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Full text of "monograph of the British marine annelids"

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THE 



RAY SOCIETY 



INSTITUTED MDCCCXLIV. 




This Volume is issued to the Subscribers to the Ray Society for the Year 1907. 



LONDON: 



MDCCCCVIII. 



PRINTED BY ADLARD AND SON, LONDON AND DORKING. 



H\5i 
/.Pi* I 



MONOGEAPH 

/ 



OF THE 



BRITISH ANNELID^. 



YOL. II.-PAET I. 

POLYCH^TA. 
NEPHTHYDID^l TO SYLLID^. 

Pages 1 — 282 ; Plates XLIII — L, Coloured, and LVII — LXX, Uncoloueed. 



by 

WILLIAM CAHMICHAEL McINTOSH, M.D.Edin., 

L.E.C.S.E., LL.D., F.E.S., F.E.S.E., F.L.S., C.M.Z.S., V.P. EAY SOC., V.P. LIT. & PHIL. SOC. St. AND., V.D., etc., 

PROFESSOR OF NATURAL HISTORY IN THE UNIVERSITY OF ST. ANDREWS ; DIRECTOR OF THE UNIVERSITY 

MUSEUM AND OF THE GATTY MARINE LABORATORY. 



LONDON : 
PRINTED EOR THE RAY SOCIETY. 



1908. 



DEDICATED 

TO 

THE BT. HON. LORD EEAY, LL.D.(St. Andrews), 

G.C.S.I., G.C.I.B., ETC., 

RECTOE OF THE UNIVERSITY 1884-86, 

WHOSE LONG-CONTINUED INTEREST IN THE MARINE WORK AT ST. ANDREWS 
HAS BEEN A SOURCE OF SINCERE SATISFACTION. 



TEMPOBABY PREFACE 



The present Part contains a larger number of species than Part II of the previous 
volume, and the majority are more or less brilliantly coloured, not a few, for example, in 
the PhyllodocidaB, Hesionidge, and Syllidse being amongst the most beautifully ornamented 
Invertebrates; indeed, many vie with the gaudy tints of butterflies and birds or the 
burnished splendour of beetles. Fortunately some of the finest of them had been 
drawn from life by the late Mrs. Gimther (Roberta Mcintosh), and their softness and 
beauty did not suffer at her hands. 

It is perhaps well to note that the author does not feel warranted in adopting any of 
the recent classifications of the Polychaata, e. g. that of Professor Benham ( £ Cambridge 
Natural History,' ii, 1896), which has certain features of Levinsen's classification, because 
none relieves the difficulties encountered in the older and more simple classification into 
errant and sedentary forms by Audouin and Edwards. Thus in such a system as 
Benham's, incongruous families are grouped together (e. g. the Nephthydidse between the 
Nereidse and Amphinomidse), and the arrangement has the support neither of structure 
nor development. The Polychseta, indeed, do not lend themselves readily to the 
systematist, and it is safer at present to place the families in series according to their 
natural and structural relationships, reserving further consideration of the subject for 
the summary. There are notable differences in the structure of the body-wall, the 
arrangement of the great nerve-cords and the neural canals, the digestive system and 
other features, but even these do not readily conduce to systematic arrangement. They 
are thus in contrast with the Nemerteaus, the classification of which in 1868-9 on a struc- 
tural basis has never been disputed, though modern nomenclature may sometimes obscure 
the original facts. 

Dr. Goodrich has kindly drawn the segmental organs typical of the Nephthydidse, 
and his experience of these is great. 

I have to thank Dr. Allen, the Director of the Plymouth Marine Laboratory, who 
exerted himself to procure such of the southern types as were not readily obtainable in 
the fresh condition elsewhere. 

To Miss A. H. "Walker, the artist who has faithfully aided me throughout the work 
on the Annelids, special acknowledgments are due. Her pencil and brush have been of 
great service. 

I have to acknowledge the courtesy of the Carnegie Trust in having given aid for 
three years in regard to the artist and in section-making. Without such help it would 
have been impossible to overtake the work in time for issue in 1907. Moreover, the 



viii TEMPORARY PREFACE. 

Carnegie Trust has granted a sum of £100 for 1907-8, and this, together with a balance, 
will be utilized for the heavy expenses of the coloured plates. By this liberality the 
Ray Society will be the more able to take in hand the next part (similar in size and in 
the number of plates), which lies ready for printer and lithographer. 

To Mr. Kappel, Librarian of the Linnean Society, and Mr. White, Librarian of the 
Royal Society, my best thanks are due for their valued aid with books. 

Lastly, I have to thank Mr. Hopkinson, the Secretary of the Ray Society, for the 
great pains he has taken in the arrangement of the figures in the coloured plates, the 
verification of certain references, for general interest and advice in the course of the 
work, and especially for his careful revision of the proofs. 

W. C. M. 

Gatty Marine Laboratory, 
St. Andrews; 
October 10th, 1907. 



FAMILIES, GENERA, AND SPECIES, OE THE BRITISH MARINE 

ANNELIDS. 

(ANNELIDA POLYCELETA, continued.) 



Family VI. — Nephthydid^e, 1 Grube, 1851. 

Head flattened, quadrangular or rhomboidal, rarely nearly hexagonal, a short tentacle 
at each anterior angle, and one (palpus, Kinberg, tentacular cirrus, Ehlers) a little 
posterior (1st segment) and ventro -lateral in position, or both may be absent. Eyes two 
or none. A small ciliated organ at each lateral groove posteriorly. 

Body elongate, almost tetragonal in section, though convex dorsally and flattened 
ventrally. Segments similar. Vent terminal with a single cirrus (rarely two, Ehlers) 
beneath. Peristomial segment fused with that behind. Proboscis large, sub-cylindrical, 

dv. 




Vrn. 



Fig. 34— Section of NepMhys caeca, about the anterior third. Id, dorsal division of the foot ; bv, ventral division ; 
d, alimentary canal ; dm, dorsal longitudinal muscle ; dv, dorsal blood-vessel ; nc, nerve-cord ; om, oblique 
muscle ; vm, ventral longitudinal muscle. 

ovate or obovate with or without a double arch of bifid papilla, a pair of horny teeth or 
none. Alimentary canal simple — slightly moniliform. 

Divisions of the foot widely separated, flattened, bearing a spine and double row of 
diverse bristles (barred and serrated or rarely with lyriform tips) in each in front, the 
dorsal lobe, further, having superiorly a lamella and other processes, and a small dorsal 
cirrus in connection with the branchia; the ventral lobe has a lamella with or without 
other processes and a ventral cirrus. 

1 The title Nephthya was used by Savigny for a group of Alcyonaria including Spongodes. 

58 



2 NEPHTHYDnLE. 

A dorsal and a ventral blood-vessel (with a cardiac dilatation in front), and two 
lateral trunks. 

Body-wall (Fig. 84) in section powerfully muscular, the dorsal longitudinal muscles 
often fusing in the middle line and having an accessory band at the shelving edge. 

Nerve-area (see Fig. 37, p. 6) shaped like an inverted crown. Four conspicuous 
neural canals — the inferior the larger, occur on each side of the median line (V. casca), 
besides a variable number of others. To the exterior lie the tough limiting membrane, 
the hypo derm, and the cuticle. 



ISO. 




E:S&. 



rvp. 

Fig. 35. — Magnified drawing of the segmental organ of Nephthys by Dr. Goodrich, co, ciliated organ ; iso, internal 
end of the nephridium with solenocytes ; npc, nephridial canal ; np, nephridispore. 

The structure of the body- wall in Nephthys Hombergii as shown by Emery * does not 
differ essentially from the foregoing. 

Segmental organs (Fig. 35) are long, slender tubes opening ventrally near the base 
of the foot ; end blindly internally in a tuft of three to five branches with rows of soleno- 
cytes (Fig. 36). An incomplete genital funnel in some passing into a sac in which are 
waste products (Goodrich). F. H. Stewart 3 pointed out at St. Andrews the exact 



1 f Mitth. Zool. Stat. JSFeapel/ VII, p. 371, pi. 13. 

2 ' Ann. Nat. Hist./ ser. 7, vol. v, p. 161, pis. ii and iii, 1900. 



NEPHTHYDKLE. 3 

position of the various parts of the organ and increased the information concerning the 
excretion of solid matters and other physiological features. The mode of exit of the 
genital products is unknown. 1 

Goodrich has again examined the structure of the organ, and is of opinion that with 
regard to the products of excretion they lie in a kind of sac formed of peritoneal 
epithelium. Louis Fage 2 does not adopt F. Stewart's view that the amoebocytes from the 
phagocyte organ enter the nephridial tube, but is inclined to think that they pass into 
it by simple osmosis. Fage concludes by mentioning that the ciliated organ, shaped 
like the shell of a Pecten, is present, and is largest in the middle and posterior regions, but 



,x x JL 



HI 




n.cajjy. 



71ZIC. 



"E.S.G, 



Fig. 36— Highly magnified representation of the solenocytes and segmental (nephridial) canal by Dr. Goodrich. 
/, cilia ; nuc, nuclei ; pp, protoplasmic process of a solenocyte ; sol, solenocytes ; t, tubes connecting solenocytes 
with the nephridial canal. 

has no internal opening, so that it is, he says, impossible for the genital products to find 
their way outward by this means. He thinks that the ciliated organ only conducts 
the solid particles of the coelom to the phagocyte organs. 

The eggs appear to be demersal, but the larvae are pelagic. 

As a family the Nephthydidae are characterised by great uniformity in the structure 
of the bristles, so that it would not be easy to distinguish the species by this means, and 
thus they differ from the majority of the Annelids. The characters of the feet, however, 
are distinct, and whilst the sexual variations may not be fully known, their structure may 
be considered reliable in discriminating the various forms. 

Few species possess eyes, yet several with such, e.g. NepMhys Verrilli, were described 
in the collection of the ' Challenger,' 3 and this was a form allied to the K picta of Ehlers 
in the structure of the feet. 

1 I am indebted to Dr. E. S. Goodrich for these figures (35 and 36). 

2 'Ann. Sc. Nat/ 9 e ser., p. 286. Figs. 11-16, pi. vi, figs. 5 and 6. He chiefly investigated 
NepMhys Hombergii. 

3 Yol. xii, p. 163, pi. xxvi, f. 6, 7, pi. xxxia, fig. 8. 



4 NEPHTHYDIDJS. 

No British species shows the large foliaceous branchiae occasionally seen in foreign 
species, as in the N. phyllobranchia of the ' Challenger.' 

Though not falling within the British area it may be mentioned that Nephthys 
(Aglaophamus) inermis, Ehlers, occurs in European waters, having been procured in the 
' Porcupine ' Expedition of 1870 off Cape Finisterre in 81 fathoms. 1 It may occur yet 
nearer Britain. Ehlers procured it from Florida. 

Nephthys caeca and others are found at all parts between tide marks in North Uist 
in muddy sand, and therefore Dr. Drummond's opinion that they are only found at very 
low- water requires qualification. Moreover, since in this island very heavy showers of 
rain fall after they are uncovered by the tide for a considerable time, they can scarcely 
be so sensitive to fresh water as supposed. 

The various members of the family are used as bait. 

Certain features, and especially the structure of the bristles, suggest affinity between 
the Nephthydidae and the Ariciidae, and future investigation may throw light on this 
subject. 

Linnaeus included the marine Annelids under the great group of the Vermes, and the 
Nephthydidae were generally ranged amongst the Nereids, one of the groups of the 
Vermes Mollusca. A similar arrangement was followed by Otho Fabricius, and by 
0. F. Muller, who placed the majority under his Vermes Helminthica. 

Savigny in his ' Systeme' (1820) arranged the Nephthydians under his family Nereides, 
but he observed their distinction from the other members of the group, and hence he 
separated them as a sub-section — Nephthys. They are characterised by a proboscis 
furnished with tentacles at the orifice; antennas — both exterior and middle — equal; no 
tentacular cirri ; all the cirri short — almost absent ; branchiae distinct. 

Of the genus Nephthys he gave a detailed and fairly accurate description so far as he 
went, though he observed only a single row of bristles. 

Notwithstanding these remarks he on a later page was inclined to place the Nereis 
Gseca of Otho Fabricius under a new genus — Aonis. As the form referred to is only 
N cdsca, it is unnecessary to refer further to this lapsus. 

Audouin and Milne Edwards (1834) placed this family along with Glycera and 
Goniada in the second tribe of the Nereidiens non tentacules. Their description was on 
the whole good, though they misinterpreted the dorsal cirrus. 

Grube (1 851) in his ' Familien der Anneliden ' grouped the Nephthydea under his 
Tribe Rapacia as the fifth family between the Lycoridea and Phyllodocea. In his 
subsequent papers he does not seem to have directed further attention to the systematic 
position of this type, though in the Annelids of the i Gazelle ' (1877) he put them 
between Goniada and Cirratulus. 

Dr. George Johnston arranged his Nephthacae between the Nereids and the 
Phyllodocids. 

Kinberg 2 did not much change the characters of the family as described by Grube. 
His chief features were — the flattened head with two antennae (tentacles) and two palpi, 

1 'Ann. Nat. Hist./ ser. 7, vol. v, p. 254. 

3 « Annulata Nova." < Ofver. Af. k. Vet-Akad/ Forh., 1865, No. 2. 



NBPHTHTDID^B. 5 

the cylindrical proboscis with its terminal and lateral papillae and its two maxillae, together 
with the fusion of the dorsal cirrus and the branchia, and the lamellar (he called it 
branchial) condition of both divisions of the foot. He adopted a similar arrangement to 
that of Grrube — placing the Nephthydea between the Nereiclea and Phyllodocea. 

Malmgren first (1865) placed the J^ephthydea (Grube) after the Phyllodocea, but in 
his later publication (1867) he altered this arrangement, for they come between the 
Sigalionidae and the Phyllodocidae. The author may have found that their relationships 
were such that they most naturally occupied this position. 

In his remarks on the general characters De Quatrefages (1865) explains the so- 
called compound bristles of certain species as the result of partial fractures of these 
organs, with a proneness to split at an angle to the long axis. He describes the cephalic 
ganglia as subdivided into a main and three accessory ganglia connected by a seventh 
ganglion elongated transversely. He finds an ocular apparatus joined with the main mass 
(cerveau) though there may be no trace of eyes. The oesophageal connectives give the 
twigs to the rudimentary feet of the buccal segment. He states that instead of one 
ventral vessel two distinct trunks exist. He divided the group into three divisions, those 
with four, those with three, and those devoid of antennae (tentacles). The latter division, 
however, was due to a misapprehension. 

Claparede (1868) emphasises the fact, first pointed out by Ley dig, that the muscular 
fibre separates into two distinct layers — one axial — the other cortical. This is well seen 
in JSTephthys. 

A distinct advance on the treatment of the Nephthydidae was made by Ehlers (1868), 
who corrected and arranged the known facts as well as added to our knowledge of the 
subject. The segmental organs he did not observe. He divided the family into two 
genera, viz., those with four tentacles and a single caudal cirrus (Nephthys), and those 
with two tentacles and two anal cirri (Portelia). The latter genus, however, has no 
British representative, and considerable confusion has been caused by the inclusion of 
the well-known N. cxca, 0. Fabricius, under it. In classifying the known species of the 
genus Nephthys he made use of the length of the posterior lip in each division of the foot. 

