acipenser), 1057.

Lilljeborgii (Cottus), 169, 191.

Limanda, 386.

limanda (Hippoglossoides, Hippoglossus), 421
limanda (Pleuronectes), 378, 386, 407.
Limandelle, 451.

limandoides (Hippoglossoides, Platessa, Pleuro
nectes), 421, 429.

Limax, 1220.
limosa (Myxine), 1208.

Lindare. 748.
linearis (Gobius), 269.
linearis (Porobranchus), 600.
lineata (Achirus, Solea), 364.
lineata (Liparis), 287.
lineatus (Beryx), 68.
lineatus (Cantharus, Sparus), 54.
lineatus (Cyclopterus, Liparis), 284.
lineatus (Cyprinus), 733.
lineatus (Exocoetus), 358.
lineatus (Labrus), 7, 10.

•lineatus (Lepadogaster), 303.
lineatus (Roccus), 44, 45, 46.

Ling, 230, 526.
linguatula (Citharus), 371.
linguatula (Pleuronectes), 421.

Linnei (Acanthias), 1158.

Linnei (iEglefinus), 466.

Linnei (Alburnus), 792.

Linnei (Anguilla), 1023.

Linnei (Aspius), 783.

Linnei (Cantharus), 54.

Linnei (Carassius, Cyprinus), 736.

Linnei (Conger), 1038.

Linnei (Galeus), 1133.

Linnei (Hippoglossus), 410.

Linnei (Labrax), 45.

Linnei (Lota), 532.

Linnei (Merlangus), 487.

Linnei (Merluccius), 516.

Linnei (Molva), 526.

Linnei (Pollacliius), 504.

Linnei (Solea), 373.

Linnei (Spinacliia), 638.

Linnei (Spinax), 1163.

Linnei (Tinea), 748.

Linnei (Thynnus), 98.

Linnei (Trachurus), 86.

lintea (Leucoraja, Raja), 1103, 1117.

lioderma (Pleuronectes), 441.

Liparis, 157, 283.

Liparis nostras, 287.

liparis (Cyclogaster, Cyclopterus), 284, 287.
liparoides (Cyclopterus, Liparis), 284.
Lip-fisli, 3.

Liza, 331.

Ljuster, 1010, 1034.

Lloydii (Coregonus), 899.

Loach, 711.

Loach, Spined, 706.

Lodda, 876, 882.

Loligo sagittata, 120.

longa (Guaperua), 634.

longibarbis (Macrurus), 583.

longifilis (Macrurus, Neinatonurus), 581, 583

longipinnis (Brama), 76, 80, 429.

longissimus (Batistes), 634.

Long-line, 1007.

155

Scandinavian Fishes.

1232

longus (Asellus), 526.

Lophiid®, 126, 136, 1012.

Lophius, 138.

Lophobranchii, 2, 661, 1043.

Lophonectes, 426.

Lophotes, 309, 320.

Lopliotidfe, 309.

Lophotiformes, 309.

Loricaria, 693.

loricatus (Gasterosteus), 648.

Lortbuk, 757.

lota (Gadus, Lota, Lotta), 531, 532.

Lotella, 521.

lotharingus (Gasterosteus), 658.

Lotinre, 464.

Lotta, 464, 520, 530.

Lubb, 519, 562.

lubb (Gadus), 562.

lubricum (Brancliiostoma), 1220.

Lucerna, 200.

lucerna (Trigla), 194, 199, 200.

Lueifuga, 462.

lucidus (Alburnus), 719, 720, 792.
Lucioperca, 36.

lucioperca (Perea, Stizostedium), 36
Lucius, 823, 998.
lucius (Esox), 998, 999.

Luddhvarf, 458.

Luden Hvarf, 456.
lugubris (Lvcodes), 613.
lumbricalis (Petromyzon), 1189.
lumbriciformis (Neropliis, Ophidion, Scyphius,
Syngnathus), 666, 683, 686.

Lumpen (Mustela), 603.

Lumpenus, 213, 224.

lumpenus (Blennius, Centronotus), 225.

Lumpenus, Blunt-Tailed, 225.

Lumpenus, Sharp-Tailed, 228.

Lumpfisk, 283.

Lump-Sucker, 294.

lumpus (Cyclopterus), 294.

luna (Chrysotosus, Lampris, Zeus), 123.

lunatus (Platophrj's), 430.

lupus (Anarrhichas), 232, 237, 608.

lupus (Centropomus, Labrax), 45.

Lupus marinus, 234.
luscus (Labrus), 5.

luscus (Gadulus, Gadus), 466, 467, 493, 495.
luscus (Platessa, Pleuronectes), 399.
lutescens (Gobio), 743.

Liitkenii (Lycodes), 609, 612.

Luzzo, 998.

Lycengraulis, 991.

Lycodalepis, 609.

Lycodes, 602, 607.

Lycodidse, 211, 463, 602.

Lycostomus, 952.
lycostomus (Gadus), 504.

Lycothrissa, 991.
lymma (Trygon), 1099.

Lyomeri, 1022.

Lyr, 506.

lyra (Callionymus, Uranoscopus), 272, 273,
279, 280.
lyra (Trigla), 195.

Lyrbleck, 506.

Lyrtorsk, 499, 504, 506.

Lysi, 519.

Lysing, 515, 516, 518.

Lysipharyngii, 2, 23.
lysan (Chorinemus), 83.

Langa, 526.

Langebarn, 230, 530.

Langebarn, spetsstjertadt, 225.

Langebarn, trubbstjertadt, 228.

Langhala, 822.
Langstjert, 822.
Langstjertgadda, 1005.
Lannare, 767.

Lodga, 832.

Lofgadda, 1005.
Lofmort, 776.

Loga, 794.

Loja, 720, 792, 794.

Ill ace r (Polyprosopus), 1144.

Machaarium, 595.

Mackenzii (Salmo), 890.

Mackerel, 110.

Mackerel, Frigate, 108.

Mackerel, Spanish, 1.

Mackerel-guide, 351.
macrocephala (Murrena), 1023.
macrocephalus (Gadus), 464, 473, 474, 475.
macrochir (Macrurus), 584.

Macroeutri, 717.

macrolepidotus (Pleuronectes), 370.
macrophthalma (Phycis, Molua), 521.
macroplithalmus (Coregonus), 899.
macrophthalmus (Cyprinus), 733.
macroplithalmus (Gobiesox), 300.
macrophthalmus (Onos), 544.
macrophthalmus (Scardinius), 779.
macrops (Lycodes), 608.
macropterus (Echelus), 1037.
macropterus (Thynnus), 91.
macrorhynchus (Laeviraja. Raja), 1123, 1125.
Macrouridaa, 581.
macrourus (Alopias), 1137.

Macruridaa, 461, 463, 539, 580.

Macrurus, 583, 1125.
maculata (Motella, Onos), 550, 551.
maculata, var. (Perea fluviatilis), 29.
maculatus (Balistes), 633, 634.
maculatus (Barbus), 718.
maculatus (Bothus, Rhombus), 366, 432.
maculatus (Callionymus), 272, 279.
maculatus (Clinus, Ctenodon, Leptoclinus, Lum-
penus, Stichaaus), 228.
maculatus (Labrus), 7.
maculatus (Lepadogaster), 303.
maculatus (Liparis), 284, 286.
maculosa (Gadus, Lota, Molva), 532.
maculosa (Perea), 18.
maculosus (Petromyzon), 1183.

Madui-marane, 902.

Mseoticus (Bothus), 437.

Maiforel, 838.

Maigre, 50.

mainensis (Gasterosteus), 658.
majalis (Clupea harengus), 959.
majalis (Leuciscus), 760.

Majfisk, 984.

major (Barbus), 540.

major (Clupea), 985.

major (Galeus stellar is), 1154.

major (Liparis) 287.

major (Molva), 526.

major (Perea), 26.

major vulgaris (Catulus), 1154.

Makrilgadda, 353.

Makrilstorje, 97, 125.

Mai, Europeisk, 693.
malabarica (Scorpaana), 154.

Malacanthidae, 127.

Malacocephalus, 583, 584.

Malacopterygii, 341.

Malncopterygii abdominales, 689.
Malacopterygii apodes, 567, 689.

Malacopterygii thoracici, 602.

Malapterurin®, 1086.

Malapterurus, 691, 692.

Malarmat, 193.

malarmoides (Aspidopliorus), 206.

Mallotus, 828, 875, 891.

Mancopsetta, 426.

Man-eaters, 1129.

manocentri (Coregoni lavareti), 902.
marsena (Coregonus, Salmo), 898, 899, 905, 907.
Maraane, 909.
maraanula (Salmo), 894.

Mareflundra, 385.

Mareflynder, 425.

Marena, 894.

Mareskadda, 398.
marginata (Anguilla), 1023.
marginata (Raja), 1117.
marginatum (Ophidium), 598.
marginatus (Conger), 1038.

Marieskadda, 398.
marina (Perea), 149.
marinus (Canis), 1167.

marinus (Lampetra, Lampreta, Petromyzon),
1182, 1183.
marinus (Sebastes), 148.

Mario, 1056.

marmorata (Torpedo), 1086.
marmorata, var. (Pleuronectes), 402.
marmoratus (Antennarius, Chironectes), 146.
marmoratus (Salmo), 851.

Marsipobranchii, 1043, 1172.
marsupiales, Syngnathi, 667, 679.

Marulk, 138.

Mastacembelidaa, 211.

Mastacembeliformes, 343.

Mastacembelus, 344.

Matfar, 558, 561.

Matjes, 968.