Langerhans divided the Nephthydidae into two groups according to the number of 
longitudinal rows of papillae in the proboscis. The first of these has 14 longitudinal 
rows of papillae on the proboscis, as in N. Griibei and others. In the second group the 
proboscis has 22 rows, as in N. cseoa. 

Levinsen l (1883) placed the ISTephthydidae (his Nephthydae) as the second group of 
his Phyllodociformia, and thus prominently kept up the relationship with that family 
without associating it with others considerably divergent in structure. 

Pruvot 3 (1885) found in Ne/phthys a stomato -gastric centre (anterior division of the 
brain), and an antennary centre (middle and posterior divisions). A nuchal organ is 
present posteriorly. The group agrees with Phyllocloce in this respect. 

1 Viclensk/'Meadel. Nat. Hist. Kjobenhavn, Aaret 1882/ 1883, p. 180. 
3 < Arch. Zool. Exper/ ser. 2, HI, p. 48-49, 1885. 



XEPHTHYS. 



Genus XXIX. — Xephthys, Guvier, 1817. 

Head as in the family ; both tentacles present. Rarely with a pair of eyes. Pro- 
boscis of two regions, with 22 distal rows of papillse in extrusion, a double arch of bifid 
papillse round the aperture, and a pair of horny teeth internally. Body and feet as in the 
family. Foot with the dorsal and ventral cirri either conical or foliaceous. LamellaB 
generally well developed. Bristles of two kinds, shorter barred bristles anteriorly in each 
division, and elongated forms with a flattened though narrow blade serrated at the edge 
(from minute rows of spikes) behind. From the formation of the foot they all occur on the 
anterior surface. Xerve-cords in the typical position. Segmental organs as in the family. 

In JSfephthys cseca the dorsal muscles fuse in the middle line which is only distinguished 
by being the thinnest part of the sheet. They widen on each side and end in transverse 



iih (L/>// /&2&?^- ~. 

11 : (i i - '."• •' -" " ••' ■ .- " 

orti 



Fig. 37. — Magnified view of the nerve-area of the ventral cord of Nephtliys cseca. The fibres of the oblique 

muscles (o.m.) occur below. 

section in an outer flap quite free from the thin circular layer of the region. From the 
shelving outer edge of the longitudinal layer on each side to the border of the foot two 
series of powerful fibres are conspicuous. The first is a horizontal layer which 
commences as a narrow continuation of the circular and passes outward in a somewhat 
fusiform manner to the dorsum of the foot. The second assumes the form of strong 
oblique bands which slope from above downward and outward from the dorsal edge to 
the foot. Besides these a separate longitudinal band occurs at the shelving tip of the 
great dorsal layer. 

The ventral longitudinal muscles form, in transverse section, two large ovoid masses 
on each side of the median nerve- cords. They are bounded externally by a strong band 
of oblique fibres from above, whilst superficially they have the cuticle, hypoderm, and 
some circular fibres. 

The fasciculi of both dorsal and ventral longitudinal muscles have their long axes 
directed inward, and thus a somewhat radiate aspect is given to the transverse sections. 

The nerve-area in the centre is shaped somewhat like an inverted crown, the convex 



NEPHTHYS. 7 

edge in section being inferior. Into the straight npper edge, as well as the upper lateral 
region, the powerful oblique muscles are inserted — separated from the nervous tissue 
only by the tough sheath. The action of these and of the adjoining longitudinal ventral 
muscles would not appear to support the theory of quiescence of the trunks advocated by 
some zoologists. 

The area (Fig. 37) is subdivided by a median septum which passes from the ventral 
fissure to the dorsal edge, though in some sections it is obscured dorsally. The broad 
circumference of the area forms a dense plexus of fibres with intervals; whilst the 
upper median region on each side is composed of more lax tissue (which stains 
differently) viz. of prominent fibres — transverse or oblique in direction — with granular and 
gelatinous intermediate substance as well as small spaces in the preparations. On each 
side of the median line in this area are two conspicuous neural canals, the inferior the 
larger, the next considerably less, besides a variable number of others. There are, 
especially near the two large inferior canals, and at the outer border of the central area, 
several deeply stained granular nerve-cells. In some sections four canals are seen 
on each side of the middle line and in a diminishing series from below upward, in others 
only two with various minor apertures are visible. 

To the exterior of the nerve-cords is the tough layer of limiting membrane which 
splits into fibres at the ventral median raphe. At the outer angle ventrally is a triangular 
area occupied by oblique fibres which slant mainly from without and below inwards. 
It is difficult to say whether any traces of circular fibres (circular coat) exist between 
the limiting membrane and the cuticle, though such is probable. 

Cunningham 1 (1888) observes : "In Nephthys, instead of the typical canal on the inner 
side of each (nerve-) cord, there are two large canals, one above the other, with a smaller 
one between them. There is also a smaller canal in the external side of each cord, and 
still smaller ones in the substance of the cords. The nerve area here is not separated 
from the epidermis." 

Following the hint of Savigny, Cuvier separated the genus (under the name of 
Nephthys) from the Nereids; and soon a more complete account of the characters was 
published by Adouin and Milne Edwards. 

The generic characters given by De Quatrefages (1865) are, briefly, those of his 
predecessors. 

Kinberg's 2 description of the genus was : — " The two antennas and the two palpi 
similar in form, marginal ; bristles both annular and smooth ; the two maxillse nail-like 
and depressed." He at the same time established two new genera, viz. Aglaophamus 
and Aglaopheme. The former, originally from Banks Strait, has also been found in 
Europe. 3 

Ehlers (1868) grouped the various forms under the genus Nephthys thus : — (1) The 
posterior lamella? in both divisions of the foot in all the segments long, e.g., N. cseca. 
(2) The posterior lamella? of the inferior division in all the segments long, N. Hombergii, ■ 

1 < Q. J. Micr. So./ n.s., vol. xxviii, p. 270. 

2 ' Ofvers. K. Yet. Akad./ Fork, No. 4, p. 239, 1865. 

3 c Ann. Nat. Hist./ ser. 7, vol. v, p. 262. 



8 NEPHTHYS (LECA. 

N. cirrosa. (3) The posterior lamellae of both divisions of the foot long — only in the 
anterior segments, K discors. (4) The posterior lamellae in both divisions of the foot in 
all the segments short, N. ciliata, N. longisetosa, JV. incisa. He comprehended under 
Nephthys, Aonis, Savigny, Biplobranchus, De Quatrefages, and Aglaophamus, Kinberg. 

In P. Tauber's critical revision 1 of ' Danish Annulata,' four species of Nephthys are 
entered, but he includes N. longisetosa, (Ersted, under N. Hombergii, and appears to be 
inclined to do the same with N. incisa, Mgrn. One of the species is N. paradoxa, Malm. 

Levinsen 3 in his systematic account of the Norwegian Annelids gives eight species 
of Nephthys, but as JSf. Malmgreni and N. minuta of Theel and N. paradoxa and 
A. emarginata of Malm are four of them, further revision is needed. 

Baron de Saint- Joseph 3 points out the importance of the foot in diagnosing the 
various forms, and the necessity for contrasting these organs from the same region of the 
body. In criticising Wiren, who groups certain species together on account of the 
variations of the feet in any given example, he truly observes that though these variations 
do occur in each, yet they essentially differ from each other. 

The large foliate organs of A. trlssophyllus, Grube, from Naples, show how these 
lamellae can be developed. Moreover, the great elongation and comparative straightness 
and slenderness of the bristles of this species are diagnostic. 



I. GrEOUP WITHOUT A LONG UNPAIRED CIERUS. 

1. Nephthys oeca, 0. F. miller, 1776. Plate XLIII, fig. 1; fig. 2 var. ciliata; 
Plate LVII, figs. 1-5; Plate LXVI, figs. 1-3— foot; Plate LXXVII, figs. 1-P— bristles. 

Specific Characters. — Head rounded in front ; eyes usually absent (two small on 1st 
segment, St. Joseph) and two papillae (sensory) at posterior part of head. Body twelve 
or more inches long, 130-136 segments ; exserted proboscis with twenty-two distal rows of 
papillae, ^ve, or occasionally six, being in each row. The lip with ten forked papillae 
and a median simple one at each side. Dorsal lamella of the foot fan-shaped and 
prominent, its deepest part being external ; dorsal cirrus short and tapering ; branchial 
process coiled externally, of moderate length, with a semicircular flap of the spinigeous 
lobe at its base anteriorly. Ventral division with a large, broadly lanceolate and pointed 
lamella continuous with the fleshy lobe of the region ; ventral cirrus of moderate size 
slightly flattened and conical. Subulate (barred) bristles typical. The capillary bristles 
have a flattened blade with a close series of fine spikes. Segmental organs opening at the 
base of the feet ventrally. Tail terminating in a single long cirrus. Muscular walls of 
body typical. 

1 Kjobenhavn, 1879. 

2 Yidensk, ' Meddel. f. d. natudh. For. Kjobenli./ 1883. 



3 i 



Ann. Sc. Nat/ 8 e ser. xvii, p. 1, 1894. 



NEPHTHYS C^ECA. 



Synonyms. 



1758. Nereis cserulea, Linn. Syst. Nat., eel. 10, i, p. 654. 

1767. „ „ idem. Ibid., ed. 12, i, pt. 2, p. 1086. 

1776. Nais caeca, 0. F. Muller. Zool. Dan. Prodr., p. 219, No. 2653. 

1780. Nereis cserulea, Fabricius. Fauna Groenl., p. 298. 

} , „ cseca, idem. Ibid., p. 304. 

1789. „ ciliata, 0. F. Muller. Zool. Dan., iii, p. 14, pi. 89, f. 1—4. 
1791. „ cserulea, Linn. Gmelin Syst. Nat., ed. 13, I, pt. 6, p. 3117. 

„ „ cseca, Gmelin. Ibid., p. 3119. 

1799. „ „ Fabricius. Skriv. Nat. Selsk., v, p. 185, Tab. iv, figs. 24—29. 

1806. „ „ Tin-ton's Gmelin, iv, p. 90. 

1820. Aonis (Nereis) cseca, Savigny. Syst. Annel., p. 45. 
1825. „ cseca, De Blainville. Diet. Sc. Nat., xxxiv, p. 451. 

1828. „ „ idem. Ibid., lvii, p. 483. 

1829. White Worm, Lurg or Lurgan, Drummond. Loud. Mag. Nat. Hist., ii, p. 121. 
1835. Nephthys margaritacea, Johnston. Ibid., viii, p. 341, fig. 33. 

1842. „ cseca, CErsted. Kroyers' Nat. Tids., p. 123. 

1843. » „ idem. Groril. Dorsibr., p. 193 (41), f. 73, 74, and f. 77—86 (excl. fig. 78). 
„ „ longisetosa, idem. Ibid., p. 195, fig. 75 and 76. 

1844. „ bononensis, De Quatrefages. Ann. Sc. Nat., 3 e ser. 14, p. 352, pi. 9, fig. 2. 

margaritacea, Thompson. Ann. Nat. Hist., xiii, p. 439, and Rept. Brit. Assoc. 1843, 
p. 273. 

1845. „ „ Johnston. Ann. Nat. Hist., xvi, p. 456. 
1851. „ cseca, Grube. Fam. Annel., pp. 53 and 128. 

1853. Nereis (Nephthys) lineata, Dalyell. Pow. Creat., ii, p. 146, pi. 21, f. 4 — 10. 

„ Nephthys ingeus, Stimpson. Mar. Invert. Grand Manan, p. 33. 
I860. „ cseca, Packard. Annel. Caribou, Canacl. Nat. and Geol., vii, p. 418 (fide aut.). 

1862. „ „ Sars. Geol. og Zool. Reise, p. 50. 

1863. „ „ Stimpson. Annel. Greenland, Proc. Acad. N.S. PhilacL, p. 140. 
1865. „ cseca, Malmgren. Nord. Hafs-Ann., p. 104, Tab. xii, fig. 18. 

„ margaritacea, De Quatrefages. Anneles I, p. 423. 

„ bononensis, idem. Ibid., p. 425. 

„ (Erstedi, idem. Ibid., p. 427. 

Portelia cseca, idem. Ibid., i, p. 433. 
Nephthys margaritacea, Carrington. Annel. of Southport, p. 8. 

„ „ Johnston. Cat. Brit. Mus., pp. 167 and 342, fig. 34. 

1866-69 „ cseca, Packard. Mem. Bost. Soc. N. H., i, p. 294. 1 

1867. „ „ Malmgren. Annel. Polych., p. 18. 

„ „ ,, Parfitt. Trans. Devon. Assoc, ii, pt. 1, p. 231. 

1868. „ „ Ehlers. Borstenw., ii, p. 588, pi. xxiii, figs. 10—34. 

1869. „ ciliata, Mcintosh. Rept. Brit. Assoc, 1868, p. 337. 

1871. „ margaritacea, Grube. Mit. ueb, St. Vaast-la-Hougue, p. 100. 

1874. „ cseca, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 195. 

« » ,, Malm. Kongl. vet. o. vitt. samhalles i Goteborg Handl., xiv, p. 77. 



1 These Annelids (of Labrador) were also mentioned by the same author in the 'Canadian 
Naturalist and Geologist/ viii, p. 418, etc, 1863. 

59 



10 



NEPHTHYS C^CA. 



1874. Nephthys cseca, Malm. Amer. Journ. Sc., x, p. 39. 

1875. „ „ Verrill. Proc. Amer. Ass. So., 1873, p. 370. 
„ „ „ Mcintosh. Mar. Invert. St. Andrews, p. 118. 
„ „ „ Mobius. Unters. Dentsch Meer., p. 168. 

1878. „ „ Mcintosh. Trans. Linn. Soc, 2nd ser., i, p. 501. 

1879. „ „ Tauber. Ann. Danic, p. 83. 

„ Theel. K. Sv. Yet. Akad. Handl., xvi, No. 3, p. 30. 
„ „ „ Yerrill. Check List., U.S. Comm. F. and F., sep. copy, p. 7. 

1880. „ {cseca) margaritacea, Grraber. Arch. f. Micros. Anat., xvii, p. 289, pi. xxviii., 

fig. 14, and pi. xxix, fig. 15 — 18. 

1881. „ longosetosa, E. Horst. ("Willem Barents'' Bxped., 1878.) Niederl. Arch. f. Zool. 

Suppl. 1, 5, p. 8, Taf. f. 1. 
„ „ ciliata, idem. Ibid., p. 7. 

1883. „ caeca, Wiren. Chastop. ' Vega' Exped., p. 392. 

„ „ „ Levinsen. Yidensk. Meddel. Foren., Kjoben, p. 217 (Syst.-geogr. Overs. Nord. 

Annel., p. 60). 
1886. „ „ Schack. Anat. Histol. Untersach., Kiel. 

1890. „ „ Malaquin. Rev. Biol., v, Annel. Bonlon, 32. 
„ „ bononensisj idem. Ibid. 

„ „ cseca, Marenzeller. Annel. Beringsm., p. 1. 

„ „ „ Giard. Bull. Sc. Fr. Belg., t. xxii, p. 76. 

1891. „ „ Hornell. Trans. Biol. Soc. Liverp., vol. v, p. 244 (exclus. synon.). 
1893. „ „ Levinsen. Yidensk. UcL, " Hauchs," p. 337. 

1896. „ „ Michaelsen. Polych. deutsch Meere., p. 25. 

1899. „ „ (sexual var.), Mcintosh. Nat. Sc. (May), p. 375. 

1900. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. v, p. 257. 