Maurolicus, 931.

maxillaris (Coregonus), 898, 902, 905.
maximus (Bothus, Pleuronectes, Rhombus),
427, 433, 434, 442.
maximus (Catulus), 1152.
maximus (Cetorhinus, Selache, Selaclius, Squa-
lus), 1136, 1143.
maximus (Hippoglossus), 410.
maximus hybridus (Bothus), 433, 444.
mediorostris (Anguilla), 1023, 1024.
mediterranea (Chimaara), 1080.
mediterranea (Julis), 21.
mediterranea (Meletta), 976.
mediterraneus (Gadus, Onos), 551, 556.
mediterraneus (Phycis), 539.
mediterraneus (Scomber), 105.
mediterraneus (Thynnus), 98.
mediterraneus (Trachurus), 88.
medius (Lumpenus), 225.

Meer-att, 1079.

megalops (Coregonus), 898, 904.
megastoma (Arnoglossus, Lepidorhombus, Pleu-
ronectes, Rhombus, Zeugopterus), 447, 448.
Megrim, 428.

Melandrys, 99.
melanochir (Anguilla), 1023.
melanogaster (Pseudorhombus), 366.
Melanogrammus, 467.
melanostomus (Gadus), 511.
malanostomus, melastomus (Galeus, Pristiurus,
Scyllium), 1125, 1149.
melanotus (Cyprinus, Idus), 724, 764.
melanura (Julis), 21.

Melet, 992.

Meletta, Melettus, 976.
meletta (Engraulis), 992.

1233

Melichthys, 631.

inelops (Crenilabrus, Labrus), 4, 18.
membranaceus (Paraliparis), 283.

Membras, 952.

membras (Clupea harengus), 959.

Menidse, 52.

meuto (Exocoetus), 355.

Menuaggia, 432.
meridionalis (Aphia), 266.
nieridionalis (Sqalius), 770.

Merkii (Coregonus), 895.

Merlangus, 487.

merlangns (Gadus), 464, 466, 467, 487, 511.
Merlu barbu, 540.
inerluccius (Merlucius), 515.

Merluciime, 464.

Merlucius, 464, 514, 515.

Merlus 515.

Merluzzo, 518.

Mertensii (Thymallus), 883.

Merulk, 138.

messanensis (Cicerellus), 573.

Messingsgadda, 1005.

Meunier, 772.

microcephalus (Acanthorhinus, Lasmargus,Scym-
nus, Somniosus, Squalus), 1127, 1168.
microcephalus (Coregonus), 899, 904, 906, 907.
microcephalus (Gasterosteus), 648.
microcephalus (Platessa, Pleuroneetes), 378,
383.

microdon (Osmerus), 876.

microlepidotus (Abramis), 812.

microlepidotus (Cyprinus), 721, 764.

microlepis (Leuciscus), 753.

microlepis (Thymallus), 882.

microps (Coregonus), 906, 907.

microps (Cottunculus), 158, 613.

microps (Gobius), 244, 255, 256, 257, 258.

microps (Salmo), 851.

microptera (Anguilla), 1023.

micropterus (Scymnus), 1168.

micropteryx (Abramis), 803.

micropus (Liparis), 292.

microstoma (Crenilabrus), 14.

microstomus (Pleuroneetes), 383.

microstomus (Salmo), 899.

migratoria (Anguilla), 1023.

milvus (Trigla), 197.

Mina, 859."
miniatus (Idus), 764.
minimus (Gadus), 559.

Minnow, 720, 754.
minor (Albula), 792.
minor (Anarrhichas), 232, 236.
minor (Couchia), 554.
minor (Harengus), 979.
minor (Perea), 41.
minor vulgaris (Catulus), 1154.
minuta (Aphya, Atherina), 265, 266, 432.
minutus (Cyclopterus), 294.
minutus (Gadulus, Gadus), 466, 467, 493,
495, 508.

minutus (Gobius), 243, 244, 251, 255, 257,
258, 262, 294, 425.
mirabilis (Clupea), 954.
miraletus (Raja), 1112.
mirandella (Alburnus), 792.

Mirbelia, 301, 303.

Mirbelii (Lepadogaster), 303.

Misgurnus, 704.
mistops (Salmo), 851.
mixtus (Labrus), 4, 6, 10.

Mocka, 859.

Mogki (Rhomboidichthys), 427.

Mola, 625.

mola (Diodon, Orthagoriscus, Tetraodon, Tetro-
don), 622, 625, 975.
moles (Cyprinus), 736.

Molina, 622.

Molua, 464, 520.

Molva, 520.

molva (Gadus, Lota, Molua, Molva), 521, 525,
526, 528, 531.

Monacanthi na3, 631.
monensis (Squalus), 1139.

Monflundra, 385.

Monkey-fish, 1079.

Monster-fish, 1079.

monstrosa (Chimaera), 1065, 1066, 1078, 1079.
1204.

Montagui (Cyclopterus, Liparis), 284.
Mordacia, 1182.

mordax (Argentina, Osmerus), 869, 872.
morena (Anguilla), 1023.
morhua (Gadus), 472.

Moringua, 1022.

Mormyridae, 1054, 1086.
morinyrus (Sparus), 56, 57.

Morone, 44.

Morrhua, 520.
morrhua (Gadus), 472.

Morris, 1038.

Morrisii (Leptocephalus), 1021, 1038.

Mort, 503.

Mo rue, 520.

Motella, 543.

Motellinae, 464.
mucosa (Leviraja), 1125
mucosus (Cyclogaster, Neoliparis), 286, 293.
mucosus (Lycodes), 608, 609, 612.
mucronata (Raja), 1125.

Mudd, 757.

Mudderskrubbe, 403.

Mud-eel, 1204.

Mud-minnows, 997.

Mugil, 328, 330, 342, 891.

Mugilidfe, 126, 327, 328.

Mugiliformes, 126, 327.

Muksun, 898, 899, 902.
muksun (Coregonus), 905, 907.

Mullet, Golden, 337.

Mullet, Red, 63.

Mullet, Thick-lipped grey, 334.

Mullet, Thin-lipped grey, 339.

Mullidse, 25, 61.

Mulloides, 61.

Mullus, 62.

multibarbata (Brotula), 595.
multidentatus (Labrus, Crenilabrus), 7, 8.
Mulus, 62.

Munflundra, 385.

Mur sen a, 1013, 1021, 1023.
murtena (Lycodes), 609, 617.

Mursenesox, 1022.

Muraenidse, 1021.

Muraenolepis, 461.

Murrayi (Ipnops), 923.

Mustela, 1183, 1188.

mustela (Gadus, Motella, Onos), 544, 545, 554.
mustelaris (Blennius), 219.
mustelaris (Gadus), 554.
mustella (Gadus, Onos), 550, 551.

Mustelus, 1128, 1142.

muticellus (Leuciscus, Telestes), 753.

Mutterloschen, 787.

Myctophum, 936.

Myliobatidse, 1071, 1093.

Myliobatis, 1094.

Mulleri (Maurolicus, Salmo), 922, 923, 931, 942.
Muller's Topknot, 458.

myrus (Murtena), 1037.

Myxina, 1208.

Myxine, 1195, 1196, 1208.

Myxinidae, 1195.

Myxus, 328.

Mort, 720, 773.

Nadd-id, Naddi, 770.
naevus (Raja), 1112.
namaycush (Salmo), 845.
narce (Torpedo), 1086.

Nasello, 518.

Nasus, 799.

nasus (Coregonus, Salmo), 899.
nasus (Diplanchias, Mola), 626.
nasus (Squalus), 1139.

Naucrates, 82, 83.

navaga (Gadus), 465, 467, 481, 514.
nebulosus (Gasterosteus), 658.

Needle-fishes, 667.

Nejonoga, 1195.

Nelma, Njelma, 875, 890.
nelma (Salmo, Stenodus), 890.

Nemachilus, 704.

Nematognathi, 690.

Nemichthyidfe, 1022.

Neoliparis, 286.

Nerophini, 679.

Nerophis, 679.

nesogailicus (Chironectes), 146.

Nestis, 328.

Neunauge, 1195.

Neustria? (Labrus), 7.
neustrianus (Gasterosteus), 648.
nidrosiensis (Raja), 1064, 1103, 1123.
niger (Astronesthes), 921.
niger (Conger, Muraena), 1037.
niger (Gasterosteus), 648.
niger (Gobius), 241, 244, 245, 255, 343
461.

niger (Petromyzon), 1189.
niger (Raniceps), 559.
niger (Spinax), 1125, 1163.
nigricans (Petromyzon), 1183.
nigripes (Trigla), 197, 201.
nigripinnis (Salmo), 851.
nigro-auratus (Cyprinus), 724.
nigromanus (Pleuroneetes), 379.
nilotica (Anguilla), 1023.

Nilsonii (Crystallogobius, Gobiosoina, Gobius
Latrunculodes, Latrunculus), 269.

Nilssonii (Coregonus), 892, 898, 904, 907.

Nissoga, 706.

nobiliana (Torpedo), 1086.

nobilior (Esox), 1003.

nobilis (Albula), 898.

nobilis (Salmo), 850.

noctula (Myliobatis), 1095.

Nonnat, 264, 432.

Nordmanni (Coregonus), 899.

Nordmanni (Cyprinus), 724.
norvegicus (Crenilabrus, Labrus, Lutjanus), 18
norvegicus (Holocentrus, Perea, Sebastes), 148
149.

norvegicus (Lepidorhombus, Rhombus, Sco
phthalmus, Zeugopterus), 427, 452, 453.
norwegianus (Acanthorhinus), 1168.

Nors, 868, 869.

Norway Haddock, 148.

Nostras (Liparis), 287.

Notidanida?, 1085, 1128, 1156.

Notopterida?, 829.

Notopterus, 947.

Nounat, 264.

1234

Novaga, 481.

nov®orleanensis (Anguilla), 1023.
nov®terr® (Anguilla), 1023.
nov®-Zelandi® (Macronurus), 581.
noveboracensis (Gasterosteus), 648.
nuda (Orcynopsis, Pelamys), 103, 104.
nudus (Cyclopterus, Gobiesox), 300.
nudus (Cyprinus carpio), 724.
uukta (Labeo), 716.