1901. „ „ Whiteaves. G-eol. Snrv. Canada, No. 722, p. 82. 

„ „ ,, Johnson. Proc. Bost. Soc. Nat. Hist. vol. xxix, No. 18, p. 401. 

1904. „ ,, Allen. Journ. M.B.A., n.s., vol. vii, p. 225. 

1906. „ ,, (movements) Bohn. Ann. Sc. Nat. 9 e ser., t. iii, p. 107. 

„ „ „ (segm. org.) Fage. Ibid., p. 286. 



Habitat. — Common everywhere around the shores of Britain from Shetland to the 
Channel Islands in sand near low-water mark, and extending some distance shorewards. 
In the outer Hebrides it would seem occasionally to be in the way of deluges of rain, 
but probably the moist sand preserves it from the injurious influence of fresh water, as 
described by Dr. Drummond. It is often accompanied by Arenicola in the sand or muddy 
sand. It is also tossed on shore in storms after dislodgment from its sandy retreats or 
its haunts amidst fragments of shells and nullipores. 

Extends to the shores of France (Zostera-fields, De Quatrefages), Northern 
Europe, Spitzbergen, and Greenland. American coasts, both east and west. — Pacific, 
Puget Sound (Johnson). Canada (W.C.M.). 

Head (Plate LVII, figs. 1 and 2) more or less square, truncate anteriorly with a 
short conical tentacle at each angle. Behind and beneath the latter is another somewhat 
broader conical tentacle (palpus, Kinberg). In lateral view the head is shovel-shaped — 
thinning off towards the tip. In the slanting (converging) furrows at each side posteriorly 
is a little elevation or flap (with a lens-like apparatus, Ehlers). Brain reddish. 



NEPHTHYS C^CA. 11 

Mouth (Plate LVII, fig. 2) situated between a pair of crescentic fleshy lips, which at 
their anterior attachment show a little fold or papilla — it may be sensory in function. 
Posteriorly, at the mouth, the lips also have a free curve in front of the median 
elevation indicating the proboscis. The latter in extrusion is a massive pinkish iridescent 
organ shaped like a float or pear (Plate LVII, figs. 3 and 4), and scabrous from flat papillae, 
especially towards its distal region, which has twenty-two longitudinal rows of them. 
These increase in length distally, and each row has about five, or occasionally six papillae, 
which in ordinary spirit-preparations have a tendency to be directed backward.. In some, 
one or two small isolated papillae are scattered here and there on the region behind the 
former. The median dorsal region has two rows which run forward to the peak at the 
wide band which marks the dorsal angle of the aperture of the organ, and the terminal 
papilla on the left or right is a little longer and more slender than the rest. On the 
ventral surface a similar arrangement holds, one of the rows having an advanced 
and somewhat longer papilla. From each row a band proceeds to the groove 
between the terminal ridges of the lips, ten of these ridges being boldly marked 
(Plate LVII, fig. 4), whilst a small median at each side completes the series. Each 
ridge is separated from the adjoining ones at the margin of the lip and is continued a 
short distance inward (in extrusion). From the outer side of the free tip arise two 
flattened tapering papillae, the outer being somewhat longer. The dorsal and ventral 
rows of the papillae diminish a little on each side of the middle line, the intermediate 
one at each side, which is single, being the shortest. On slitting the proboscis a shelf 
occurs inside the dorsal and ventral lips, then the smooth wall of the canal trends inward 
for a short distance and breaks up into four prominent muscular ridges, with deep inter- 
mediate grooves. One furrow therefore occurs opposite the middle of each lip, and 
another at the intermediate line. At the distal end (in extrusion) of each central furrow 
is a conical blackish horny tooth, the base having a visible spur directed to the left 
(Plate LVII, fig. 5). The four muscular ridges, in all probability, can be protruded to 
the tip of the organ, and thus the horny teeth will grasp the prey. When it penetrates 
the sand uncovered by the tide it strikes the surface with the proboscis more frequently 
than in Glycera, and is soon immersed. 

In some large examples the proboscis is deeply coloured with black pigment best 
marked in the long papillae of the lips and distal rows, and which increases the iridescence 
of the organ. 

In an example from Salthill, Co. Dublin, a hollow process projected from the base of 
the extruded organ, the tip of the diverticulum being warty. 

Body. — Elongated (ranging to a foot or more in length), slightly tapered anteriorly, 
and more distinctly so posteriorly, and with numerous segments, 140 — 150. Colour pearl- 
grey with pinkish iridescence. This is but slightly marked in the middle line either 
dorsally or ventrally, for a comparatively smooth surface of a fine pearly lustre occupies 
those regions. The dorsum is convex, the ventral surface usually has a median groove, 
which in front splits at the central prominence and runs forward on each side to terminate 
at the mouth. The region in the fork is marked by a close series of longitudinal furrows, 
which in extrusion of the proboscis are much stretched so as to reach from side to side. 



12 NEPHTHYS (LECA. 

The peristomial segment is peculiarly flattened, and separated dorsally by a more 
distinct lateral cleft. It bears dorsally a foot-papilla with a spine and a palisade of barred 
bristles (curved backward) in front, and a line of slender elongate simple bristles behind 
the spine. To the exterior is a flattened dorsal cirrus with a subulate tip. Below and 
slightly in front of the foregoing is a tuft of simple, slender and straight bristles which 
taper to a point, while at the outer border is a short cirrus — flattened like the foregoing 
and with its upper basal edge slightly extended or bulging. The two cirri form the 
tentacular cirri of E biers. 

The second foot presents well-marked dorsal and ventral divisions, though smaller 
than in the typical form. Dorsally the end of the spine is in the centre of a rounded 
area or papilla which has in front a backwardly curved row of long barred bristles of large 
size and with long filiform tips. On the posterior curve of the papilla is a row of long 
tapering bristles of the same type as the serrated forms, but the serrations have not yet 
attained distinctness, though traces are visible. They also curve backward. A short 
cirrus flattened especially at the base arises from the posterior and outer edge of the 
division. A small lamella with a free edge passes from the inner border of the spine- 
papilla. The inferior division has a broadly lanceolate lamella of smaller size than in the 
succeeding feet, but is similar in outline. The somewhat flattened ventral cirrus occurs 
at its inferior border. The fleshy part of the foot terminates superiorly, in front of the 
foregoing, in a short blunt cone. The long tapering bristles, with slightly flattened blades 
above the shaft, hold their normal position in front of the lamella, while the barred kind 
— still with long hair-like tips — form an anterior row separated from them by the spine- 
papilla. 

The third foot has superiorly a somewhat elongated fan-like lamella to the inner side 
of the lanceolate and flattened dorsal cirrus. The bristles have the same disposition — the 
anterior barred forms being long and curved backward. The small lamella to the inner 
border of the spine-papilla is better marked than in the former foot. The dorsal cirrus 
is much flattened and of irregular outline, being bifid at the tip and extended at its outer 
and basal edge. The inferior division has a larger lamella, and the ventral cirrus is 
lanceolate. The fourth foot shows the branchial lamella dependent from the lower edge 
of the base of the dorsal cirrus. It is a comparatively small process, but has the character- 
istic hook-like curve. The dorsal cirri diminish in size while the branchiae and lamellae 
increase from before backward. 

Anteriorly the typical foot (Plate LXVI, figs. 1 and 2) presents a large fan-shaped 
lamella projecting like a crest from the posterior border of the upper division, its free 
inferior edge coming off at a blunt angle from the outer. The fleshy base of the foot, with 
the posterior part of which this is continuous, slopes from above downward and outward, 
and terminates in a rounded disc or lobe above the cirrus. Immediately below is a short 
tapering cirrus, and after an interval or notch the hook-shaped branchial process, which is 
of moderate length and has a slight papilla at the base externally. A semi-circular flap 
lies over the base of the branchial process anteriorly, being the inferior termination of the 
flattened lamina between the two rows of bristles. No free flap guarding the camerated 
bristles anteriorly occurs in this species. In front of the lamella is a row of pale elongate 



NEPHTHYS GMGA. 13 

serrate bristles (Plate LXXVI, fig. 1), which extend outward fully three times the longer 
diameter of the fan. The serrated edge, simple in lateral view, presents a close series of 
transverse spikes (Plate LXXVI, fig. d) as indicated by various authors. Beyond the 
serrated region the bristle extends as an extremely attenuate process and tapers to a fine 
smooth point. A narrow fleshy ridge with the tip of the spine projecting externally 
next follows with a row of the pale barred tapering bristles in front. These show a series 
of transverse bars (Plate LXXVI, fig. 1 b, and c, c 1 ) which in lateral view (d) have an 
interrupted edge. The bars are probably modifications of transverse rows of spikes — 
forming the so-called camerated arrangement. Dr. Thomas Williams was thus incorrect 
in saying that one of the rows of bristles is placed in front and the other behind the cirral 
lamina. 

The foregoing constitutes the dorsal division of the foot, and it is separated by a 
considerable interval from the ventral division, the intermediate region being occupied by 
the branchial process. 

The inferior lamella is broadly lanceolate, pointed, and with the longer edge ventral- 
ward. It is continuous with the fleshy lobe, the tip of which slopes from below upward 
and outward, the reverse of the arrangement in the upper lobe. The fleshy part 
terminates superiorly in a thickened adherent point. The long pale bristles similarly 
pass out close to the base of the lamella, and extend about the same distance. A narrow 
fleshy ridge (corresponding to the foot proper) separates them from the row of barred 
bristles in front. The conical ventral cirrus is slightly flattened and of moderate size. 

In the posterior feet both dorsal and ventral lamellse, branchiae, and dorsal and 
ventral cirri diminish as in iV. Rombergii (Plate LXVI, fig. 8), and in the last feet only 
a minute trace of the cirri remain. The bristles also decrease in length but retain the 
same fundamental arrangement. 

The body terminates in an anus, with a median cirrus ventrally. 

Loxosoma3 were studded on the branchiae of a large example sent from Montrose 
Bay by the late Dr. Howden. De Saint Joseph mentions Cothurnia maritima as common 
on the bristles of those at Dinard, and a form probably identical is found at St. Andrews 
and elsewhere, in great numbers, on the same organs; indeed Infusoria are very 
frequently found parasitic on the Nephthydidee. 

A form (Var. ciliata, Plate XLIII, fig. 2) which approaches N. caeca in the 
feet, and which may be Dr. Johnston's A", longisetosa, occurs at St. Andrews and 
Montrose occasionally in May and June. The largest examples were about six inches long, 
and they ranged from this downward to two or three. In structure the foot (Plate LXVI, 
fig. 3) has longer divisions, a long, narrow dorsal cirrus, and the superior lamella is 
slightly less. The length of the slender bristles, which give a remarkably hairy appear- 
ance and great breadth to the animal, is diagnostic. The foot generally agrees with that 
of N. cd3ca, but the superior lamella is often distinctly less and does not project so far 
outward. The spinigerous region, moreover, ends in a more distinctly rounded flap than 
in the ordinary form— projecting outward above (and in front of) the dorsal cirrus. No 
special differentiation occurs in the fleshy parts of the inferior lobe, although the upper 
part of the fillet from the barred bristles is somewhat more distinct. The bristles consist 



14 



NEPHTHYS (L3ECA. 



of long delicate simple bristles without the marked blade of the serrated forms, and 
tapering to a fine point. They are of great length, are pale (Plate LXXVI, fig. 1 e), and 
never attain the diameter of the more distinctly serrated forms, a few of which are found 
at the ventral margin of the inferior division and elsewhere. The latter are recognised 
by their yellowish hue, a colour which enables even broken fragments in the upper 
division to be readily recognised. Why these long, smooth, or very minutely serrated 
bristles should be developed in this way is unknown. ~No sexual elements were 
observable in the examples after preservation. No change occurs in the barred bristles. 
It may be noted that Ehlers likened the foot in Nevldhys to the epitokous condition in 
Nereis. 1 If such be an epitokous condition it would not therefore be complete. 

The weight to be placed on small changes in the development of parts of the foot, 
e. g. enlargement of lobes, is not yet sufficiently understood. 

Reproduction. — At the reproductive season (e. g. towards the end of March at St. 




so 



ov. 



vm orrv ric. 



Fig. 38. — Section of a female Nephthijs cseca distended with ova. s o. Segmental organ (nephridium). d. Alimentary 
canal, d.m. Dorsal longitudinal muscle, d.v. Dorsal blood vessel. n. c. Nerve cord. o.m. Oblique 
muscles, ov. Ova. p. v. Perivisceral (ccelomic) space, v.m. Ventral longitudinal muscles. 

Andrews) the body-cavity is filled with large ova (Fig. 38), which in section are found 
chiefly below the gut and in the cavities of the feet, though they also pass above the gut. 
The changes in the thickness of the muscular development in this condition may be 
ascertained by contrasting this figure with Fig. 34. In the males a similar condition 
prevails, the sperms likewise appearing above the gut and the dorsal vessel. 

The larva3 of JNTephthys have been long known, though their association with the 
genus was not always understood. Thus a larva described by Busch 2 apparently agrees 
with a stage figured by Claparede and Mecznikow with curved tufts of cilia. 

Claparede and Mecznikow 3 found several stages, though they were uncertain as to 
the relationships of some, and the same may be said in regard to Fewkes. 4 



1 ' Borstenwiirmer/ ii, p. 584. 

2 'Beobach./ p. 68, Taf. viii, fig. 7. 

3 'Beitrage/ 1869, p. 187, figs. 3, 3a, and 3d. 

4 < Stud./ 1884—85, p. 180, Plate iv, figs. 1—12. 



NEPHTHYS GMCA. 15 

An early stage is described and figured by Claparede and Mecznikow in the form of 
the usual trocophore. It is pear-shaped without trace of segmentation, and with a ring 
of cilia in front of the mouth, and a pencil of cilia anteriorly at the apex- The 
alimentary canal bends forward toward the sensory area anteriorly — the dilated region 
being greenish — and then curves backward to the conical posterior end. A brown 
eye-spot is on each side. 

Both Claparede and Mecznikow, and Hacker, describe and figure the next stage. 
Hacker (1896) gives the first metatroch stage of Nephthys scolopendroides as some- 
what pear-shaped with a green-flecked anterior end and a pair of eyes. Two brown 
pigment-lines occur behind the prototroch, and green and red pigment behind the 
paratroch. The stomodeum is transverse with a three-lobed upper lip. The mid-gut has 
brown and blue pigment and oil globules, and the anus has blue pigment. This author 
finds that the prototroch has an upper small and two lower streaks or grooves (Reifen). 
The paratroch is one-rowed. The sides of the larva are crenate (four segments), but 
without bristles in one elsewhere procured. 1 

A stage in which the large globules of yolk occur far forward, just behind the eyes, 
is given (Plate L, fig. 1). ISTo colour is present. Short bristles are distinct in three 
segments, and they do not yet show the transverse strise. Round diatoms occurred in 
the alimentary canal. The madder-brown tip to the tail is more diffused, not clearly 
forming two bands. The mouth in the sketch is on the under surface, so only the 
large globules are seen. In a larval form with eight bristled segments (Plate L, 
fig. 2) the head is broad and horseshoe-shaped, with a general pinkish-yellow hue, 
and two madder-brown eyes. The proboscis is indicated behind — as far as the 
commencement of the second body-segment. The alimentary canal following has 
numerous large clear globules, and terminates posteriorly in a narrow rectum, and a 
vent at the median short style. The colour of the gut is bluish-green. Two boldly 
ciliated regions are present — the preoral and anal. The latter is marked by a double 
madder-brown band — one on each side of the cilia. A short foot behind the head bears 
the first bristles, and thereafter seven pairs occur. At the side of each segment is a 
small foot-lobe. The larva swims actively by aid of its cilia. The lateral bristles are strong 
and have the characteristic transverse markings of Nephthys. Diatoms and debris were in 
the alimentary canal. In Fig. 3 is an example captured a few days later (28th October) 
in which the general form of the head and body has undergone alterations. 

Two later stages are figured and alluded to by Claparede and Mecznikow, one in 
which there are eight pairs of feet, eyes with lenses, and teeth in the proboscis. Both 
rings of cilia persist. In the more advanced stage the body is elongate and the preoral 
ring of cilia less distinct. The characteristic bristles occur on the feet. The authors 
suppose that the species is Nephthys scolopendroides — a common one at Naples. 

The older stage, of date October 28th (Plate L, fig. 3) presents a more distinctly 
differentiated head, which is separated from the peristomial segment, and the area of the 
proboscis is clearly outlined. The greenish-blue colour of the intestine is still conspicuous. 
Though only nine pairs of bristle-tufts occur, the form is considerably older than that 

1 Plankton Exped. ' Pelagic Polych./ p. 10, taf. i, fig. 4. 



16 NEPHTHYS (LSECA. 

represented in Fig. 2. Similar advanced stages were procured in the Bay of Kiel by 
Leschke in November and December. 

De Saint Joseph says that, like Nephthys scolopendroides, N. cseca is mature from 
November to April. 

Habits. — Abundant in sand near low water-mark along with Mag dona and other 
forms, and it bores through the moist sand with great rapidity. It is likewise procured 
in considerable numbers under stones on a sandy and gravelly (shell) surface with a 
slight coating of mud, in company with a few specimens of Girratulus. In a vessel 
of sea-water the branchiae are borne in a loop at each side, and now and then jerked 
a little. 

It forms an excellent bait for flat and other fishes, and is used whenever it can be 
obtained. As might be anticipated, therefore, it is not uncommon in the stomachs 
of cod and haddock. 

The Dutch author Seba 1 (1734) figures what has been considered a Nephthys, 
but as it is from Amboina and resembles an Amphinome or Hermodice, the interpretation 
is somewhat doubtful. 

The early description of 0. Fabricius (1779) is fairly accurate, both in regard to 
external appearance and habits, but he represents two caudal cirri in his figure — an 
error probably due to the artist. 

Colonel Montagu (1808) in all probability represents this species (MS. vol. Linn. 
Soc, Plate VIII, fig. 3) as Nereis cseca. 

Savigny (1820) simply copied the observations of Fabricius, but he saw that a 
new genus was necessary. 

Dr. Gr. Johnston (1835) corrected several points in the earlier descriptions, and 
increased our knowledge of its habits and haunts. He thought the lamellae of the 
feet were useful in swimming, which probably they are, though the powerful muscles 
of the body-wall are even more important. 

CErsted (1845) made further improvements in the description and figures, and 
differentiated the species from N. Hombergii. The finer characters of the bristles 
escaped him. 

Busch's fig. 6, Taf. VIII s (1851), shows a young larva of Nephthys or Nereis with 
the hook-like tuft of cilia on each side in front of the eyes, and two posterior papillae. It 
may however relate to another form, as the tufts are behind the eyes in those figured by 
other authors. Fig. 5 of the same memoir would seem to be a younger stage. 

The Nereis (Nephthys) hirsuta of Daly ell 3 may be a small example of this species or of 
an allied form. 

De Quatrefages (1865) seemed to be in considerable perplexity concerning this form 
— of which he made three species, viz. N. margaritacea, N. CErsteclii, and Portelia cseca. 
He had perhaps relied too much on uncertain figures. From the same author's descrip- 
tion and the remarks of M. Malaquin," N. bononensis, De Quatref ., is probably the same 

species. 

1 ' Thesaurus/ i, pi. lxxxi, fig. 7. 

2 ' Beobach. u. Anat. u. Entwickel. Wirb. Seethiere/ 

3 < Pow. Great./ ii, p. 145, pi. xxi, fig. 1—3, 1853. 



NEPHTHTS HOMBERGII. 17 

Ehlers (1868) gives a minute account of the structure of this species, though he did 
not make out that of the segmental organs. 

V. Grraber * makes the oesophagus of Nephthys caeca one of his types in describing the 
structure of the region in the annelids, the layers from within outwards consisting of the 
intima, the pigmented protoplasmic (hypodermic ?) layer, with glands, the subfibrosa, 
the muscular, and externally the peritoneal. 

Wiren 2 considers, after quoting the descriptions of the feet, that the N. ciliata 
of Malmgren (1865) is this form, as is also the N. cirrosa and N. ciliata of Ehlers, 
the N. emarginata of Malm, and the N. longisetosa of Horst. 

Theel's N. Malmgr eni from the Norwegian North Atlantic Exped., 1879, and 
Hansen's N. atlantica are united by Wiren. 

Mobius 3 was of opinion that this species included N. ciliata, N. Hombergii, and 
N. incisa, but the finer points of distinction were not considered. Theel 4 was inclined 
to a similar view, but thought further researches were necessary. 



2. Nephthys' Hombergii, Lamarck, 1818. Plate XLIII, fig. 6; Plate LVII, figs. 6-7 a; 
Plate LXVI, figs. 4-8— feet ; Plate LXXVIL figs. 2-5— brist. 

Specific Characters. — Head rectangular, more elongated than in N. caeca, tentacles 
more slender ; posterior pair larger than the anterior (Ehlers) ; inferior tentacular cirrus 
with its wide base merging into the anterior lip. Body with 130 segments. Proboscis 
with a slightly longer terminal papilla in the mid-dorsal line. Dorsal lamella broad and 
low, projecting beyond the tip of the foot; dorsal cirrus short. Branchia (commencing 
at the fifth foot) large, with a process at its base, and often curved inward; tongue- 
shaped inner flap guarding base of bristles, and another, somewhat pointed, occurs at 
the tip of the foot. Lamella of the ventral division large, directed upward and out- 
ward, broad and truncated toward the tip. Ventral cirrus short and broadly lanceolate. 
Capillary bristles shorter than in NT. caeca, shafts long with an expanded blade bent down- 
ward at an angle and tapering somewhat quickly to a delicate point. The edge is finely 
serrated. The barred (anterior) bristles approach those of N. caeca, the tips perhaps 
being shorter. Bristles throughout dark. 

Synonyms. 

1555. Scolopendra marina, Rondelet. Insect, et Zooph., in Piscibus Marinis, pt. 2 (Univ. Aquat. 

Hist.), p. 108, fig. I. 
1818. Nephthys Hombergii, Lamarck. Anim. s. Vert, v, p. 314. 



1 " Die G-ewebe u. Driisen d. Anneliden-CEsophagus," lxvii Bd. der ' Sitz.-ber K. Akad. d. 
Wiss. ; 1 Abth. Marz.-Heft., 1873, 2 pis. 

2 Chastop, ' Sibirisk och Berings Haf. "Vega" Exped/ Stockholm, 1883. 

3 ' Die zweite dentsch JSTordpol./ 1869-70. 

4 'Kongl. sv. vet. Akad. Handl./ B. 16, p. 24. 

60 



18 



NEPHTHYS HOMBERGII. 



1820. Nephthys Hombergii, Savigny. Syst. des Annel. p. 34. 

1825. Nereis scolopendroides, Delle Chiaje. Memoire, ii, pp. 401, 424, Tav. xxviii, (1) fig. 8. 

„ „ Hombergii, De Blainville. Diet. Sc. Nat., t. xxxiv, p. 438. 

1828. Nephthys „ idem. Ibid., t. lvii, p. 483, and Chsetop., Tab. 18, f. 1. 

1833. „ „ Aud. and Ed., Ann. Sc. Nat., t. 29, p. 257, pi. xvii, figs. 1 — 6, and copied 

in Cuv. Keg. Anim., pi. xv, fig. 2, Edit. Illnst. 

1834. „ „ Aud. and Ed., Annel., p. 235, pi. vb, figs. 1—6. 

1837. „ „ Cuvier. Keg. Anim., torn, iv, p. 173, pi. xv, f. 2 (and Edit. 1820). 

1838. „ „ (circulat.), H. Milne Edwards. Ann. Sc. Nat., 2 e ser., x, p. 211, pi. xii, 

figs. 3 and 3a. 

„ idem. Kegne Anim. 111., pi. la, figs. 3 and 3a. 

1840. .. neapolitana, Grube. Actin., Ecliin., n. Warmer, p. 71. 
Hombergii, idem. Ibid., p. 71. 

1841. „ scolopendroides, Delle Chiaje. Descrizione, iii, p. 99, v, p. 105, Tav. 99, f. 11, 

Tav. 102, f. 8, 23, 24. 

1842. ,, longisetosa, CErsted. Kroyers Naturh. Tidsskrift, iv, 2, p. 123. 

1843. ,, „ idem. Gronl. Dorsibr., p. 195, figs. 75 and 76. 
assimilis, idem. Annul. Dan. Consp., p. 33, figs. 93, 100. 

1851. „ Hombergii, Grube. Fam. Annel., p. 53, 128. 

neapolitana, idem. Ibid., p. 53, 128. 

assimilis, Williams. Kep. Brit. Assoc, pp. 188, 199, 235. 
1853. „ „ idem. Ann. Nat. Hist., ser. 2, vol. xii, p. 404, pi. xiv, f. 7a and b. 

1865. ,, Hombergii, Carrington. Annel. Southport, p. 8. 

longisetosa, Johnston. Cat. Brit. Mus., p. 172. 

Hombergii, idem. Ibid., p. 173. 

longisetosa, Malmgren. Nord. Hafs.-Ann., p. 106, tab. xii, fig. 20 {fide Tauber). 

Hombergii, De Quatrefages. Annel., i, p. 420. 

assimilis, idem. Ibid., p. 429. 

scolopendroides, idem. Ibid., p. 429. 

longisetosa, Malmgren. Annel. Polych., p. 19 (fide Tauber). 

Hombergii, Ehlers. Borstenw., ii, p. 619, taf. xxiii, figs. 7 and 42. 

scolopendroides, Claparede. Annel. Nap., p. 176, pi. xvi, fig. 1. 

„ Claparede and Mecznikow. Zeits. f. w. Zool., xix, p. 25 (sep. abdr.), 

taf. xiv, f. 3. 

assimilis, Kupffer. Jahresb. Com. deutsch. ' Pommerania/ 1871, p. 150. 

„ Malm. Kongl. Yet. o. Yitt. Samhailes i G-oteborg, Handl. xiv, p. 78. 

Hombergii, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 196. 

emarginata, Malm. Op. cit. Coteb., xiv, p. 77, tab. i, fig. 1. 

Hombergi, Ehlers. Zeits. f. wiss. Zool., xxv, p. 19. 

„ Mcintosh. Invert, and Fishes, St. Andrews, p. 118. 

cseca, Mobius. Jahresb. Com. deutsch, p. 168. 

emarginata, Malm. Americ. J. Sc. and Arts, x. p. 39 (fide ant). 

ciliata, Lenz. Jahresb. Com. deutsch, p. 13 (?). 

atlantlca, Hansen. Nyt Mag. f. Nat., xxiv, p. 4. 

assimilis, idem. Ibid., p. 268. 

Hombergii, Langerhans. Zeits. f. w. Zool., xxxiii, p. 302, pi. xvi, fig. 38. 

Hombergi, Tauber. Annul. Danic, p. 84. 

Theel. Annel. Nouv. Zemb. (K. Sc. Yet. Akad. Handl., t. xvi, No. 3), p. 26. 
1879. ,, assimilis, Hansen. Nyt Mag. f. Naturvid, xxiv, p. 268. 



1867. 
1868. 

>) 
1869. 

1873. 

1874. 



1875. 



1878. 
1879. 



NEPHTHYS HOMBEEGII. 



19 



1879. 



1883. 



1885. 



1886. 
1886- 
1890. 
1891. 
1893. 
1894. 
1896. 
1900. 
1901. 
1902. 
1904. 
1905. 
1906. 



Nephthys scolopendroides, Marion and Bobretzky. Ann. So. Nat. 6 e ser. viii, Art. No. 7, p. 16, 

pi. xv, fig. 2. 
„ Hombergii, Levinsen. Yidensk. Meddel. Foren Kjobenh., p. 217 (Syst.-geogr. Overs. 

Annel. nord., p. 60). 
scolopendroides (Hombergii), Prnvot. Arch. Zool. Exper., 2 e ser., iii, p. 225. 
„ Cams. Fauna Med., p. 223. 

„ Jaquet. Mitth. Zool. Stat. Neap., vi, p. 364 and pi. xxii, fig. 75 — 81. 

longisetosa, Harvey Gibson. Verm. Liverp., p. 152. 
7. ,, scolopendroides (struct, muscles), Emery. Mitth. Zool. Stat. Neap., vii, p. 371, p. xiii. 

Hombergii, Malaquin. Annel. d. Boulon., p. 33. 

„ Hornell. Trans. Liverp. Biol. Soc, vol. v, p. 244. 

assimilis, Levinsen. Vidensk. Ud. " Hauchs," p. 337. 

Hombergii, De Saint Joseph. Ann. Sc. Nat., 8 e ser. xvii, p. 3, pi. i, figs. 1 — 13. 
scolopendroides, Boule. Camp. ' Caudan/ p. 449. 

„ Mcintosh. Ann. Nat. Hist., ser. 7, vol. v, p. 258. 

„ idem. Ibid., vol. viii, p. 222. 

„ Marenzeller. Polych. Grund., p. 12. 

Hombergii, Allen. Journ. M. B. A., n. s., vol. vii, p. 225. 

scolopendroides ( = N. longisetosa Johnst.), Grraeffe. Arbeit. Zool. St. Triest, xv, p. 322. 
Hombergii, Fage. Ann. Sc. Nat., 9 e ser., iii, p. 287, figs. 11 — 14. 



Habitat. — From Shetland to the Channel Islands — along both shores, and occnrring 
alike in the tidal region and in deep water. W. coast of Ireland. Dredged at a depth of 
96 fathoms in the ' Porcupine ' Expedition of 1S69. 

Frequent on the French shores, and in the Mediterranean. Rare on the shores of 
Sweden and in the Baltic (Malmgren). Madeira (Langerhans) where it frequents water 
from 10 to 20 fathoms in depth. 3 — 400 metres (Roule) ' Caudan ' in the Gulf of 
Gascony. 

Head pentagonal (Plate LVII, fig. 8), more elongated than in N. cseca, and with 
very well marked tentacles. 

In the preparations no sense-organ or ciliated papilla is observable. Ehlers states 
that it is present on each side in the converging groove. The lobes at the sides of the 
mouth are more pointed. Delle Chiaje gives his form two distinct eyes, and though such 
are not often seen, yet in a good example from St. Magnus Bay, they are present at the 
base of the anterior tentacles (Plate LVII, fig. 8). 

The peristomial segment has the foot divided into a dorsal and a ventral division, 
each of which has a spine-papilla and the two kinds of bristles. The dorsal cirrus is 
minute, but the ventral is large. The long dark bristles are directed forward on each 
side of the head. 

The branchiae begin on the fifth foot (St. Joseph says fourth segment). 