Nurse-Hound, 1152.

Nyctophus, 936.

Nalle, 503.

Nabbgadda, 347.

Nfibbsik, 900, 902.

Natting, 1188.

Obesus (Cyprinus), 724.
oblongirostris (Anguilla), 1023.
oblongus (Cyprinus), 736.
obolarius (Gasterosteus), 648.
obtusirostris (Saliuo), 882.
obtusus (Alburnus), 792.
occidentalis (Conger), 1038.
occidentalis (Gasterosteus), 658.
oceanica (Clupea harengus), 959.
oceanica (Limanda), 386.
ocellata (Torpedo), 1086.
ocellaris (Blennius), 212, 219.
ocellatus (Lepadogaster), 303.
ochrodon (Leuciscus), 792.
ocla (Salino), 850.
ogac (Gadus), 464, 465, 467, 479.
ogat, 480.

Olfersii (Argyropelecus, Pleurothyris, Sterno-
ptyx), 921, 923, 925.

Omalii (Petromyzon), 1189.

Ombre, 841.

Ombrina, 50.

Onchus, 1085.

Oncorlivnclius, 830, 839, 854.

Onion-fish, 589.

onitis (Labrus, Tautoga), 11.

Onomorphi, 461.

Onos 464, 515, 518, 543, 567.
ontariensis (Tliymallus), 883.

Opah, 123.

ophidii, Syngnathi, 667, 679.

Ophidiid®, 461, 463, 567, 594.

Ophidiin®, 595.

Ophidium, 595.

ophidion (Nerophis, Soyphius, Syngnathus), 666,
680, 683, 686.

Ophiocephalid®, 328.

Opliiur®, 425.

Ojfistomi, 211.

Oppror, 1010.
orbicularis (Cantharus), 54.
orca (Gobius), 244, 259, 260.
orcadensis (Salmo), 851.

Orcynopsis, 91, 102.

Orcynus, 91, 92, 96, 99.

Oreosoma, 305.

Oreosomatid®, 305.

orfus (Cyprinus), 764, 767.

orientalis (Thynnus), 91, 98.

Ormfisk, 711.

Orsini (Ozodur i), 626.

Ort , 768.

Orthagoriscid®, 622.

Orthagoriscus, Orthragoriscus, 622, 625.
Orthocormi, 24.

Osmeroides, 827, 912.

Osinerus, 827, 828, 863, 866, 867, 869, 875.
ossifagus (Labrus), 10.

Osteoglossum, 947.

Ostracion, 1012.

Ostraciontina, 622.

Otolithus, 51.
ouak, 479.

Outil, 909.
ovak, 479.
ovis (Sargus), 53.

Owsianka, 754, 787.
owsianka (Aspius), 787.

Oxsimpa, 187.

oxygeneios (Epinephelus), 48.
oxyrhiua (Mur®na), 1023.

Oxyrhynchus, 898, 902.
oxyrhynchus (Acipenser), 1057.
oxyrhynchus (Coregonus), 902, 905.
oxyrhynchus (L®viraja, Raja), 1103, 1125.
oxyrhynchus (Salmo obtusirostris), 882, 899,
902, 905.

oxyrinchus (Isurus), 1139.
oxyrinchus (Raja), 1117.

Ozodura, 626.

ozodura (Orthagoriscus), 626.

Ozzone, 331.

Pacifici (Batrachus), 133, 135.
pacificus (Cynicoglossus), 386.
pacificus (Thaleichthys), 875.
pacificus (Thynnus), 91.

Paddtorsk, 558.

Paddulk, 158.

Paddal, 561.

Pagellus, 56.

Pagrus, 56.

Pagurus, 425.

Pala, Palla, 898.

Pale, 503.

Pal®onotidani, 1128.
palea (Coregonus), 898.

Pallasii (Clupea), 954.

Pallasii (Pleuronectes), 392, 395.

Pallasii (Thymnllus), 883.
palleus (Leuciscus), 774.
pallidus (Lycodes), 612, 613.
pallidus (Salmo), 842.

Pallonii (Labrus), 4, 5.

palmicornis (Blennius, Chirolophis), 218.

Panka, 806, 815.

pantherinus (Anarrhichas), 236.

papalina (Alosa), 976.

Paralepidin®, 923, 936, 945, 997.

Paralepis, 945.

Paralichthys, 366, 408, 420.

Paraliparis, 283.
parallelus (Macrurus), 583.

Pareti (Leuciscus), 770.

Parexocoetus, 356.

Parnelli (Gobius), 261.

Parophrys, 377.
paroticus (Labrus), 21.

Parr, 833, 855.
parva (Albula), 898.
parvipinnis (Fierasfer), 600.

Pasciuti, 1033.

passer (Flesus, Platessa, Pleuronectes), 398,
399, 441.

Passer fluviatilis, 402.

Pastinaca, 1098.

pastinaca, (Raja, Trygon), 1097, 1098.
Pastinac®, 1097.

Pata, 859.

patris (Acanthocottus) 160.

Pausingeri (Leuciscus), 774, 777.
pavonina (Pleuronectes), 441.

pavoninus (Cyclopterus), 294.

Pearl-Side, Boreal, 931.
peckianus (Syngnathus), 664.

Pediculati, 136.

pediculati, Acanthoptherygii, 24.
pediculati, Anomali, 24.

Pejsa, 837.

pekinensis (Carassius), 733.
pelagicus (Lampris, Scomber), 123.
pelagicus (Syngnathus), 672, 674.

Pelainis, 99.

Pelamis, Plain, 103.

Pelamis, Stripe-backed, 105.

pelamis (Euthynnus, Scomber), 91, 95.

pelamis (Orcynus, Sarda), 92, 105.

Pelamys, 104.

pelamys (Euthynnus, Thynnus), 95.
pelamys (Sarda), 105.

Pelecus, 721, 790, 822.
pelegrinus (Squalus), 1144.

Pelerin (Squale), 1144.

Pelet, 891.

pellucidus (Aphia, Apliya, Gobius, Latrunculus),
266.

pellucidus (Thyris), 363.

Pennanti (Argentina, Maurolicus, Scopelus), 932.
Pennantii (Crenilabrus), 18.

Perea, 25, 26.

Percesoces, 328.

Perch, 26, 44.

Perch, Carass, 29.

Perch, Deep-water, 30.

Perch, Leaf, 30.

Perch, Mud, 30.

Perch, Pike, 36.

Perch, Sea, 45.

Perch, Sedge, 30.

Perch, Spotted, 29.

Perch, Stone, 30.

Perches, Sea-Pike, 49.

Percid®, 25.

Percin®, 25.

Percis, 127.

Percomorphi, 25.
peregrinus (Cetorhinus), 1144.
peregrinus (Thynnus), 103.

Periophthalmus, 240.

Peristedion, 193.

Permuck, 554, 557.
personatus (Ammodytes), 577.
perspicillum (Lycodes), 608, 612, 615.

Peruk, 1107.

Petromyzon, 1172, 1182.

Petromyzonid®, 1172.

phalerica (Aphya, Clupanodon), 976.

Pharyngognathi, 2.

Pharyngognathi malacopterygii, 342.

Phobetor, 159.

Pholis, 213, 220.
pholis (Blennius), 214.

Phoxinus, 752, 753.

phoxinus (Cyprinus, Leuciscus), 721, 754.
Phrynorhombns, 452.

Phycis, 464, 531, 538.
phycis (Blennius), 539.

Physiculus, 538.

Phj'soclysts, 1, 689.

Physoclysti hemibranchii, 635.

Physoclysti lophobranchii, 660.

Physoclysti plectognathi, 618.

Physostomi, 689.

Picked Dog Fish, 1158.

pictus (Chironectes), 146.

pictus (Gobius), 241, 244, 255, 257, 258.

pictus (Dranoscopus), 272, 273.

Pifvare, 441.

Pigg, 657.

Pigghaj, 1158, 1161.

Pigghvarf, 368, 434, 441.

Pigghvarfvel, 441.

Piggha, 1161.

Piggmakrel, 87.

Piggrocka, 1107.

Piggvarr 441.

Pihral, 1208.

Pike, 998.

Pike, Brass, 1005.

Pike, Bony, 1044.

Pike, Flower, 1005.

Pike, Gertrude’s, 1005.

Pike, Grass, 1005.

Pike, lee, 1005.

Pike, Leaf, 1005.

Pike, Long-tailed, 1005.

Pike, Meadow, 1005.

Pike, Saury, 353.

Pike, Short -tailed, 1005.

Pike-net, 1007.

Pike-spear, 1010.

Pilchard, 979.

Pilchardus, 979.

pilchardus (Alausa, Alosa, Clupea), 952, 979.
Pilot-fish, 83, 1130.

Pilal, Piral, 1208.

Pingelii (Triglops), 167.

Pinguipes, 127.

pinguis (Hippoglossus, Platysomatichthys,
Pleuronectes), 417.
pini (Trigla), 194, 195.

Pipefish, iEquoreal, 680.

Pipefish, Deep-nosed, 674.

Pipefish, Great, 668.

Pipefish, Lesser, 672.

Pipefish, Straight-nosed, 683.

Pipefish, Worm, 686.
piscatorius (Lophius), 136, 138.
piscatrix (Lophius, Rana), 138, 139.
Pisciculus varius, 754.

Pisoodonophis, 1022.

pistilliger (Cottus, Gynmocanthus, Phobetor).
160, 161.

Pjon, 794.

Placoidei, 1043.

Placodermi, 1044.

Plagiostomi, 1085.

Plagiostomi batoidei, 1086.