The Body has similar proportions to that of iV. cseca, but is considerably less, no 
example yet met with having attained the proportions of the species just mentioned. It 
is iridescent pinkish in life, bluish-white along the median line dorsally and whitish 
laterally, with bright red branchiae along the sides. 

The proboscis (Plate LVII, fig. 6a) in extrusion presents similar rows of papillae 
dis tally to those in N. cmca, but the last or terminal of the median dorsal is considerably 



20 NEPHTHYS HOMBERGIL 

longer and more slender than the others, so that it is a conspicuous process. The number 
of papilla in each row is two or three, and thus the species, as De St. Joseph observes, 
is distinguished from N. caeca. Moreover, each often rises from the distal edge of a 
thickened base like the stump of an injured papilla. In some only the truncated stump 
remains. In a large Neapolitan example the two median dorsal rows converge distally, 
and beyond the point of junction the long cirrus arises. On the ventral surface the 
median row agrees with its fellows. 

The typical foot (Plate LXVI, fig. 4) is distinguished from that of N. cseca by the 
less prominent superior lamella, which has a shorter vertical diameter though its 
elongation transversely carries its outer edge beyond the tip of the foot. The bristles 
are also darker (brownish). In front a tongue-shaped flap, guarding the base of the 
bristles, occurs superiorly, while another small flap of similar shape is situated 
inferiorly, a view from the front thus presenting a bifid surface. Below the point 
of the spine is a prominent papilla. No part of the foot is more diagnostic when 
contrasting it with the homologous region in N. cseca. Its free margin is often sinuous 
or incurved. Beneath the short dorsal cirrus, which is separated by a deep notch, is the 
richly ciliated (two rows of cilia, St. Joseph) branchia often curved inwards in the prepara- 
tions, and a little less massive than in N. cseca though not shorter. The outer part of its 
base is enlarged into a projecting papilla, which in the variety kersivalensis 1 (Plate LXVI, 
figs. 5 and 6) forms a short cirrus. The serrate bristles are much shorter than in N. cseca, 
have long cylindrical shafts, and an expanded blade — bent downwards at an angle — 
tapering somewhat quickly to a delicate point (Plate LXXVII, fig. 2), the edge being 
finely serrated, and thus in all respects contrasting with the corresponding bristle in 
N. cseca. St. Joseph speaks of minutely spinose bristles in the superior division of 
the posterior segments — modifications of the elongated forms. The barred bristles 
(Plate LXXVII, fig. 3) do not offer much for comment except perhaps that the bars are 
somewhat closer, and the region is shorter. They form a vertical curved palisade in front 
of the spine-papilla, while the long serrated forms present in the line of insertion a 
considerably smaller arc, and the tips are incurved superiorly and inferiorly, so that the 
form of the bristle is diagnostic— a flattened and pointed hair-pencil. The longer dorsal 
bristles are near the inferior margin, the longer ventral near the superior margin. 
Moreover, the ellipse formed by the insertions of the two groups of bristles differ in so 
far as the dorsal have a gap superiorly, while the ventral have it inferiorly. In the 
ventral division the lamella is largely developed, directed upward and outward, and with 
a characteristically broad and somewhat truncated tip. In front are two small lamellae — 
flap-like in character and spatulate in outline — which guard the bristle-bundles, and 
make a duplication of the bifid arrangement in the superior division. Instead of the 
elongated spinigerous region of N. cseca, the part, as in the superior division, is ovoid, but 
the papilla is superior instead of inferior. The bristles agree with those of the dorsal 
division, except that the tips of the serrated forms are longer. The ventral cirrus is 
short and broadly lanceolate in outline. 

1 From the old name of Lochmaddy, North-TJist, where it is plentiful. 



NEPHTHYS HOMBEEGIL 21 

Posteriorly the lamellae, branchiae, and cirri diminish, and the gap between the 
dorsal and ventral divisions is greater. The bristles, however, retain their characteristic 
features. 

Occasionally a specimen, e. g. from Bressay Sound, shows the lower lobe of the foot 
somewhat like that of N. caeca, but the dorsal lobe and the structure of the bristles 
enable ready diagnosis to be made. A similar condition is observed in examples from 
Denmark. 

Many years ago a series common in the Outer Hebrides and elsewhere had been 
separated from N~. Hombergii by the fact that the ventral lamella in the anterior third was 
much less than in the typical form, and both lamellae posteriorly decreased more 
decidedly. The process at the base of the branchia is also longer (Plate LXVI, fig. 5 
anterior and fig. 6 posterior foot). Extended examination, however, shows this to be 
a younger stage in the growth of K Hombergii, intermediate forms leading from the one 
to the other, and it may be called variety hersivalensis. The chief distinction is that in 
the variety hersivalensis the ventral lamella in the anterior third is much less than in 
the type, and both lamellae posteriorly decrease more decidedly. 

Such distinctions, however, may be sexual. The capillary bristles of the variety 
(Plate LXXVII, figs. 4 and 5) show a somewhat more marked curve. 

Reproduction. — Specimens under two inches in length from the west Voe of Burra, 
Scalloway, in July, had eggs far advanced and floating freely in the perivisceral 
chamber. 

Claparede (1868) mentions that the ova are red, as indeed was shown by Delle 
Chiaje. 

Claparede and Mecznikow 1 describe the telotrochous larva of this species at Naples. 
The youngest stage is monotrochous with a whip of cilia in front and a ring behind the 
mouth, in front of which is a pale reddish pigment-band. The alimentary canal has 
mouth, pharyngeal region, stomach (with greenish pigment), and intestine terminating in 
an anus at the posterior end and ciliated throughout. A pair of red eyes is present in 
the larval prostomium in which the greater part of the alimentary apparatus lies. The 
monotrochous becomes a telotrochous larva by the appearance of rings behind that at the 
mouth, and the development of the anal ciliated ring. The lateral rings by-and-by 
develop bristles, the alimentary canal becomes elongated as in the figures of the British 
examples, and a form like the Naveda-l&rva of A. Agassiz and others is found. Such 
young forms are remarkable for the greenish or bluish colour of the stomachal region of 
the gut, whilst in the pharyngeal two teeth appear. Their oldest form had seven pairs of 
feet with bristles. The authors probably procured these larvae at Naples in November or 
early in December. 

Var. vasculosa. 

Fine examples of Nejphthys Hombergii were forwarded from Montrose Bay by the 
late Dr. Howden in 1870. The same year it was dredged in 25 fathoms in Bono Bay by 
the < Porcupine.' The head agrees with the typical form. The feet (Plate LXVI, figs. 
1 ' Zeitschr. f. w. Zool./ Bd. xix, p. 25 (sep. abdr.), taf. xiv, figs. 3, 3a— 3d, 1868. 



22 NEPHTHYS HOMBEKGIL 

7 and 8) are distinguished by the great development of the inferior lamella. Dorsally 
the lamella agrees very closely with that of N. Hombergii. The spinigerous lobe shows no 
boss or process below the spine as in the Neapolitan form, but this feature is absent in 
various examples both from the Mediterranean and elsewhere. The free flaps from the 
fillet — superiorly and inferiority — are somewhat less developed than in the ordinary 
examples, some having a tendency to diminution. The dorsal cirrus is shorter and less 
filiform, but the branchia is similar. The inferior division again presents a much larger 
foliaceous lamella, forming a thin fan which often overlaps the adjoining edge of the 
upper lobe. The greater part of the lamella is occupied by arborescent blood-vessels — 
leaving a narrow marginal belt of translucent tissue. The brownish capillary bristles 
are in two groups in each division of the foot, the upper division having the upper group 
shorter, while in the ventral division the lower is the shorter. They and the barred 
forms appear to correspond with those of N. Hombergii, though they are somewhat more 
slender. Posteriorly all the processes of the foot are diminished, especially the branchia 
and the inferior lamella. 

It is interesting that this vascular modification of the inferior lamella occurs not 
infrequently in examples from Montrose, and yet has not been found elsewhere than in 
the Mediterranean. 

In the intestine of an example from Montrose were muddy sand, bristles, and shreds 
of tissue of its prey (other Annelids). 

In all probability Montagu's Plate LXIX 1 represents this form. The branchia is 
apparently turned outward. 

If Cuvier's species as described by Savigny (1820) be the common form then the 
title JV". Hombergii, Cuv., has the priority. Savigny's remarks are by no means dis- 
tinctive, since he does not indicate the characters of the ventral division of the foot with 
sufficient precision in contrast with the dorsal. As French authors, however, identify 
Cuvier's specimens with this species it is well to accept it. Besides, this decision is 
substantiated by the adoption of the same title by Lamarck in 1818. 

The early description and figure of Delle Chiaje (1825) give no diagnostic features 
for the species, nor does his later publication (' Descrizione ') add further information. 
G-rube, however, had in the meantime found the same form in the Adriatic and 
Mediterranean, as Delle Chiaje himself notes, and though he separated it from Nephthys 
Hombergii, Aud. and Ed., it would appear, and as Claparede considered, that it refers 
to that (the present) species. Grube, indeed, subsequently associated it with (Ersted's 
N. assimilis. It would have been difficult to connect Delle Chiaje's species with that 
of Milne Edwards' if the workers at Naples had not satisfied themselves, for he describes 
and figures two very distinct eyes which are not often seen in the British representatives, 
and the presence of two caudal cirri in the figure is another discrepancy. 

Audouin and Milne Edwards (1834) give a vivid description of the movements of 
this species and its powers of burrowing in the sand. They allude also to its use as bait 
by the fishermen. 

De Quatrefages (1865) separated the K Hombergii of Audouin and Milne Edwards' 

1 MS. drawings by Miss Dorville, 1808, Library Linnean Society. 



NEPHTHYS CILIATA. 23 

from Cuvier's form, naming the latter N. Cuvieri, but it is doubtful from the description 
of the author or of Cuvier whether any distinction exists. 

After describing the species, Ehlers (1868) furnishes a digest of the literature, 
pointing out the confusion into which it had lapsed. His view that the Nephthys 
longisetosa of Johnston was probably this species was in consonance with Malmgren's 
opinion and with the appearance of the examples in the British ~ Museum. He further 
thought that the Nephthys Cuvieri of De Quatrefages was the same or a closely allied 
form. 

Some authors place the Nephthys emarginaia of Malm (1874) under N cdsca, but a 
careful examination of his figure would lead to the view that it rather pertains to 
this species. 

Langerhans (1879) gives an interesting account of the connective-tissue apparatus 
(Band apparatus) of this species, with various figures. The central apparatus is 
internal to the nerve-cord, and branches proceed to both lobes of the foot. 

Ehlers and Theel (1879) consider that this species is indicated in (Ersted's ' Grronl. 
Dorsib.,' pp. 195-6, figs. 75 and 76, and the figure given by the author is certainly 
ambiguous — more resembling an imperfectly preserved example of N. Hombergii than 
N longisetosa, though the dorsal lamella is stated to be triangular. 

Tauber (1879) seems inclined to include the Nephthys Johnstoni, N. nudipes, and 
N. pansa of Ehlers under this form, and is doubtful as to the separation of N incisa, 
Malmgren. 

De St. Joseph (1894) gives a somewhat minute account of this species, but un- 
fortunately his figures are indifferent. Like Delle Chiaje, he mentions the occurrence 
of two small eyes at the posterior border of the first segment. He supposes the A 
longisetosa of Johnston to be the same, but this is doubtful, since the description may 
refer to the long bristled variety of Nephthys caeca. Besides the structure of the exterior 
he describes the circulatory, muscular, and nervous systems, the presence of striated 
muscular fibres in the median ventral bands being a feature of note. He found the 
sexual elements developed from November to April. 

A careful account of the nervous system of this species is given by Pruvot l in which 
he points out the single cerebral mass without inferior cerebral lobes or secondary ganglia. 
Each oesophageal connective, double at its origin, is single thereafter, and gives off the 
stomato-gastric nerves. The ventral ganglionic chain is norma], and it has one neural 
canal inferiorly (the others apparently having escaped his notice). 



3. Nephthys ciliata, 0. F. Mutter, 1789. Plate LXVI, fig. 9— foot ; PI. LXXVI1, figs. 6, 

6 a, and 7 — brist. 

Specific Characters— -Head longer than broad, with a wide anterior border and more 
slender tentacles than in N cseca. Proboscis in extrusion generally papillose, with 



i ( 



Arch. Zool. Exper/ 2 e ser, in, p. 225, pis. xi and xii, 1885. 



24 NEPHTHYS CILIATA. 

twenty-two distal rows of slender, elongated papilla? — five to seven in number in each. 
A median cirrus in front of the rows in the mid-dorsal line. Body 90 — 135 segments, of 
medium size and generally typical in outline. Foot with the dorsal lamella rounded, its 
greatest vertical diameter being median, and its axis directed upward. At its outer edge 
is a smaller rounded lamella. Dorsal cirrus slender and long. Branchial process of 
moderate length coiled outward. Inferior lobe sinuous below the small terminal lamella, 
with a small papilla at the inner border of the latter superiorly. Ventral cirrus large and 
conical. Bristles brownish, comparatively short, the serrated forms having a distinct 
curvature of the dilated region beyond the shaft, and a serrated edge. The barred forms 
are finely tapered. 



Synonyms. 

1789. Nereis ciliata, 0. F. Miiller. Zool. Danica, iii, p. 14, Tab. lxxxix, figs. 1 — 4. 

1791. „ „ Gmelnr's Linnaeus Syst. Nat., ed. 13, I, pt. 6, p. 3120. 

1806. „ „ Turton's Gmelin, iv, p. 90. 

1843. Nephthys longisetosa, CErsted. Gronl. Dorsib., Tab. vi, fig. 78. 

„ „ borealis, idem. Annul. Danic. Consp., p. 32, fig. 4. 

„ „ ciliata, H. Kathke. Beitr. Fauna Norweg., p. 170. 

1851. „ „ Grube. Fam. Annel., p. 53, 128. 

1853. „ „ (borealis GErsted), Stimpson. Mar. Invert, of G-r. Manan, p. 33. 

1865. „ borealis, De Quatrefages. Annel. i, p. 428. 

„ „ ciliata, idem. Ibid., p. 429. 

„ Diplobranchus ciliatus, idem. Ibid, p. 434. 

„ Nephthys ciliata, Malmgren. Nord. Hafs-Ann., p. 104, Tab. xii, fig. 17. 

1867. „ „ idem. Annel. Polych., p. 17. 

1868. „ „ Ehlers. Die Borst., ii, p. 629, Taf. 23, f. 36. 

1872. „ „ G. O. Sars. Bid. Kundskab Christ. Fauna, iii, p. 23. 

1873. „ „ Kupffer. Exped. Ostsee " Pommerania," p. 150. 

„ „ „ Yerrill. Ajner. Journ. Sc. and A., vi, p. 411, pi. iv, f. 4. 

„ „ „ Mobius. Jaliresb. Com. deutsch Meere, p. 113. 

,, „ ,, Kupffer. Ibid., p. 150. 

„ „ „ Yerrill. Invert. Yinyard Sound, U.S. Comm. F. and F., p. 583. 

1874. „ „ Smith and Harger. Tr. Conn. Acad., iii, p. 16, pi. v, f . 1. 

„ „ „ Malm. Kongl. Yet. o. Yitt. Samh. i Goteborg Handl., xiv, p. 76. 

„ „ „ Yerrill. Proc. Amer. Assoc, etc., for 1873, pp. 351, 367, pi. v, f. 7. 