Plagiostomi selachoidei, 1127.

Plagusia, 363.

Plaice, 392.

planci (Tympanomium), 626.

Planeri (Petromyzon), 1179, 1189, 1191, 1193.
planifrons (Batrachus), 135.

Plank, 506.

Platessa, 392.

platessa (Pleuronectes), 365, 378, 392, 401, 407.
platessoides (Citharus, Drepanopsetta, Hippo-
glossoides, Pleuronectes), 376, 387, 408,
421, 429, 454.

Platophrys, 364, 426, 427.
platycephala (Anguilla), 1023.
platycephalus (Cottus), 181, 182.
platyrhina (Mursena), 1023.
platyrhynchus (Anguilla), 1023.
platyschistse (Munenidaa), 1022.
Platysomatichthys, 408, 416.
plebejus (Gasterosteus), 648.

Plectognates, 618, 619, 1043.

Plectognathi, 1, 344.

Plectospondyli, 690, 702.

Plestya, 803.

plestya (Cyprinus), 803.

Pleuracauthus, 1066.

Pleuronectes, 377, 378.

Pleuronectidte, 366, 377.

Pleuronectina, 371, 377.

Pleuronectini, 370.
pleurostictus (Triglops), 167.

Pleurothyris, 925.

Plogjernsrocka, 1125.
plotizza (Scardinius), 779.

Plotosus, 690, 692.

Pluddennun, 385.
plumbea (Chimaera), 1078, 1080.
plumbeus (Petromyzon), 1189, 1193.
pneumatophorus (Scomber), 110.
podas (Rhomboidichthys), 366.

Podothecus, 206.
poeciloptera (Trigla), 195, 201.
pcecilopus (Cottus), 169, 173.

Pogonias, 49.

pola (Platessa, Pleuronectes), 379, 383.
polaris (Boreogadus, Gadus), 484.

Pol cur, 891, 898, 899, 902.
polcur (Coregonus), 891, 899, 904, 906, 907.
Pole, 378.

Pollachius, 504.

pollachius (Gadus, Merlangus), 466, 467, 504.
Pollack, 499, 504.

Pollack, Norwegian Whiting-, 508.
pollan (Coregonus), 896.

Pollinii (Gobio), 743.

Polycentrus, 11.

Polyodontid®, 1044.
polyosteus (Salmo), 851.

Polyprion, 25, 47.

Polyprosopus, 1144.

Polypterus, 343, 946, 1012, 1044.
Pomatomus, 82.
pontica (Raja), 1104.
ponticus (Gasterosteus), 648.

Pope, 41, 44.

Porbeagle, 1069, 1138.

Porcellus, 41.

Porichthys, 133.

Porobranchus, 600.
porosus (Cottus), 180.

Potta, 441.

Poutassou, 511.

poutassou (Boreogadus, Gadus, Merlangus),
465, 466, 467, 511.

Powan, 909.
prsestabilis (Alosa), 985.

Prfflstefiyndre, 396.
prasinus (Leuciscus), 773, 777.
princeps (Brama), 80.
pricka (Petromyzon), 1189.

Pride, 1189, 1194.

Prionodon, 1130.

Pristidas, 1093.

Pristiophoridas, 1128.

Pristiurus, 1148.
probatoccphalus (Diplodus), 53.
productus (Gadus, Merlucius), 464, 516.
Proppe, 786.

Protocampus, 667, 679.

Protochordata, 1210.
proximus (Gadus), 467, 490.

Psammodiscus, 377.

Psettichthys, 408.

Psettodes, 408.

pseudoflesus (Platessa), 393, 403.
pseudogastropteri, Orthocorrni, 24.
Pseudorhombus, 366, 408, 420.

Pteraclis, 70, 71.

Pterichthys, 1044.

Pteropliryne, 145.

Pterurus, 680.

Pterycombus, 70, 71, 72.

Ptyobranchidse, 1022.

Pucelle, 984.
pulcher (Scomber), 95.

Pulsvad, 1035.
pumila (Spratella), 974.
punctata (Labrus, Seisena), 1 1 .
punctatissima (Anguilla), 1023.
punctatus (Batrachus), 135.
punctatus (Dasybatis, Raja), 1104.
punctatus (Helmictis, Leptocephalus), 1038.
punctatus (Lepadogaster), 303.
punctatus (Pleuronectes, Rhombus, Zeugopterus),
383, 427, 456.
punctatus (Salmo), 851.
punctatus, var. (Scomber), 111.
punctulatus (Batrachus), 135.
pungio (Zeus), 308.

Pungitius, 638.

pungitius (Gasterosteus), 264, 658.
purava (Stolepborus), 991.
purpuratus (Salmo), 830.
purpureus (Gadus, Merlangus), 500.
pusillum (Acanthidium), 1163, 1166.
pusillus (Labrus), 7.

Putnami (Euchalarodus), 405.

Puttning, 1035.

pycnocentri (Coregoni lavareti), 899, 902.
Pycnodonta, 71.

Pycuodontoidei, 71.

Pyn, 794.

Padrifvare, 832.

Palkrok, 1008.

(Juadracus (Apeltes), 649.
quadricornis (Cottus), 169, 175.
quadricornis (Motella), 550, 553.
quadridens (Pleuronectes), 383.
quadrilateralis (Coregonus), 899.
quadrilobatus (Cyprinus), 733.
quadrimaculatus (Gobius), 261.
quadripunctatus (Scomber), 93.
quadrituberculatus (Pleuronectes), 392, 395.
quadrivittatus (Gobius), 263.

Quenselii (Pleuronectes), 383.

Querimana, 328.

quinquecirrhatus (Gadus, Motella), 554.
Qvabbso, 297.

Qveise, 131.

Qveite, 416.

Qvidd, 757.

Rabbit-fish, 1084.

Rabboxe, 893, 898.

Radflackiga Tungehvarfven, 428.
radiata (Amblyraja, Raja), 1088, 1103, 1108.
radiatus (Trachinus), 128.
radula (Raja), 1112.

Rrebus, Raspuschka, 898.

Raja, 1101.

Raji (Brama, Sparus), 76, 77, 80, 429.
Rajidae, 1100.

Raniceps, 464, 558.

raninus (Blennius, Gadus, Raniceps), 558, 559.
ranula (Liparis), 287.

Ranzania, 624.

rapax (Aspius), 720, 783.

raptor (Gadus, Molua), 526, 527.

Raschii (Brama), 80.
rashleighanus (Polyprosopus), 1144.

Rav, 415.

1236

Ray, Deep-sea, 1111.

Ray, Eagle, 1071, 1095.

Ray, Electric, 1071, 1086, 1092.

Ray, Sandy, 1089, 1112.

Ray, Starry, 1087, 1108.

Ray, Sting, 1097, 1098.

Rays, 1043, 1063, 1085, 1086, 1100, 1104.
Redi (Ortliagoriscus), 626.

Regalecus, 320.
regina (Cyprinus), 724.
regina (Rhodichthys), 596.
regulus (Sebastes), 150.

Reinhardi (Careproctus, Liparis), 292.
Reisingeri (Coregonns), 898.
remipes (Gymnetrus, Regalecus), 322.

Renke, 909.

repandus (Trachypterus), 313.

Repsen, 898.

resplendens (Lampanyctus, Scopelus), 937.
reticulatus (Callionymus), 279.
reticulatus (Lepadogaster), 303.
reticulatus (Liparis), 286.
reticulatus (Lycodes), 609, 611.

Retropinna, 827, 912.

Retzii (Ortliagoriscus), 626.

Rex Cyprinoruiii, 724.

Rhamphistoma, 344, 667.

Rhinidae, 1128.

Rhinobatida?, 1093.

Rhinodon, 1073, 1142.

Rbinolepis, 691.

Rliinomugil, 328, 333.

Rhodeus, 715.

Rhodichtbys, 596.
rhomboides (Rhombus), 452.

Rhomboidiclitbys, 364, 427.

Rhombus, 432.

rhombus (Bothus, Pleuronectes), 427, 433, 441.
rhombus hybridus (Bothus), 433, 445.
Rhynchichthys, 24.
riali (Onus), 515.

Richardsonii (Coregonus), 898.

Richardsonii (Retropinna), 827, 912.

Ringbug, 283.

Ringbuyk, 283.
ringens (Balistes), 633.

River Trout, 856.
rivularis (Cyprinus), 754.

Roach, 720^ 773.

Roach, Gold, 777.

Roach, Ice, 776.

robustus (Clypeocottus), 187.

Roccus, 25, 44.
rocheanus (Thynnus), 108.

Rochei (Auxis, Scomber), 108.

Rockhok, 1107.

Rock-cook, 14.

Rockling, Five-Bearded, 554.

Rockling, Four-Bearded, 544.

Rockling, Northern, 548.

Rockling, Three-Bearded, 550.
rodens (Leuciscus, Squalius), 760, 761.
Rogenia, 954.

Rogn-kal, 297.

Rogn-kexe, 297.

Rondeletii (Carcharodon), 1139.

Rondeletii (Exocoetus), 358.

Rondeletii (Orthragoriscus), 626.

Rondeletii (Scombresox), 353.

Rondeletii (Syngnathus), 677, 678.
rone (Labrus), 18.

Rootaug, 779.

Rose-fish, Rosenfisk, 596.

Rosenhaueri (Alburuus), 795.
roseus (Pleuronectes), 401.

Rossi (Lycodes), 612, 615.
rostellatus (Syngnathus), 666, 672, 674.
rostrata (Belone), 347.
rostrata (Mursena), 1023.
rostratus (Acanthorhinus), 1167, 1169.
rostratus (Cetorhinus, Squalus), 1144.
rostratus (Leuciscus, Squalius), 760, 761.
Rotskar, 503.

Rough Hound, 1154.

Rua, Ruda, 780.

Rubellus, 773.
ruber (Gadus), 473.
ruber (Petromyzon), 1189.
ruber (Urocentrus), 67.
rubescens (Syngnathus), 669, 671.
rubieundus (Acipenser), 1056.
rubro-fuscus (Cyprinus), 724.
rubus (Raja), 1104, 1112.

Ruda, 720, 735.

Rudabborre, 29.

Rudd, 720, 779.

Rudskalle, 780.

Ruff, 41.

rumulo (Bothus), 441.

rupestris (Coryphaenoides, Macrurus), 584, 588,
590.

rupestris (Crenilabrus, Ctenolabrus, Labrus,
Perea, Scimna), 4, 6, 16.

Ruslsa, 403.

Ruskeskadda, 403.

Russellii (Regalecus), 321.

Russnase, 801.

Ruthensparri (Gobius), 251.
ruthenus (Acipenser), 1051.
rutilo-bjorkna (Leucisco-Blicea), 807.
rutilo-blicca (Bliccopsis), 811.
rutiloides (Leuciscus), 773, 777.
rutilus (Cyprinus, Leuciscus), 186, 720, 759,
773, 775.

rutilus (Salmo), 842.

Ryner, 1147.

Ryssja, 1007.

Rabo Herring, 959.

Ranka, 297.

Rafhaj, 1136.

Raiding, 415.

Robug, 255.

Rodfena, 781.

Rodfisk, 152.

Rodhajar, 1152.

Roding, 832, 834, 841, 853.

Roding, Lapplands, 830.

Roaing, Vetterns, 830.

Rodknot, 195.

Rodmort, 781.

Rodnabba, 14.

Rodsnacka, 14.

Rodspatta, 369, 392.

Rodspatta, 398.

Rodtorsk, 477.

Roe, 846.

Rofisk, 831.

Rotsimpa, 180.

Sabbik, 262.

Saccobrancliinas, 692.

Saccobranchus, 691.

Saccopharyngidae, 1022.

Saelbling, 830.

Sagre, 1163.

Sagris, 352.

Saibling, 830.

saida (Boreogadus, Gadus), 463, 464, 465
467, 484, 514.

Sail-Fluke, 451.

Salanx, 827.

Salar, 841, 851.

salar (Salmo, Trutta), 830, 833, 849, 851.
Salarias, 212.

Salilota, 531.
salmarinus (Salmo), 842.

Salmlet, 851.

Salmling, 841.

Salmo, 828, 829, 841, 866, 867.
salmo (Salmo), 850.

Salmon, 829, 832, 833, 849, 851, 853, 883.
Salmon, Quinnat, 829.

Salmon, Rainbow, 830.

Salmon, Siberian White, 890.

Salmon, Spanish, 507.

Salmon stair, 860.
salmonata (Trutta), 851.
salmonea (Perea), 37.

Salmonidse, 827.

Salmons, 826.

Salmulus, 850.

salnuilus (Salmo), 832, 850, 855.

Salvelinus, 841.

salvelinus (Salmo), 830, 831, 835, 842, 867.
Salviani (Raja), 1125.

Sand-Eel, 570.

Sand-Eels, 567.

Sandflundra, 385, 386, 390.

Sandflundre, 403.

Sandflynder, 390.

Sandharr, 887.

Sandbvarf, 443.

Sandkrypare, 720, 743.

Sand-Launce, 574.

Sand-Launce, Smooth, 573.
sandra (Lucioperca), 36.

Sandskrubbe, 403.

Sandskadda, 390, 425.

Sandstubb, 262.

Sandtunga, 375.

Sandy Ray, 1112.
sanguisuga (Petromyzon), 1189.

Sanktepersfisk, 306.
sapidissima (Clupea), 985.

Sarachus, 823.

Sarda, 91, 102, 104.

sarda(Orcynus, Pelamys,Scomber,Thynnus),105.
Sardell, 992.

Sardine, 979.

sardina (Alosa, Clupea), 979.

Sarf, 720, 779, 781.

Sarfvel, 781.

Sargini, 53.

Sarsii (Lycodes), 609, 616.

Saurina?, 923, 925.

Saurus, 352, 353, 936.
saurus (Esox, Scombresox), 353.
saurus (Trachurus), 86.

Saussurii (Drama), 81.

Savignyi (Anguilla), 1023.

Sawfish, 1086.

saxicola (Glyptocephalus, Platessa, Pleuronectes),
379.

saxatilis (Canicula), 1152.

Sayi (Raja), 1099.
scaber (Chironectes), 146.
scabra (Raja), 1104.

Scad, 86.

Scaphiryn chops, 1054.

Scarabreus, 54.

Scar dine, 778.

Scardiniopsis, 795.

Scardinius, 719, 720. 752, 778.

Scardo-Blicca, 807.

scardofa (Scardinius), 779.

Scardola, 778.

Scarus, 2, 4.

Scharretong, 378.

schelderensis (Clupea hareugus), 959.
Schiefermiilleri (Salmo), 850.

Schilus, 36.

Schinzii (Coregonus), 899.
Schismatorhynclius, 716.

Schmupel, 909.

Schneider, 798.

Scholle, 398.

Schomburgkii (Polycentrus), 11.

Schoneveldi (Clupea), 974.

Scliusslaube, 798.

Schwebforell, 838.

Scimna, 50.

Sciaenidm, 25, 49.

Sclerodermes, 622.

Scleroparei, 146.

Scomber, 110.
scomber (Scomber), 111.

Scomberesoces, 342.

Scomberesox, 352.

Scomberomorus, 89, 91, 105.
Scombresocidis, 343.

Scombresociformes, 343.

Scombresox, 352.

Scombridse, 70, 89.

Scombroidea, 121.

Scombromorphi, 69.
scoinbrus (Scomber), 83, 92, 110.
Scopelkhe, 826, 827, 920, 923.
Scopelomorphi, 826.

Scopelus, 936.

Scopelns, Arctic, 941.

Scopelus, Greater, 937.

Scophthalmus, 452.

Scorpsena, 143, 146, 153.
scorpama (Cottus), 180.

Scorpfenidae, 126, 147.
scorpioides (Gobius), 244, 260.
scorpius (Cottus), 157, 169, 180.
scriptus, var. (Scomber), 112.
sculponeatus (Cyprinus), 724.
scutellatuin (Scombresox), 353.

Scylliidfe, 1128, 1 147.

Scylliorhinus, 1148, 1151.

Scyllium, 1152.

Scymuube, 1166.

Scymnus, 1066, 1156, 1166, 1167, 1169.
Seyphius, 683.

Sea-Bream, Black, 54.

Sea-Bream, Common, 59.

Sea-Bream, Rasch’s, 80.

Sea-Bream, Ray’s, 77.

Sea-Bream, Spanish, 58.

Sea-Cat, Blue, 237.

Sea-Cat, Common, 232.

Sea-Cat, Spotted, 236.

Sea-Pheasant, 440.

Sea Pike-Perches, 49.

Sea-Scorpion, 180.

Sea-Snail, 283.

Sea-Snail, Common, 287.

Sea-Snail, Montagu’s, 284.

Sea-Snail, Slimy, 291.
sebago (Salmo), 850.

Sebastes, 146, 148, 1 125.

Sebastodes, 147, 148.
secundo-dorsalis (Thynnus), 91, 98.

Sej, 500, 503.

Sejlyra, 506.

Selache, 1144.

Selachoidei, 1127.

Seldetka, 896.

Selene, 83, 85.

Selysii (Leuciscus), 773, 7 77.
semiarmatus (Gasterosteus), 648.
seiniloricatus (Gasterosteus), 648.
seminudus (Lycodes), 612.
semispinosus (Caranx), 86.
septentrionalis (Motella, Onos), 544, 548, 557.
septentrionalis (Mugil), 334.
septoeuil (Petromyzon), 1189.
serpentinus (Acus), 686.

Sevprda, 332.

Serrani, 3.

serrato-granulata (Perea), 27.
serratus (Gasterosteus), 648.

Setipinna, 990.
setipinnis (Selene), 83.

Seuruga, 1054.

Sewin, 856.

Shad, 983, 984, 988.

Shad, Allice, 984.

Shad, American, 985.

Shad, Twaite, 984.

Shadow-fish, 50.

Shanny, 215.

Shagreen Ray, 1115.

Shark, Basking, 1143.

Shark, Blue, 1130.

Shark, Fox, 1136.

Shark, Greenland, 1068.

Sharks, 1043, 1063, 1085, 1127.

Sharks, Cow, 1157.

Sharp-nosed Ray, 1117.

Shavianus (Cetorhinus), 1144.

Sheatfish, 693.

Shelly, 909.

Sheppy Argentine, 932.

Sherg, 1054.
sibi (Thynnus), 91.

Sibirian Huch, 830.
siculus (Ammodytes), 574.
signifer (Thymallus), 883.

Siil, 579.

Siil-Smelt, Greater, 913.

Siil-Smelt, Hebridal, 917.

Sik, 883, 891, 898, 900.

Sikloja, 893.

Sikinatk, 832.

sikus (Coregonus), 898.

Sikvimma, 893.

Sildelodde, 882.

Silfverfisk, 87.

Silfverlax, 856.

Sill, 954.

Sillack, 898.

Sillapipare, 1187, 1195.

Sillkung, 322.

Sillmor, 507.

Silurichthys, 693.

Siluridae, 66, 692.

Siluridse heteropterse, 692.

Siluriformes, 692.

Silurus, 691, 693.

Silus, 914.

silus (Acantholepis, Argentina, Coregonus),
Salmo), 913, 917.

Sivnia marina, 1079, 1084.

Simpa, 179.

Simpknot, 167.

Simpstubb, 260.

sinensis (Gasterosteus), 647.

sinuatus (Merlucius), 516.