1875. „ cseca, Mobius. Jahresb. Comm. w. Untersuch., etc., p. 166. 

1878. „ ciliata, Leuz. Jaliresb. Comm. w. Untersuch. deutsch, 1874 — 75, p. 13. 

1879. „ „ Yerrill. C. List, U.S. Comm. F. and F. (sep. cop.), p. 7. 
„ „ „ Mcintosh. Trans. Linn. Soc, 2 ser., vol. i, p. 501. 

„ „ „ Hansen. Nyt. Mag. f. Naturvid., xxiv (Annel. Norske Nordhav. Exped.), 

p. 268. 
„ „ „ Theel. Acad. Handl. Stockholme, Bd. xvi (Annel. JSTouv. Zemb.), p. 24. 

1881. „ „ Horst. Mederland. Archiv f. Zool., Suppl. Bd., 5, p. 7. 

1883. „ „ Levinsen. Yidensk. Meddel. Foren. Kjoben., p. 217 (Syst-geogr. Overs. 

Nord. Annel., p. 60). 



NEPHTHYS CILIATA. 25 

1886. Nephthys ciliata, idem. Ann. Kara-Hav. Led., p. 8. 

189 °- „ „ Malaquin. Eev. Biol, du Nord Fr., p. 32 (Annel. Boulon.). 

1^93. „ „ Levinsen. Vidensk. Ud. " Hauchs," p. 337. 

1898. „ „ Micliaelsen. Zool. Ergebn., ix, G-ronl. Annel., p. 126. 

1900. „ „ Fauvel. Annel. Cherbourg, Mem. Soc. nationale des Sc. nat., etc., Cherbourg, 

tome xxxi, p. 309 et seq. 
„ „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. v, p. 258. 

1901. „ „ idem. Ibid., vol. viii, p. 222. 

» >, „ Whiteaves. G-eol. Surv. Canada, No. 722, p. 82. 

1902. „ „ Marenzeller. Polych. des Grundes, Denkschr. d. K. Akad. d. Wiss., Wien, 

lxxiv Bd., sep. abdr., p. 11. 

1903. „ „ Moore. Polych. Japan, Proc. Acad. Nat. Sc. Philad., June, 1903, p. 433. 



Habitat — Bressay Sound, 10 fathoms; St. Magnus Bay, 90 — 100 fathoms; generally 
on muddy ground or in sandy mud. 

O. F. Miiller procured it in the first instance from the Faroe Islands, and it stretches 
to Greenland and the eastern Canadian waters as well as to America. 

Malmgren gives Spitsbergen, Scandinavia, and Iceland, and Ehlers the European 
and American shores of the North Atlantic (Verrill). Kara Sea, Nova Zembla 
(Theel). 

The most abundant Norwegian species collected by Canon Norman; Behring's Sea 
(Marenzeller) . 

Head longer than wide, with a broad and slightly convex anterior border and 
more slender tentacles than in K cseca, the anterior being lateral in position, the posterior 
ventrolateral. 

The peristomial segment shows the dorsal and ventral spine-papillae and the two kinds 
of bristles in each. 

Proboscis. — In extrusion the terminal papillae are more slender and elongated than in 
2V. cseca or N. Hombergii, and the body of the organ therefore is generally more papillose. 
There are about five papillae in each row, the anterior being longer and more slender 
than in the common species. Moreover, shorter papillae are dotted over the organ almost 
to the base. In the median line dorsally a long slender anteriorly directed cirrus occurs 
a little in front of the other rows. 

Body slightly tapered in front, more so posteriorly, and terminating in a caudal 
cirrus. 

The typical foot (Plate LXVI, fig. 9) is distinguished from that of N. cseca by the fact 
that the superior lamella is smaller and more rounded, its greatest vertical diameter being 
towards the middle, and the axis directed dorsally (upward), whereas the greatest vertical 
axis in N. cseca is directed obliquely outward. The slight development of the lamellae, 
indeed, at once differentiates the species. The dorsal lamella is continued to the tip of the 
foot where the edge bends downward, and its tissue becomes continuous with that of the 
dorsal cirrus and branchia. The dorsal cirrus at the base of the branchial process is 
rather slender and long. The branchial process is of average length, though in some it is 
short. Immediately in front is the spinigerous region of the foot which has the peculiarity 

61 



26 XEPHTHYS CILIATA. 

of forming a free lamella guarding the capillary bristles in front, so that they lie in a 
fissure between it and the posterior lamella. This spinigerous flap has the spine a little 
beyond its middle, and thereafter it ends in a rounded or disc-like free lamella at the edge 
of the foot. The anterior base of the foregoing has a slight fillet from which the barred 
or camerated bristles project. The bristles have a distinct curvature (Plate LXXVII, 
fig. 6) along the dilation above the shaft, and this is spinose or serrated. The barred 
forms (Plate LXXVII, fig. 7) are finely tapered, and perhaps less robust than those of 
N. caeca. 

The inferior division has a short posterior lamella which superiorly in the large 
examples rises as a free rounded lobe. The spinigerous region forms another lamella 
anteriorly from the upper border to the lower, an interval, however, occurring between 
it and the cirrus. The capillary bristles arise between it and the posterior lamella, and 
thus lie in a deep pit. The barred bristles spring from a line marked by a ridge of skin, 
which superiorly ends in a distinct flap or lamella at the commencement of the spinigerous 
region. 

The conical ventral cirrus is comparatively large. The ventral bristles agree with 
the superior in having a well-marked series of spikes on the curved and dilated portion 
above the shaft. On the whole N. ciliata has comparatively short and slender 
bristles. 

The figures of 0. F. Miiller in the ' Zoologia Danica ' (1789) are recognizable not so 
much in minute detail, as in the firm outline and general contour of figs. 1 and 2. The 
description of the plate, moreover, bears out this interpretation, though the author adheres 
to there being two caudal cirri, but as only traces of these were left in the specimens which 
Mohr had sent him from Faroe, the mistake is explained. He correctly discriminated it 
from the N. caeca of 0. Fabricius. He distinguished it from the gigantea (Seba Thesaur, I, 
81, fig. 7) of Linnaeus by the third series of bristles, whilst from Nereis caeca (Fabricius, 
'Fauna Groenlandica ') it differs by the absence of "scales" between the feet, Midler's 
form having only fleshy fimbriae. 

H. Eathke (1843) described this as a new species, but in all probability it is the 
same form. He gives no figures. 

(Ersted (1843) at first confused the species with N. longisetosa, but soon corrected 
himself, and gave an excellent figure of a foot in his ' Conspectus.' 

The formation of a new genus by De Quatrefages (1865) for this species rested on 
misapprehensions, somewhat increased by his entering Rathke's Nephthys ciliata and 
(Ersted's N. borealis as separate forms. 

Dr. Johnston did not separate the species from N. cseca. 

Malmgren's account and figures, with the supplementary note in the 'Annulata 
Polychgeta,' readily define the species from allied forms. 

Ehlers (1868) corrected the synonymy up to date, and gave a reliable description 
without figures. 

Smith and Harger's figure (1874) might do for either Nejphthys ciliata or incisa, 
though it more nearly approaches the latter. These and other Annelids described in their 
paper were dredged on St. George's Bank. 



NEPHTHYS HYSTRICIS. 27 

Dr. H. Theel (1879) criticises under this species Malmgren's figures of the feet, 
since he did not accurately compare the same feet in each species, or at least from the 
same region of the body. 

Tauber (1879) includes Nephthys cirrosa, Ehlers, and probably N. lactea, Malmgren, 
under this species. 

Shepotieff describes and figures the bristles and bristle-sacs with their muscles x — his 
careful investigations being made chiefly on sections of the parts. 



4. Nephthys hystjbtois, Mcintosh, 1900. Plate LVII, figs. 8, 9 ; Plate LXVI, figs. 10 and 

10 a — feet. 

Specific characters. — Head elongated from before backward, with rather pointed 
subulate tentacles anteriorly, the broader second pair following after an interval. 
Peristomial segment ventrally with two broad flaps and a symmetrical series of furrows. 
Body resembling that of N. ciliata. Proboscis comparatively short, with slender papillae 
in twenty-two rows. The mid-dorsal pair converge on the long tentacle immediately in 
front, and the rows on each side of the mid-ventral line converge in a more marked 
manner, but there is no median filament. As a rule, four papillaB occur in each row, but 
in some traces of a fifth are seen, and in the ventral pair (on each side of the median 
line) are six short papillaB. Foot with a long, little elevated, dorsal lamella, which does 
not extend so far outward or droop at the tip, as in N. Hombergii. It also approaches in 
form that of N. ciliata, but differs in the shape of the dorsal lamella, and in the fact that 
in front of the capillary bristles the spinigerous region forms a low cone with the spine at 
the apex, the barred bristles arising in the fissure between it and a large flap or lamella 
formed by the fillet guarding the bristles, whereas in N. ciliata the spinigerous region 
itself forms the free flap. The dorsal cirrus is somewhat thick, separated only bv a 
shallow notch from the branchia, which is of moderate size, has a papilla at its outer 
base, and is curved outward. Bristles similar to those of N. Hombergii, but thinner. 
Inferior division of the foot with a small lamella posteriorly, another nearly as prominent 
formed by the fillet for the barred bristles, the conical spinigerous lobe lying between. 
In several views, therefore, the tip is trifid. 



Synonyms. 

1900. Nephthys hystricis, Mcintosh. Ann. Nat. Hist., ser. 7, vol. v, p. 259. 

1901. „ „ idem. Ibid., vol. viii, p. 222. 

1902. „ „ Marenzeller. Polych. Grand., p. 14. 

Habitat. — Dredged in the 'Porcupine' Expedition of 1869, 110 fathoms; 370 
1 'Zeitsch. f. wiss. Zool./ Bd. lxxvii, p. 593, Taf. xxvii, f. 5—17, 1904. 



28 NEPHTHYS HYSTBICIS. 

fathoms ; 422 fathoms ; off Ireland. From various stations in the Mediterranean 
during the 'Porcupine' Expedition of 1870; 9 miles off Cape Finisterre; 81 fathoms 
east of Cape de Gatte, 6 miles from shore, 60 — 160 fathoms; Stations 6 and 8, 
1870. Dredged in the ' Porcupine ' Expedition of 1870, Bono Bay, 25 fathoms ; No. 36, 
128 fathoms, 1870; No. 2, 305 fathoms, and off Cape Sagres, 45 fathoms; Sidi 
Ferrara? 45 fathoms. 'Knight Errant,' August 24th, 1882, Station 18, 516 fathoms. 
Boyal Irish Academy's Expedition, 1885, Berehaven. A closely allied if not identical 
form was obtained by Canon Norman in Lervig Bay, Norway, in 1878, in considerable 
numbers. 

Head (Plate LVII, fig. 8) elongated from before backward, with rather pointed 
anterior tentacles, the second pair being a little behind and ventro -lateral in position. 
Posteriorly the head is bounded by the collar with a median notch, and a four rayed 
mark is situated on the head a little in front. 

The peristomial segment shows two broad lips anteriorly and a symmetrical series 
of furrows posteriorly. The ventral cirrus anteriorly is small, as in A. ciliata. 

In all the Irish specimens the proboscis is ejected, so that the head is distorted 
(Plate LVII, fig. 8). Its transverse diameter is greater than the antero-posterior. 
The anterior edge is straight with a small subulate tentacle at each side. A broader 
ventral one follows after a short interval. 

The body is incomplete in all the examples, but it resembles that of N. ciliata. 

The proboscis (Plate LVII, figs. 8 and 9) is comparatively short with slender bifid 
papillae in twenty-two rows. The median dorsal pair converge on the long tentacle 
immediately in front. The rows on each side of the ventral median line converge even 
in a more marked manner, but there is no long median filament. The papillae are 
longest distally and diminish backward. Four are distinct and in some is a trace of a 
fifth, though in the pair on each side of the ventral line there are more (at least six), 
but they are short. Under the lens the rest of the basal region is smooth. The acute 
bifid papillae which form a border to the aperture at the tip are very distinct, either 
projecting prominently or folded over the aperture, the tips crossing in a regular and 
symmetrical manner. The long dorsal tentacle projects as far as the latter (/. e. to the 
closed aperture). In some the distal region is pale madder brown. 

The typical foot (Plate LXVI, fig. 10) approaches that of certain varieties of 
A. Hombergii, such as var. Jcersivalensis, insofar as it has a long, little elevated lobe, 
which, however, does not pass so far outward or droop so much at the outer end as 
in - the form mentioned. It stands indeed horizontally. The aspect of the foot also 
approaches that of A. ciliata, but it differs in minute characters, such as in the form 
of the dorsal lamella, the condition of the spinigerous region, and other particulars. In 
front of the capillary bristles the spinigerous region forms a low cone or pyramid of 
considerable thickness with the spine at the apex, the barred bristles arising in the 
fissure between it and a large flap or lamella formed by the fillet guarding the bristles 
(Plate LXVI, fig. 10 a). There is thus an essential difference between it and A. ciliata, 
in which the spinigerous region itself forms the free flap. The dorsal cirrus is somewhat 
thick, and is only separated by a shallow notch from the branchia, which is large and 



NEPHTHYS LONGISETOSA. 29 

curved outward. The bristles are similar in curvature to those of N. Hombergii, but 
are more translucent (thinner), and the serrated edge is less rigid, and often disappears, 
apparently by friction. The barred bristles offer no feature of note. The inferior 
division has a small lamella posteriorly, another nearly as prominent formed by the 
fillet for the barred bristles, while the conical spinigerous lobe lies between. In many 
views, therefore, the tip is trifid, and quite diverges from that of JSf. ciliata. 

The foot of the Mediterranean examples agrees with that in the Irish forms, the 
only difference being the greater distinctness of the spinigerous papilla between the 
lamellge in the upper division, but as the Irish specimens are badly prepared and 
preserved this may be more apparent than real. The bristles of those from the 
6 Porcupine ' are certainly longer. In both their colour is dull yellow, and their structure 
seems to agree. 

The branchial process, which is of moderate size, has a well-marked papilla or 
process at its outer base ; as shown for instance in Malmgren's figure of N. assimilis, 
(Erst. 1 It is, however, more distinctly defined than the latter. 

This form approaches N. ciliata in the structure of the foot, yet differs in the longer 
dorsal lamella (which thus extends further outward), in the form of the dorsal cirrus, and 
in the basal process of the branchia. The terminal lamella of the foot is different. 
The median dorsal cirrus of the proboscis also seems to be longer. It is no nearer 
N. cirrosa, Ehlers, from which it differs in the great width of the dorsal lamella. 

A Norwegian example collected by Canon Norman shows a more distinctly bilobed 
fillet in front of the camerated bristles superiorly. 

A variety procured in the ' Porcupine ' east of Cape de Gatte, nine miles from shore, 
in 6 to 160 fathoms, had more slender bristles, which apparently had undergone some 
change and had. lost the serrations on the edge. 



5. Nephthys longisetosa, (Ersted, 1842. Plate LVII, fig. 10-12; Plate LXVI, fig. 11- 
foot; Plate LXXVII, figs. S-8b— brist. 