Siphagonus, 203.

Siphonostoma, Siphostoma, 668.

Siren lacertina, 1204.

Sittichkarpf, 719.

Sjurygg* 294, 297.

Sjofjiiril, 212.

Sjohane, 306.

Sjokarp, 727.

Sjokock, 151, 272, 273.
Sjoruda, 736, 780.
Skaboskadda, 396.

Skalle, 780.

Skall-id, 763, 767.

Skall-jer, 763.

Skalmussla, 471.

Skarpsill, 974.

Skata, 1120.

Skate, Common, 1120.

Skate, Black-bellied, 1123.
Skate, Long-nosed, 1125.
Skate, Shagreen, 1115.
Skate, Sharp-nosed, 1117.
Skates, 1114, 1119.

Skates, White, 1114.
Skattbonde, 41.

Skavskadda, 396.
Skinnskrabba, 185.
Skinnalling, 660.

Skipjack, 988.

Skipper, 353.

Skitspigg, 657, 757.
Skjadbrosme, 543.
SkjEer-ising, 425.

Skolast, 590.

Skomakare, 752.

Skotta, 18.

Skottnat, 741.

Skrabb, 185.

Skrej, 87, 476.

Skrobba, 398.

Skrubba, 403.

Skrubbskadda, 398, 403.
Skuggfisk, 50.

Skyggfisk, 214.

Skalla, 398.

Skalling, 29, 30.

Skadda, 377.

Skaggsimpa, 208.
Skaggtorsk, 493.

Skalryta, 185.

Skar-auer, 155.

Skarflundra, 378.
Skargardssill, 959.

Skarknif, 823.

Skarlanga, femtommad, 554.
Skarlanga, fyrtommad, 544.
Skarlanga, nordisk, 548.
Skarlanga, tretommad, 550.
Skiirsn ultra, 18.

Skotspigg, 657.
Skotstromming, 965.

Slant, 1009.

Slemsugare, 291.

Slom, 867.

Slukal, 1033.

Slutenblasingar, 461.
Slathvarf, 441, 443.
Slathvarfvel, 443.

Slatrocka, 1120.

Slatta, 390.

Slatte, 391, 398.

Slattika, 391.

Slatting, 743, 746.
Slat-tobis, 573.

Slatt-vahr, 392.

Slattvar, 443.

Smear Dab, 383.

Smed, 561.

1238

Smelt, 863, 869.

Smelt, Arctic, 867.

Smelt, Atlantic, 867, 869.

Smelt, Pacific, 867.
smiridus (Merluccius), 515.

Smahvarf, 453.

Smalax, 832.

Smaling, 898.

Smaroding, 842.

Smasej, 503.

Smaspigg, 658.

Smatorsk, 476.

Smarling, 898.

Smorbult, svart, 245.

Snsebel. 909.

Snake-fish, 711.

Snorfojbul, 297.

Snorgers, 41.

Snorluf, 41.

Snorpels, 41.

Snultra, 2.

Snurrevad, 1035.

Snorpevad, 972.
socialis (Salmo), 876.

Sockerskadda, 372, 382.

Sole, 386.

Sole, Smooth, 428.

Solea, 371, 372.

solea (Pleuronectes), 372.

soleeeformis (Arnoglossus, Rhombus), 429.

Soleina, 371.

Soleini, 370.

Solenostoma, 662, 663.

Soles, Double, 375.

Soles, Reversed, 375.

Solsensodg, 757.

Somniosus, 1167.

Sorthaa, 1163.

Speer, 511.

Spallauzani (Leptocephalus), 1038.

Spanjor, 1119.

Sparidse, 25, 52.

Sparini, 53.

Sparlingus, 974.

Spams, 56.
speciosa (Julis), 21.
speciosus (Labrus), 10.
specularis (Cyprinus), 724.

Sperlin-Bleak, 797.
spet (Sphyraena), 327.

Spetsstjertadt Langebarn, 225.

Sphagebranchus, 1022.

Sphyraena, 917, 945.
sphyrasna (Argentina), 912, 917.

Sphyrsenidee, 327.

Spliyrnae, 1128.

Spigg, 2, 657.

Spillanga, 530.

Spilsteenghyse, 1084.
spilurus (Aspius), 782.

Spinachia, Spinochia, 637, 638.

spinachia (Gasterosteus, Gastraea), 638, 660.

Spinacidse, 1128, 1157.

Spinax, 1162.

spinax (Acanthias, Etmopterus, Squalus), 1157,
1163.

Spindelflynder, 379.
spinosus (Cyclopterus), 298.
spinosus (Echinorhinus), 1157.
spinosus (Zeus), 306.

spinosissimus (Aspidophorus, Leptagonus), 206.
spinulosus (Gasterosteus aculeatus), 646, 648.
Spirinchus, 869.

spirinchus (Osmerus, Salmo), 867, 870, 875.
Spirlin, 798.

Spirlinus, 790, 796.
splendens (Bervx), 68, 429.

Spjutrocka, 1098.

Spotted Dog Fish, 1154.

Spotted Dog Fish, lesser, 1154.

Spotted Goby, 262.

Sprat, 953, 974.

Spratella, 974.

Spratelloides, 952.

Sprattus, 974.

sprattus (Clupea, Harengula, Meletta), 952,
974, 979.

Springkrok, 1009.
spurius (Salmo), 851.

Spatta, 398.

Squalius, 760, 770.

Squalo, 770.

Squalus, 1157.
squalus (Leuciscus), 778.

Squamipinnes, 49, 52, 71.
squatic-ola (Alepas, Anelasma), 1166.
squatina (Rhina), 975.

Stafsill, 989.

Stagg, 657.

Staggsill, 87.

stagnalis (Salmo), 835, 843, 867.

Stainbuck, 297.

Stainbajtare, 297.

Stakanat, 859.

Staksill, 87, 984.

Stamp, 530.

Stamsill, 983.

Stavsild, 919.

Steensuer, 1208.

Steinbits-Broder, 236.

Steinbutt, 441.

stellaris (Scylliorhinus, Scyllium, Squalus), 1076,
1148, 1152.

stellatus (Acipenser), 1055.

stellatus (Liparis), 290.

stellatus (Platessa, Pleuronectes), 399, 403.

Stenbider, 235.

Stenbit, 297, 711, 831—833.

Stenbrosme, 543.

Stenflundra, 441.

Stengers, 174.

Steulake, 172, 711.
stenobrachii, Acanthopterygii, 24
Stenodus, 875, 889.
stenorhynchi (Coregoni lavareti), 902.
stenoripidi, Acanthopterygii, 24.

Stensimpa, 170.

Stensnultra, 16.

Stenspotta, 441.

Stensugare, 172, 385, 746, 1195.

Stensut, 172.

Sterility of Salmons, 838.

Sterlet, 1049, 1055.

Sternoptychidae, 305, 826, 923, 947.
Sternoptychinae, 923, 924.

Sternoptyx, 305, 925.

Stickleback, Fifteen-spined, 638.

Stickleback, Ten-spined, 658.

Stickleback, Three-spined, 647.

Stickleback-net, 657.

Sticklebacks, 2, 635.

Stigmatophora, 687.

Stincus, 869.

Stirr, 833.

Stizostedium, 25, 36.

Stockfisk, 503.

Stoer, 1056.

Stolephorus, 990.
stomachicus (Salmo), 840, 851.

Stone-Bass, 47.

Stone-biter, 711.

Stor Flygfisk, 357.

Storgap, 425.

Storguoding, 200.

Stormun, 425.

Storuors, 868.

Storryssja, 859.

Storroding, 842.

Storsik, 900.

Storslom, 868.

Storspigg, 647.

Stortorsk, 476.

Strandsatt, 859.

Strandvad, 1035.

Strinsia, 463.

Striped Wrasse, 10.

Stroeinii (Carelophus, Gunnellus), 218.
Stroemii (Macrurus), 590.

Strafling, 763.

Stromharr, 887.

Stromming, 959, 960.

Stromming, Autumn, 960, 966.

Stromming, Ice, 965.

Stromming, Net, 965.

Stromming, Spring, 960, 965.
Strommingsvarp, 970.

Stromsill, 917, 919.

Sturgeon, 1056.

Sturgeon-fishes, 1043.

Sturgeons, 829, 1054.

Sturio, 1056.

sturio (Acipenser), 1046, 1056, 1204.
sturioides (Acipenser), 1057, 1061.

Stuvitzii (Boieogobius, Gobiosoma, Gobius
Latrunculus), 266.

Stylophorus, 314.

Stymphalicus (Leuciscus), 787.

Styria, 1056.

Stangkrok, 1008.

Staffing, 763.

Stam, 720, 760.

Stor, 1056.

Sucker, Bimaculated, 302.

Sucker, Lump, 294.

Suder, 752.

Suderna], 14.

Sudis, 827, 945.

Sugare, 283.

Sugfisk, 283.

Suidteri (Coregonus), 899.
suillus (Ctenolabrus, Labrus), 16.

Suiefianus (Blennius), 213.

Sula, 375.

Sulzeri (Coregonus), 899.
sundaica (Clupea), 952.

Sunfish, Short, 625.
superciliosus (Clinus), 219.
suppositus (Gasterosteus), 648.
surmuletus (Mullus), 63.

Sutare, 720, 748, 752.
svalliza (Squalius), 770.

Svartbuksrocka, 1123.

Svart Smorbult, 245.

Svartspoling, 815.

Svarttorsk, 477.

Svardfisk, 222.

Swordfish, 118, 2 2 1 .

Synaphobranchidae, 1022.

Synentognathi, 342.

Syngnathi marsupiales, 667.

Syngnatlii ophidii, 667.

Syngnathidae, 662, 663.

Syngnathinas, 663, 696.

Syngnathus, 347, 661, 667.

Sypiga, 14.

1239

syrtensium (Argentina), 914.
Sal a, 375, 425.

Salbling, 881.

So-abborre, 10.

Soe-Naal, 680.

Solvfisk, 1084.

Solvhaae, 1079.

Solvqveite, 314, 315.

Somus, 1084.

Sorotte, 1084.

Soriiv, 1084.

Taeand, 493.

Tadpole Fish, 558.

Taenia, 314.

taenia (Acanthopsis, Botia, Cobitis, Gobitis),
703, 705, 706, 714.
tgeniensis (Clupea harengus), 959.
Taeniiformes, 309.

Tgenioides, 309.
tasnionotus (Syngnathus), 669.
taeniopterus (Cottns), 181.
taenuis (Arnoglossus), 428.

Taerbe, 1107.

Taggmakril, 86.

Taggsill, 87.

Taggsimpa, 191.

Tajraeu, 832, 837, 856.
tancoides (Labrus), 7.

Tandfierasfer, 601.

Tangbrosme, 223.

tapeinosoma (Arnoglossus), 428, 430, 431.
tapinorhynchi (Coregoni lavareti), 900.

Tara, 482.

Taretorsk, 477.

tan (Batrachus, Gadus), 134.

tazo (Scomber), 108.

Tectobranchii, 1063.

Tectospondyli, 1128.

Tejstefisk, 222.

Telara, 990.

telara (Setipinna, Stolephorus), 991.

Teleostei, 1.

Teleostei physostomi, 689.

Teleostomi, 1044.

Telescope-fish, 734.
telescopus (Cyprinus), 733.

Telestes, 753.

Temnodon, 82, 85.

Tench, 720, 748.

Tennfisk, Tennfisk med spjut, 657.
tenuirostris (Anguilla), 1023.
tenuirostris (Syngnathus), 671.
tenuis (Encheliophis), 601.

Tepel, 1120.

Tephritis, 408.
tessellatus (Labrus), 11.

Teste, 222.

Testefisk, 220.

tetracanthus (Gasterosteus), 644, 645, 648.
Tetragonuridse, 327.

Tetragonurus, 327.

Tetrapturus, 117.

Tetrodon, 619, 664.

Tetrodontina, 622.
texana (Anguilla), 1023.
texanus (Gasterosteus), 648.

Thalassophryne, 133.

Thaleichthys, 875.

thazard (Auxis, Scomber), 89, 92, 108.
Tholichthys, 24.

Thompsonii (Acipenser), 1057.

Thomsonii (Cottus), 157.

Thon blanc, 1138.

thoracatus (Cyprinus), 733.
thoracici, Acanthopterygii, 24.
thoracopteri, Orthocormi, 24.

Thornback, 1074, 1087, 1104.

Thresher Shark, 1137.

Thrissa, 952, 984, 990.

Thrissomorphi, 690, 826.

thunina, thunnina (Euthynnus, Orcynopsis,
Scomber, Thynnus), 93.

Thyinallus, 828, 882, 884, 891, 892.
thymallus (Coregonus, Salmo), 884.

Thymus, 884.

thynnus (Orcynus, Scomber, Thynnus), 91, 97,

101.

Thyris, 363.

tiberinus (Leuciscus), 770.

Tilesia, 482.

Tinfish, Speared, 657.

Tinea, 720, 722, 747, 748.
tinea (Cyprinus, Leuciscus, Tinea), 748.
tinea (Labrus), 7, 18.
tinea (Strinsia), 463.

Tigerflaekad hafkatt, 2.36.

Tistefisk, 222.

Toadfish, Common, 135.

Toadfish, European, 134.

tobianus (Ammodytes), 568, 570, 574.

Tobiskung, 570.

Tok, 1059, 1060.

Tope, 1072, 1132.

Topknot, Ekstrom’s, 453.

Topknot, Muller’s, 458.

Torpedinidfe, 1093.

Torpedo, 1071, 1086.

Torr-borr, 1107.

Torsk, 2, 364, 368, 472, 562.
torvus (Caranx), 85.
torvus (Cottunculus), 157.

Townsendii (Chimsera, Ischyodon), 1078.
Trachinidic, 126, 127.

Trachinini, 127.

Trachinomorphi, 126, 127.

Trachinus, 128, 272.

Trachipterus, 314.

Trachurus, 84.

trachurus (Caranx, Scomber, Trachurus), 83,
84, 86, 975.

trachurus (Gasterosteus aculeatus), 645, 648.
trachurus (Hemilepidotus), 156.

Trachydermus, 162.

Trachynotus, 83, 85.

Trachypterida?, 126, 309.

Trachypteromorphi, 126, 309.

Trachypterus, 304, 310, 312, 314.
Trammel-net, 741, 1007.

Triacanthinge, 622, 631.

Triacanthus, 622.

Triadontina, 622.

Trichis, Trichius, 952.

Trichiuridic, 309.

Trichonotidge, 211.

tricirratus (Gadus, Motella, Guos), 544, 550,
551, 556.

tricuspis (Cottus, Phobetor), 160.
tridactylus (Salarias), 212.
trifurcus (trifurcatus, Blennius, Raniceps), 559.
Trigger-fishes, 632.

Trigla, 146, 194.

Triglinte, 193.

Triglops, 157, 166.
trimaculatus (Labrus), 1 0.
tripunctulatus (Maurolicus), 931.

Trout, 851, 853, 855.

Trout, Common, 855, 856.

Trout, River, 856.

Trout, Sea, 860.

Trubbstjertadt Langebarn, 228.

Trumfiskar, 49.

Trumpet-fishes, 637.

Trutta, 841, 849, 884.

trutta (Salmo, Trutta), 830, 831, 833, 851.

truttula (Salmo), 851.

Trygon, 1092, 1097.

Trygonidae, 1096.

Trygonopterte, 1097.

Tryte, 28.

tschawytscha (Oncorhynchus), 830.

Tschir, 899, 904.

Tubfish, 200.

tudes (Zyggena), 1075.

tumidus (Chironectes, Lophius), 145, 146.

Tumula, 331.

Tunfisk, 97.

Tunga, 372, 385, 386.

Tungehvarf, radflackig, 428.

Tungens hoer-unge, 425.

Tunglik Fluudra, 391.
tunicatus (Liparis), 287.

Tun -bellies, 622.

Tunnina, 93.

Tunny, 90, 97.

Turbot, 368, 432, 434, 435.
turdus (Labrus), 18.

Turneri (Lycodes), 609, 612.

Tushaj, 1155.

Tusk, 562.

Tvarspol, 832.

Twaite Shad, 984.

Typhle, Typhline, 667.

typhle (Siphonostoma, Siphostoma, Syngnathus),
663, 665, 666, 669, 672, 674.
typicus (Rhinodon), 1073.
typus (Carangichthys), 83.

Tanglake, 603, 711.

Tangsnipa, 683.

Tangsnalla, liten, 672.

Tangsnalla, stor, 668.

Tangsnarta, 218.

Uakal, 481.

Uaegte Tunge, 425.

Ulk, 185, 190.

Umbla, 841.

umbla (Salmo), 830, 831, 841, 842, 867.
Umble, TJmbre, 841.
umbra (Sciasna), 51.

Umbridae, 997.

uncinatus (Centridermichthys, Cottus, Icelus),
163.

undulata (Raja), 1120.
unicolor (Ammocoetes), 1183, 1186.
unicolor (Orcynopsis, Pelamichthys, Pelamys,
Scomber), 91, 103.
unicolor (Sparus), 53.

unimaculatus (Rhombus, Scophthalmus), 452.
unipuuctatus (Gobius), 262.

Upengeus, 61.

Ur, 152.

Uraleptus, 539.

Uranidea, 157, 169, 170.

Uranoscopus, 271, 272.
uranoscopus (Gobio), 742.

Uraspis, 83.
uraspis (Caranx), 83.

Uria grylle, 222.

Urocentrus, 67.

Urochorda, 1210.

Urogymni, 1097.

Urokadlin, 481.

156

Scandinavian Fishes.

1240

Urolophi, 1097.

Ursini (Ozodura), 626.
Uakan, 481.

Vaagnner, 315.

Vad, 971.

Valilii (Lycodes), 609, 610, 613.

Val, 889.'

Vampire-Ray, 1094.
vampyrus (Ceratoptera), 1094.

Vandesius, 894.
vandesius (Coregonus), 894.

Vanlig Fjarsing, 128.

Vanlig Hafkatt, 232.
variabilis (Acanthocottus, Cottus), 180.
variabilis (Labrus), 7.
variabilis (Trutta), 851.
variegatus (Labrus), 10.

Vaslodde, 882.

Vegse (Pleuronectes), 390.

Veling, 763.

velivolans (Lepidorhombus, Zeugopterus), 447
451.

Vendace, 893.

venernensis (Salmo), 851.

ventralis (Cottus, Gymnocanthus,Phobetor), 160

ventricosus (Salmo), 842.

Vermlandsroding, 842.
vernalis (Merlangus), 511.

Veron, 754.

Verrillii (Lycodes), 617.
vespertilio (Lophius), 136.
vetula (Abramis), 812.
vexillifer (Thyinallus), 884.

Vieille, La, 7.

Vildkraks-simpa, 185.
villosus (Clupea, Mallotus, Salmo), 876.
Vimba, Vimma, 720, 799.
vimba (Abramis, Leucabramis, Cypriuus), 798
799.

vimba (Coregonus, Salmo), 893, 896.
viola (Antimora, Haloporphyrus), 581.
vipera (Trachinus), 131, 266.
virens (Gadus, Pollachius), 466, 467, 500
viridescens (Osmerus), 869, 872.
viridi-violaceus (Cyprinus), 724.