Specific Characters. — Head somewhat shield-shaped ; the anterior tentacles marked 
by a translucent stripe at the base in front, long and slender ; second pair also long and 
lanceolate. Proboscis distinguished by fifteen (Theel says fourteen) rows of numerous 
papillse, viz. eleven to fifteen in each row. Dorsal lamella of the foot elevated and 
prominent (triangular, (Ersted), with an accessory rounded process. Dorsal cirrus large 
and lanceolate, closely connected with the base of the branchia, only a shallow notch 
intervening, and the branchia is often curved inward. Ventral lobe with a tongue- 
shaped lamella above the dorsal edge at the tip, and only a narrow lamella beyond the 
foot. Both barred and serrated bristles are yellow or brownish with a metallic sheen, 
and largely developed, the latter presenting a gentle curve from base to apex, the 
minute spines being traceable from the short shaft to the tip. 

1 Op. cit., Taf. xii, fig. 19 6. 



30 NEPHTHYS LONGISETOSA. 



Synonyms. 



1842. Nephthys longisetosa, CErsted. Kroyers Nat. Tids., iv, 2, p. 123. 

18 43. „ j} idem. Gronl. Dorsibr., p. 195, figs. 75 and 76. 

1851 • „ „ Sars. Nyt Mag. vi, p. 208 (Reise i Lofoten og Finmarken), 1849. 

1865. „ longosetosa, De Quatrefages. Annel. i, p. 428. 

» )) „ Malmgren. Nord. Hafs-Annul., p. 106, Taf. xii, fig. 20. 

1866-69. „ longisetosa, Packard. Mem. Bost. Soc. N. Hist., i, p. 293. 

1867. „ }J Malmgren. Annel. Polychget., p. 19. 

» » „ Parfitt. Cat. Annel. in Tr. Dev. Assoc, ii, pt. 1, p. 231. 

» ft longosetosa (Stimpson) Packard. Mem. Bost. Soc. N. H., p. 294. (?) 

1871. „ ,, Elilers. Annel. from Spitzbergen, Sitzungsb. Phys.-Med. Soc. 

Erlangen, iii, p. 79. 

1878. „ i} Marenzeller. Nordpol. Exped., Denkschr. Mat. -Nat. class d. Kaiserl. 

Akad. Wien, p. 395. 

1879. „ „ Yerrill. C. List U. S. Comm. Kept., p. 7. 

» 99 99 Hansen. Nyt Mag. f. Naturvid. (sep. copy), xxiv, p. 268. 

« 99 Malmgreni, Theel. Annel. Nouv. Zemb., p. 26, pi. i, f. 17 5 , and pi. ii, f. 17 10 . 

1881. „ „ Horst. Niederl. Arch. Zool., Suppl. Bd. i, 5, p. ]0. 

1883. „ 9i Levinsen. Syst. G-eogr. Nord. Annel (Vidensk, etc.), p. 216. 

1896. „ longisetosa, Michaelsen. Polych. deutscli. meere, p. 24. 

1 9 °0- „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. v, p. 260. 

1901- „ „ Whiteaves. G-eol. Surv. Canada, No. 722, p. 82. 

Habitat— St. Magnus Bay, Shetland, 100 fathoms, and in 70 fathoms, Outer Haaf, 
Skerries (J. G. J.). Station 8 in the Faroe Channel ('Knight Errant'). Station 10 in 
the Faroe Channel ('Knight Errant'), March 2nd, 516 fathoms; August 24th, 82 
fathoms. 'Porcupine,' 1870, 447 fathoms, No. 31; Sidi Ferrara?, 45 fathoms; 374 
fathoms, No. 25. 

(Ersted's specimen came from Godthaab. Extends to the Atlantic shores of America 
(Verrill), Strait of Magellan (Ehlers). Canada (W. CM.). Fragments were procured off 
Anticosti in 100—210 fathoms, 1871, and off Port Hood, Canada, 1873, by Dr. Whiteaves. 

The head (Plate LVII, fig. 10) is somewhat shield-shaped, with a slightly convex 
anterior margin. The anterior tentacles are marked off by a translucent stripe in front ; 
and thus appear to rise considerably within the margin. They are comparatively long 
and slender. The second pair are lanceolate and also long. The dorsal cirrus of the 
first foot is remarkably large, its outline being lanceolate. 

The body is slightly narrowed in front, more so posteriorly, and distinguished by the 
strong curved bristles which stand somewhat stiffly out on each side. Ventrally again 
the enlarged lanceolate lamella formed by the ventral cirrus is characteristic. 

The proboscis in extrusion (Plate LVII, fig. 10) is distinguished by the well-marked 
basal region; and the very numerous papilla in its distal rows, which are fifteen in 
number. Malmgren states that there are but fourteen. Moreover, the latter author 
also limits the number of the papillaB in these rows to seven, whereas they are much 
more numerous in the present examples, viz. eleven to fifteen. 

When partly withdrawn, the organ in situ (Plate LVII, fig. 11) presents a button- 



XEPHTHYS LONGISETOSA. 31 

shaped tip of ruptured radiate muscular bands which are fixed to the next region. On 
each side a strong series of fibres forms a triangular muscular sheet, narrow posteriorly, 
which is inserted into the hollow of the constriction about a third from the front. 
Thereafter the organ gently dilates, again to contract posteriorly where it joins the gut. 
A slight ridge, commencing behind the pit at the muscular belt, marks the dorsal median 
line, and a slight furrow the corresponding part of the ventral surface. This region has 
fine transverse lines as in the proboscis of the Aphroditidse. 

When the organ is completely retracted, the protractor muscles form long bands 
(Plate LVII, fig. 12) from the snout backward to the dense proboscis, the special triangular 
bands at each side, again, acting on the latter in eversion, so as to tilt it outward. 

The typical foot is not much produced, and shows dorsally an elevated and conspicuous 
lamella of a semicircular or truncated broad ovoid form (Plate LXVI, fig. 11) with a 
continuation appearing as a small rounded process or flap while the interfascicular lobe 
slants from above downward and outward with a prominent papilla for the spine, after 
which the margin recedes. A small free flap occurs at the outer margin of the fillet 
guarding the base of the barred bristles. The base of the branchia is closely connected 
with the dorsal cirrus, only a shallow notch separating them. The cirrus is lanceolate, 
broad and flattened at the base, pointed at the tip. It is interesting that the same 
structure in A T . trissophythus, Grube, from Naples, is largely developed into a thin lamella 
like the cordate leaf of the lime. The branchia is often coiled inward. The barred 
bristles offer no diagnostic feature except their strength (Plate LXXVII, fig. 8a). The 
long serrated bristles are remarkably strong, with a pale yellow metallic sheen, and 
curve backward. The flattened blade gently bends from the shaft, its convex edge being 
minutely serrated with rows of spikes (Plate LXXVII, fig. 8 and 8b). 

The inferior division presents two flaps superiorly, the upper edge of the bristle-fan 
passing between them. The anterior lamella is the larger, is tongue-shaped, and has a 
further modification of the small process observed in other forms, e.g., K ciliata, 0. F. M. 
The lower angle of the same fillet often has a small free flap. The spinigerous region forms 
an acute triangle with the spine at the apex. The inferior lamella is comparatively small 
and somewhat resembles an ovate leaf, but there are considerable differences in regard to 
the development of this lamella, the Zetlandic form especially having it well-marked with 
the tip pointing upward and outward. The ventral cirrus is large and lanceolate with a 
central rib, and an articulation at its base. Both barred and serrated bristles are largely 
developed, the latter presenting a gentle curve from base to apex, the minute spines being 
traceable from the short shaft to the tip. 

In contrast with K caeca, the inferior oar of the foot in N. longisetosa is less 
prominent, and does not project so far outward. The bristles of the entire foot are also 
much more strongly marked (not longer), and have a decided curve backward. In 
K longisetosa those with the serrations at the side are broader at the dilated portions. 
The inferior lobe is little developed, and thus strongly contrasts with several of its allies. 
The inferior cirrus is lamellate, and broader than in most. The serrations on the edges of 
the bristles are less conspicuous than usual; or they resemble the slated markings of 
wool rather than sharp and independent serrations. This gives a character to the bristles. 
The process at the base of the sickle-shaped cirrus is rather long and lanceolate. 



32 NEPHTHYS LONGISETOSA. 

The head is less rounded in front than in A r . cseca, and the anterior tentacles more 
prominent and larger. 

In what appears to be the northern variety of this species from Finmark the dorsal 
lamella is smaller, the accessory lobe little developed, the dorsal cirrus shorter and 
broader, the branchia much larger, but still rolled inward, the ventral lamella much less 
and the cirrus smaller and less foliaceous. The bristles in this form are more slender 
than in the British, but the serrations are distinctly separated. The barred forms are 
also more slender. While the lamella both dorsally and ventrally are thus smaller, the 
setigerous region in both divisions is more prominent and acute. 

A form dredged by Canon Norman near Bergen in 1878 differs from the foregoing 
in the much darker bristles, which have, however, the same backward curve. The body 
somewhat resembles that of K cseca, though the specimens are smaller. The head shows 
in one of the preparations two opaque specks posteriorly, but they do not seem to be eyes. 
The proboscis is enclosed. The foot presents a considerable dorsal lamella of a rounded 
form and extending to the tip. In front of it are the dark capillary bristles, then a much 
less spinigerous region than in N. longisetosa so that the barred bristles are quite close to 
the others. What is, however, the most marked feature is the presence of a small rounded 
flap superiorly and inferiorly in front of all the bristles, and this is apparently a develop- 
ment of the fillet in front of the barred bristles. The dorsal cirrus is small instead of 
large, and separated by a slight notch from the branchia, which is large, though not long, 
and curved outward. The inferior division has a somewhat smaller posterior lamella 
than K longisetosa, and the cirrus is much less — forming a simple conical process. Both 
kinds of bristles appear to be more slender than in N. longisetosa, but the larger size of 
the latter probably makes the difference more pronounced. 

The foot thus differs in the shape of the dorsal lamella, which is somewhat higher 
than in the example from Finmark, and in the pointed nature of the superior lobe. The 
dorsal cirrus and branchia are similar. The ventral lobe is also acutely pointed. The 
ventral cirrus is acute, and the long curved bristles have remarkably distinct serrations. 
The specimens are small and imperfect. 

Nephthys longibranchiata, of the ' Porcupine ' Expedition, differs from the allied form, 
N. longisetosa, (Erst., by the broader and less elevated superior lamella, by the shorter and 
more obtuse process at the base of the branchia, by the greater size and length of the 
branchia, by the rounded tip to the inferior lobe, the projection of the spine-tip, and the 
much smaller inferior cirrus. 

Malmgren (1865) considered that GErsted had this species in view in his ' Gronlands 
Annulata Dorsibranchiata,' but the figures are so unsatisfactory that doubt must always 
remain. He certainly describes the dorsal lamella as triangular. Malmgren also (1867) 
pointed out that the specimens so labelled in the British Museum by Dr. Johnston, from 
Berwick Bay, Holy Island, and the Firth of Forth, pertained to Nephthys Hombergii 
and a new species. 

Such forms as Nephthys phyllocirra, Ehlers, 1 make a close approach to this species. 

1 c Florida Annel./ p. 131, Taf. 38, fig. 7—11, 1887. 



NEPHTHYS GEUBEL 33 

6. Nephthys Gettbei, 1 Mcintosh, 1900. Plate LVII, figs. 13 and 14; Plate LXVII, 
fig. 1— feet; Plate LXXVI, figs. 9 and 9 a— bristles. 

Specific Characters. — Head elongated from before backward, with a straight anterior 
edge, which is somewhat narrower than in JV. longisetosa ; tentacles at the outer border, 
conical, tapering, with a translucent area at the base internally. Second pair follow close 
on the first, and have the form of an acuminate leaf, whereas in JV. longisetosa they are 
further removed from the first, as well as proportionally longer and more pointed. 
Distinct papilla, probably sensory (nuchal organ), at each posterior angle of the head. 
Proboscis included, but the arrangement of the parts around the mouth is similar to that 
of JV. longisetosa. First foot diverges from that mentioned in the presence of a subulate 
dorsal process or cirrus, besides the lanceolate ventral one. The typical foot has the 
dorsal lamella narrowed at its attachment, broader and somewhat truncated distally, the 
reverse occurring in JV. longisetosa. Instead of a distinct flap externally there is only a 
trace in the shape of a fillet. Dorsal cirrus less, branchia longer than in that species and 
curved inward. Pointed spinigerous lobe produced further outward, and at a different 
angle: fillet guarding bristles convex upward. In the inferior division the posterior 
lamella smaller, spinigerous lobe longer and more acute, whilst anteriorly the upper 
fillet is smaller. Ventral cirrus smaller and less tapered. Both barred and capillary 
bristles are more slender than in JV. longisetosa, though in both the transverse bars of 
the serrated bristles extend downward till the full diameter of the shaft is reached. 



Synonym. 
1900. Nejohthys Grubei, Mcintosh. Ann. Nat. Hist., ser. 7, vol. v, p. 260. 

Habitat.— Dredged at Station 8 by the ' Knight Errant,' 17th August, 1880, in 
540 fathoms. 

The head (Plate LVII, figs. 13 and 14) is elongated from before backward, with a 
straight anterior edge which is somewhat narrower than in JV. longisetosa. The tentacles 
at the outer angle are conical, tapering, and have a translucent area at the inner border. 
The position of the second pair differs from that in JV. longisetosa, for they follow 
inferiorly close on the anterior pair, and have the form of an ovate acuminate leaf; 
whereas in JV. longisetosa they are separated by a considerable space from the anterior 
pair, and they are proportionally longer and more pointed. At each angle of the head 
posteriorly is a conspicuous papilla, probably a sense-organ (nuchal organ), the corre- 
sponding organ in JV. longisetosa being inconspicuous. The lips inferiorly (Plate LVII, 
fig. 14) somewhat resemble those of N. Hombergii, but have a more acute filament. 

The proboscis is included. 

The first foot differs from that in iV. longisetosa, for it has a well-marked subulate 

1 Named after Professor Adolpli Edouard Grube, of Breslau, than whom no more earnest 
student of the Annelids existed in his day. 

62 



34 JNEPHTHYS JOHNSTONI. 

dorsal process or cirrus, besides the lanceolate ventral one ; whereas in N. longisetosa 
only a small papilla exists dorsally, and the ventral cirrus is much longer. 

In the typical foot (Plate LXVII, fig. 1) the dorsal lamella is narrowed at its 
attachment, broader and somewhat truncated distally, the reverse being the case in 
N. longisetosa. Instead of a distinct flap externally, only a trace in the shape of a fillet 
occurs : the dorsal cirrus is evidently less, while the branchia is larger, though it is 
likewise curved inward. The spinigerous lobe is produced much further outward as a 
pointed process, and has a different angle. Moreover, the fillet guarding the barred 
bristles has a convexity directed upward. 

In the inferior division the posterior lamella is considerably smaller, the spinigerous 
lobe longer and more acute, whilst anteriorly the upper fillet is somewhat smaller. The 
ventral cirrus is lanceolate, and shorter and less tapered than in N. longisetosa. 

Both capillary and barred bristles (Plate LXXVI, figs. 9 and 9 a) are more slender 
than in JSf. longisetosa, and in both the transverse bars of the serrated bristles extend 
downward till the full diameter of the shaft is reached. 



7. Nephthys Johnstoni, Elders, 1875. Plate LVII, figs. 15 and 16; Plate LXVII, 
figs. 2 and 2 a— feet ; Plate LXXVI, figs. 10 and 11— bristles. 