Visse-al, 1032.

vitrea (Perea, Stizostedium), 37.
vittatus (Cyprinus), 724.
vittatus (Labrus), 10.

viviparus (Blennius, Encbelyopus, Zoarcasus,
Zoarces), 603, 604.
viviparus (Sebastes), 148, 149.

Vogmar, 314, 315.
vogmarus (Trachypterus), 315.
volgensis (Perea), 37.
volitans (Exoccetus), 357, 358.
vomer (Raja), 1125.
vomer (Selene), 83.
vomerinus (Lopbius), 144.
vorax (Aspius), 782.

Vrakfisk, 47.

Vrakbus, 859.

vulgaris (Acanthias), 1158.

vulgaris (Acerina), 41.

vulgaris (Alausa, Alosa), 984, 985.

vulgaris (Anguilla), 1011- — 1021, 1023.

vulgaris (Auxis), 108.

vulgaris (Batis), 1121.

vulgaris (Belone), 347.

vulgaris (Brosmius, Brosmus), 562.

vulgaris (Cantbarus), 54.

vulgaris (Carassius), 735.

vulgaris (Conger), 143, 975, 1038.

| vulgaris (Engraulis), 992.
vulgaris (Eperlanus), 870.
vulgaris (Flesus), 399.
vulgaris (Galeus), 1133.
vulgaris (Gobio), 743.
vulgaris (Gunnellus), 220.
vulgaris (Hippoglossus), 408, 409.
vulgaris (Julis), 21.
vulgaris (Leuciscus), 761.
vulgaris (Limanda), 387.
vulgaris (Liparis), 284, 287.
vulgaris (Lota, Molva), 526, 532.
vulgaris (Merlangus), 487.
vulgaris (Merluccius), 515.
vulgaris (Motella, Onos), 550.
vulgaris (Mustelus), 1128.
vulgaris (Perea), 27.
vulgaris (Platessa), 392.
vulgaris (Solea), 365, 371, 372, 373.
vulgaris (Spbyrama), 327.
vulgaris (Spinachia), 638.
vulgaris (Thyinallus), 884.
vulgaris (Thynnus), 98.
vulgaris (Tinea), 720, 748.
vulgaris (Trygon), 1099.
vulpecula (Thalassorhinus), 1075.
vulpes (Alopecias, Alopias, Carcbarbinus, C;t r-
charias, Squalus), 1075, 1135, 1136.
Valgild Torsk, 476.

Wabasliensis (Anguilla), 1023.

Waclina, Wacbnja, 481.

Wagneri (Petromj'zon), 1187.

Wartmanni (Coregonus, Salmo), 892, 898, 900,
905, 907.

Weever, Great, 128.

Weever, Lesser, 131.

Whiff, 447, 448.

whiff (Lepidorhombus, Pleuronecfes), 427, 447,
448, 455.

Whitebait, 954.

White Bream, 803.

Whitefish, 759, 909.

Whiting, 465, 487.

Whiting, Pollack, Norwegian, 508.

Whiting pout, 493.

Widegreni (Coregonus), 898.

Willuglibii (Coregonus), 894.

Willughbii (Trematopsis), 626.
Winter-Herring, 959.

Wobbegong, 1147.

Wolf, 46.

Wolf-fish, 232.

Wrasse, Ballau, 7.

Wrasse, Comber, 7.

Wrasse, Rainbow, 21.
Wrasse, Scale-rayed, 5.
Wrasse, Small-mouthed, 14.
Wrasse, Striped, 10.
Wrasses, Parrot, 2.
Wreck-fish, 47.
wjmaadensis (Silurus), 697.

Xiphias, 117, 118.
Xiphiidse, 70, 116.

yarkandensis (Cobitis), 705.
Yarrelli (Argentina), 918.

Zachirus (Glyptocephalus), 386.
zebra (Brachirus), 371.
Zenarchopterus, 343.

Zenidae, 70, 126.

Zenopsis, 305.

Zeugopterus, 426, 456.

Zeus, 304.

Ziege, 823.

Zoarcfeus, Zoarces, 211, 602.
Zope, 720, 721, 819.
zunasi (Clupea), 978.

Zygfena, 1071.

Zarthe, 720, 799.

Zarthe, Black, 801.

Zarthe, Pale, 801.

ibuk, 772.

Adrag, 763.

Albrosme, 609.

Alglip, 1035.

Algard, 1035.

Alhomma, 1035.

Alkista, 1034.

Alkupa, 1034.

Alkussa, 607.

Allubb, 609.

Alryssja, 1035.

Alvad, 1035.

Alvimma, Alvinna, 1032.
Angerpiga, 390.
Anims-vimma, 894.
Arannare, 772.

Angsgadda, 1005.

Oienpaal, 512.

oresundica (Clupea harengus), 959.
Ore val, 889.

Oring, 832, 833.

Ornrocka, 1095.

Ort, 768.

Errata.

Page 594, column 2, line 12 from above: commensual, read commensal
Page 623, column 2, line 23 from above: aponeuros, read aponeui'osis

Page 1031, column 1, line 23 from above: 6 — 8 mm. (quoted from Jacoby) should probably be 6 — 8 cm., so note c oij the
following page should be omitted.

SMITT. SCANDINAVIAN FISHES

A HISTORY OF

SCANDINAVIAN FISHES

BY

B. FRIES, C. C. EKSTROM, AND C. SUNDEVALL

WITH COLOURED PLATES

BY

W. von WRIGHT

AND TEXT ILLUSTRATIONS

SECOND EDITION

REVISED AND COMPLETED BY

Professor F. A. SMITT

MEMBER OP THE ROYAL SWEDISH ACADEMY OF SCIENCE

TEXT

PART II

P. A. NORSTEDT & SONER
STOCKHOLM

BERLIN LONDON PARIS

R. FRIEDLANDER & SOHN SAMPSON LOW, MARSTON & COMPANY, Limited LIBRAIRIE C. REINWALD & CIE

Carlstrasse ii St. Dunstan’s House, Fetter Lane is, rue des Saints-P^res

PROSPECTUS.

“The illustrated work now placed in the hands
of the public is designed not only to supply scientific
men with accurate and trustworthy figures of the
piscine species of Scandinavia, and to throw greater
light on this section of the Scandinavian fauna; it
has a further object, no less important — to assist
the general reader in recognizing the different species,
and to render the study of them practicable to all
who desire to join to the pleasure and profit of the
fisherman the zoologist’s knowledge of the fishes and
of their life and habits.” Such was the plan of Scan-
dinavian Fishes, painted from living specimens
and engraved on stone by W1LH. von WRIGHT,
and described by B. FR. FRIES and C. U. EK-
STROM, at the first appearance of this work in 1836.
Three friends, all with the same object at heart, and
each among the foremost in his department of know-
ledge, had joined forces to render scientific ichthyology
a popular study. The work was interrupted in 1839
by the death of one of their number; but C. J. SUN-
DEVALL stepped into the place left by Fries, the
acute man of science again stood shoulder to shoulder
with the practical zoologist, and the artist continued
to wield his brush with unsurpassed skill, until his
hand was enfeebled by sickness, and the work was
abandoned in 1857. In its incomplete form, however,
it gained a world-wide reputation.

Every age has its characteristic peculiarities.
All the efforts of modern natural science are bent on
unravelling the problem of evolution. The claims on
a zoological description at the present day are not
what they were fifty-seven or even thirty-six years
ago; but with comparatively slight alterations and
additions the text of ‘Scandinavian Fishes’ will still
rank as a pattern of excellence. As several paintings
by v. Wright, hitherto unpublished, were besides
preserved in the archives of the Royal Swedish Aca-
demy of Science, the publishers determined upon
issuing a new edition of the work, and with this aim
consulted the present occupier of Sundevall’s post
at the Royal Zoological Museum. With the type-
specimens of the former descriptions at his disposal,
and with the other rich collections of fishes possessed
by the Museum to draw upon, Professor F. A. Smitt
has endeavoured to revise the text so as to satisfy

the requirements of modern science, and in accord-
ance with the opinions which he has maintained in
his previous zoological writings. According to the
opinions pronounced in several quarters by competent
judges his researches have elicited so much new
and valuable information that the revised edition
may with full justice be regarded as an entirely
new work. The plates are composed principally of
v. Wright’s paintings, reproduced in colours; but
where coloured figures, drawn from living or perfectly
fresh specimens, were to be found among the collec-
tions of the Royal Museum, or where such figures
could be procured from other sources, these are also
given, printed in colours and executed with all the
accuracy and finish attainable in this country, the
lithography and printing being the work of the Litho-
graphic Press of the Royal Swedish Ordnance Survey.
Where such drawings could not be procured, or
where the species in point is not of essential im-
portance either in the Scandinavian fauna, or from an
economic point of view, zincotypes, cast at the same
establishment, are inserted in the text, from drawings,
executed with all possible accuracy, of specimens
preserved in spirits at the Royal Museum, or of
examples borrowed from other collections. Most of
these figures have been drawn by Carl Erdmann,
an artist whose early loss to science and art must
be greatly deplored.

The piscine fauna of Scandinavia includes 224
species. In the former edition of this work 64 species
were figured; in the present edition reproductions are
given of 223 species. Of one species it has proved
impossible to obtain any figure, no example of the
said fish being now to be found in any museum. The
new edition besides contains representations of 5
Cyprinoid hybrids and 9 Arctic species. The total
number of coloured figures is 190, of zincotypes in
the text 380.

The situation of the Scandinavian Peninsula is
such that nearly all the piscine species of Central
and Northern Europe occur in its inland waters or
off its coasts. As almost all these species find a
place in the new edition, the work is consequently
a handbook of importance for our part of the
globe.

o o

o

The work, containing 1239 pages of text and 55 plates, is published, in paper covers complete, at a
price of 12 Guineas — 224 Mark — 280 Francs.

Separate parts not supplied.

4-5 25 '