Specific Characters. — Head shield-shaped, with a broad and somewhat convex anterior 
border ; a slender tapering tentacle at each angle, and an oblique translucent patch at its 
inner base. The second pair of tentacles arises latero-ventrally after a considerable 
interval. Proboscis with rather long rows (twenty-two) of papillae distally, and a median 
cirrus dorsally and ventrally. Between the rows of the distal bifid papillae of the 
aperture is a simple papilla dorsally and ventrally. The anterior folds or lips in front 
of the mouth are large. The dorsal lamella of the foot has a similar curvature to that 
of N~. ciliata, but it extends further outward, and only a trace of the rounded terminal 
lobe appears. The setigerous region stands prominently upward and is smoothly 
rounded externally. No free flap to the anterior fillet guards the barred bristles. 
The dorsal cirrus is a variable filiform process. The branchia is not large and is curved 
externally. The ventral lamella is large and ovato-lanceolate, and the ventral cirrus 
lanceolate, larger than in N. ciliata, though less than in N. longisetosa. The capillary 
bristles are longer and less rigid than in N. longisetosa, and the serrations very fine, 
the long distal region being smooth. The barred region in the anterior bristles is 
long, and slightly fusiform. 

Synonyms. 

1843. Nephthys longisetosa, CErsted. Gronl. Annul. Dorsibr., p. 195, figs. 75 and 76. 
1865. „ „ Johnston. Cat. Brit. Mus., p. 172. (?) 

1874. „ Johnstoni, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 196. 

1875. „ „ idem. Invert, and Fishes, St. Andrews, p. 119. 

Ehlers. Zeitsch. f . w. Zool., Bd. xxv, p. 38, Taf. hi, f. 3 and 4. 
1900. „ „ Mcintosh. Ann. Nat. Hist., ser 7, vol. v, p. 261. 



XEPHTHYS JOHNSTONI. 35 

Habitat. — Southport Sands (Dr. Carrington) ; West Sands, St. Andrews, after 
storms (E. M.) ; St. Magnus Bay and Balta, Shetland (J. Gr. J.); 'Porcupine,' 1869, 
85 fathoms, on " Porcupine Bank," off Ireland. 

Greenland (CErsted). 

The head (Plate LVII, fig. 15) has a broad and slightly convex anterior border 
with the slender and tapered tentacle at each angle. A feature, apparently not 
unfrequent, is the presence of a translucent, oblique, elongated patch at the inner base 
of this tentacle. The second tentacle is broader, with a pointed tip, and occurs after 
a considerable interval. Posteriorly the elongated head terminates in two flattened 
lobes with a slight median furrow. A sense-papilla occurs at each side. The lips 
(Plate LVII, fig. 16) inferiorly approach the type of A 7 . Hombergii, but the filament is 
longer, and the arrangement of the furrows is different. The latter, however, is not 
of much value. 

The body has the usual shape and terminates posteriorly in the somewhat short 
anal cirrus (Ehlers). 

The proboscis has rows of rather long papillae distally, about six in each row, and 
besides, a long median cirrus both dorsally and ventrally just beyond the rows. There 
are y£ bifid papillae guarding the aperture in the distal region, and the gap between 
them has a single simple papilla. 

The first bristled foot has its dorsal fascicle directed upward and the tips turned 
backward, the whole forming a prominent fan. No dorsal cirrus is present, but the 
ventral is long, lanceolate, and pointed. 

The typical foot (Plate LXVII, figs. 2 and 2 a) has considerably longer bristles 
than in N. ciliata. The dorsal lamella has a similar curvature to that in N. ciliata, but 
has, in addition, a small and inconspicuous accessory lobe beyond its diminished outer 
end — continuous with the base of the cirrus. The spinigerous region forms a prominent 
ridge and ends externally in a smoothly rounded border without the distinct lobe as 
in N. ciliata. The fillet at the base of the barred bristles is distinct but does not rise 
into a flap. The dorsal cirrus is a filiform process, though this does not seem to be an 
important character, since, as in the dorsal lamella, considerable variety exists in the 
same specimen. The branchia is not large, and is curved externally. 

The lower division has a large posterior lamella, often sinuous inferiorly and of a 
broadly ovato-lanceolate form. The setigerous flap is oblique externally, and the fold 
running from the base of the barred bristles presents a flap superiorly. 

The capillary bristles both dorsally and ventrally are comparatively long but have 
very minute serrations (Plate LXXVI, fig. 10), a feature when contrasted with such as 
occur in N. longisetosa. The serrations, moreover, are limited only to the lower region 
of the bristle, the long tip being bare. In the camerated 1 bristles (Plate LXXVI, 
fig. 11) the transverse bars proceed far downward, the widest region of the bristle being 
thus included in the camerated part. When contrasted with those of N. longisetosa, 
they are considerably shorter and smaller. The ventral cirrus is lanceolate, larger than 
in N. ciliata, but considerably less than in N. longisetosa,. 

1 So termed from the peculiar arrangement of the rows of spikes. 



36 NEPHTHYS CIRROSA. 

Loxosomas occur on the feet of those from St. Magnus Bay, Shetland. 

It is possible, as Ehlers thought, that Dr. Johnston's K longisetosa refers to this 
species. He considered that it approached K Hombergii (Aud. and Ed.), though 
differing from it, and he courteously named it after the British author. 

(Ersted's Nephthys longisetosa (1843) makes so close an approach to this species that 
it may be included under it. 



8. Nephthys cierosa, Ehlers, 1868. Plate LVII, figs. 17 and 18; Plate LXVII, fig. 3 
—foot; Plate LXXVI, figs. 12 and 13— bristles. 

Specific Characters. — Head longer than broad, seven-sided, with a pair of conical 
tentacles in front, and a longer pair, latero-ventrally situated, considerably behind, A 
small (nuchal) papilla on each side posteriorly. The translucent area to the inner side of 
the anterior tentacles is distinct in some examples. Mouth a triradiate slit, for a fissure 
runs forward between the anterior flaps. Proboscis of the normal shape, with twenty- 
two rows of rather long papilla anteriorly in extrusion — each row having six or more 
papillae, which diminish in length proximally. Shorter papillee occur over the general 
surface of the organ. A single dorsal median cirrus is present, as in N. ciliata, only it 
is slender. The terminal arches of bifid papilla? are normal. The first foot has two 
bristle-tufts and a comparatively large ventral cirrus. Eoot with an arched dorsal 
lamella of considerable breadth, highest in the middle. No other flap. Dorsal cirrus 
of average length, subulate and rather slender. Branchia of moderate size, slightly 
coiled outward. The ventral division has a posterior lamella of moderate size, directed 
upward, and with a small flap superiorly over the base of the bristles. The ventral 
cirrus is prominent and somewhat conical. Bristles rather long and curved, pale golden; 
longer forms show a marked curve of the blade and a finely serrated edge. Barred 
forms with a spindle-shaped "camerated" region. Moreover, a little modification of 
the parts of the foot brings it close to N. hystricis, which, however, may readily be 
distinguished by the longer bristles, the larger spinigerous lobe, the smaller lamella? 
superiorly and inferiorly, and the free flaps from the fillet guarding the barred bristles 
in both divisions of the foot. 



Synonyms. 

1868. Nephthys cirrosa, Ehlers. Die Borst. ii, p. 624, Taf . xxiii, figs. 6, 38, and 39. 
1883. „ cseca, When. Clmetop. 'Vega' Exped., p. 392. 

1890. „ „ Malaquin. Annel. Polycli. Boul., p. 33. 

1894. „ „ De Saint-Joseph. Annel. Pol. Dinard, Ann. Sc. Nat., 8th ser., xyii, p. 20, pi 

fig. 19. 

1900. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. v, p. 261. 

1901. „ „ idem. Ibid, vol. viii, p. 222. 



1904. 



Allen. Journ M.B.A., n.s., vol. vii, p. 225. 



NEPHTHYS CIRROSA. 37 

Habitat. — The Channel Islands, Herm and Guernsey. In sand under stones at 
Galway, Ireland. 

Shores of France, Dinard and Croisic (Baron de St. Joseph) ; Norway (Canon 
Norman) ; Strait of Magellan (Ehlers). 

The head (Plate LVII, fig. 17) is longer than broad, somewhat translucent, and 
seven-sided. The anterior border is curved between the conical tentacles, which usually 
have a small translucent speck at their inner base. A longer pair occurs behind 
after a considerable interval, and latero-ventral in position. 

The mouth has a triradiate outline, for between the two triangular flaps in front is 
a fissure which runs into the transverse furrow behind. Posteriorly a striated elevation 
occurs, which tapers off about the fourth foot. The proboscis (Plate LVII, figs. 17 and 18) 
has twenty-two rows of rather long papillae, each row having six or more papillae. 
Shorter papillae are distributed over the general surface. A single dorsal median cirrus 
is present, as in A. ciliata, but it is more slender. The terminal arches of bifid papillae 
are normal. 

The typical foot (Plate LXVII, fig. 3) has a rather long low lamella, which extends 
outward beyond the tip, its highest point being about the centre, and thus differing from 
that of A. Hombergii. No other flap is present. The dorsal cirrus is of average length 
and subulate. The branchia is of moderate size and curved outward. The spinigerous 
lobe is narrow and flattened. The fillet guarding the base of the barred bristles projects 
as a small ridge externally. 

The inferior division posteriorly has a terminal lamella of moderate size directed 
upward. It differs in shape, and is less developed than that of A". Hombergii. A small 
flap, a process of the fillet guarding the base of the bristles, occurs superiorly over the 
base of the upper bristles. The spinigerous lobe is bluntly conical and of some thickness. 
The ventral cirrus is of considerable size. 

The capillary bristles (Plate LXXVI, figs. 12 and 12 a) have a rounded shaft 
and a long flattened curved blade tapering to a fine point, the edge being minutely 
serrated. Under the microscope many of these are yellow. They are comparatively 
long and pale golden to the naked eye, with a marked curve backward. The camerated 
bristles have a cylindrical shaft — dilating as it approaches the barred region, which 
forms a long spindle terminating in a tapering tip (Plate LXXVI, fig. 13). The 
crescentic fold between the feet ventrally probably indicates the opening of the 
segmental organ. 

This form comes near A. longisetosa. 

In an example from Herm, about four inches long, the superior lamella rose higher 
than usual, and had a different shape, since the outer edge was abrupt, and the inner edge 
also in some was more or less steep. The inferior lobe was less developed than in some 
examples of A. Johnstoni. A little further modification of the foot would bring this form 
within touch of A. ciliata. 

This species and A. Johnstoni come very near CErsted's A. longisetosa 1 from 
Greenland, the former, perhaps, most closely. 

1 ( G-ronl. Annul. Dorsibr./ p. 195, figs. 75, 76. 



38 NEPHTHYS INCISA. 

The species was first differentiated by Prof. Ehlers (1868), who had procured a 
small example from England through Mr. W. Stimpson of the United States. 

Wiren (1883) considers this to be only a variety of Nephthys caeca, Fabr., just as he 
joins to the same species Nephthys ciliata, 0. F. Miiller, N. longisetosa, (Ersted, and 
N. emarginata, Malm. 

De St. Joseph (1894) describes a pair of minute eyes at the inferior border of the 
first segment. He also points out the larger serrations at the flattened commencement 
of the blade of the capillary bristles. He is of opinion that the Portelia rosea of 
De Quatrefages may be this species. He distinguishes this form by the larger size of the 
branchiae about the thirtieth segment. 



9. .Nephthys incisa, Malmgren, 1865. Plate LVII, figs. 19 and 20; Plate LXVII, fig. 4 
—foot; Plate LXXVI, figs. 14 and 15— bristles. 

Specific Characters. — Head somewhat shield-shaped, slightly wider in the middle, and 
marked by two pale lateral areas which map out the surface. Anterior and ventro-lateral 
tentacles small and conical. Proboscis with twenty-two rows of minute papillae, and a 
long dorsal and ventral cirrus. Foot with a rounded dorsal lamella extending nearly 
to the tip of the organ. Small dorsal cirrus leaf -like and pointed, separated by a 
shallow notch from the somewhat massive though short branchia, which has the tip 
bent outward and forward. It diminishes and disappears posteriorly. Spinigerous lobe 
small; hidden between the rows of bristles. Fillet guarding the base of the barred 
bristles developed into a free flap superiorly and interiorly. Inferior division trifid, 
the anterior fillet projecting most superiorly. Neither flap is much developed, and the 
spinigerous lobe is conical. Bristles dark-brown, the anterior having the barred region 
beyond the widest part, and the capillary forms being curved and minutely serrated. 



Synonyms. 

1853. Nephthys ingens, Stimpson. Mar. Invert. Gr. Manan, p. 33 (fide Yerrill ?) ; and N. circinata, 

n.s. idem., and ditto, Amer. J. Sc. and Arts, 1875, vol. ix, June, 
p. 414. 

1865. „ incisa, Malmgren. Nord. Hafs-Annul., p. 105, Tab. xii, f. 21. 

1867. „ ., idem. Annul. Polych., p. 18. 

1868. „ „ M. Sars. Fortsatte Bemoerk., p. 10. 

1873. „ „ Kupffer. Exped. Ostsee, 'Pommerania/ p. 150 (Jahresb. Com. deutsch.). 
„ „ „ Gr. O. Sars. Bid. Kundskab. Christ. Fauna, iii, p. 23. 

„ „ ingens, Yerrill. Amer. Journ. Sci. and Arts, vol. v, p. 103. 

1874. „ incisa, Malm. Op. cit., Goteb., p. 77. 

„ „ „ (or ingens), Smith and Harger. Trans. Conn. Acad., iii, pp. 16, 18, 22, and 39. 

,, „ ingens, Yerrill. Proc. Amer. Assoc. Sc, 1873, p. 347, pi. ii, fig. 2. 

„ „ „ idem. Peabody Acad. Sc, vi Rep., p. 60. 



NEPHTHYS INCISA. 39 

1875. Nephthys incisa, Ehlers. Zeitscli. f. w. Zool., xxv, p. 19. 

1876. „ „ Mcintosh. Proc. Roy. Soc, xxv, p. 216. 

„ „ ingens, Yerrill. U. S. Comm. F. and Fish., p. 583. 

1879. „ incisa ( = ingens, Stimpson), Yerrill. C. List, U. S. Comm. F. and F., p. 7. 

„ Webster. 32nd Rep. N. Y. Mus. Nat. Hist., p. 104. 
„ „ ingens, idem. Trans. Albany Instit., p. 213. 

1883. „ incisa, Levinsen. Yidensk. Meddel. Foren. Kjoben., p. 217. 

1884. „ „ Webster and Benedict. Ann. Chart. Mass. (from U. S. Rep. F. and F. for 

1881), p. 702. 
1893. ,, „ Levinsen. Yidensk. Ud. 'Hauchs/ p. 338. 

1900. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. v, p. 262. 

1901. „ „ Whiteaves. Geol. Surv. Canada, No. 72^, p. 83. 



Habitat. — Connemara, Galway, 1871 (A. Gr. Moore). 

Extends to Greenland ('Valorous' Expedition), East American shores (Verrill), 
Canada; 'Porcupine' Expedition, 1869, 6—80 fathoms; 'Porcupine,' 1870, nine miles 
off Cape Finisterre, 81 fathoms. 

Head (Plate LVII, figs. 19 and 20) somewhat shield-shaped, nearly of the same 
breadth from front to back, though it is a little wider in the middle ; marked by two pale 
lateral areas, which map out the surface. Anterior and ventrolateral tentacles small 
and conical. 

The mouth (Plate LVII, fig. 20) has a short filament towards the anterior part of 
each lip. 

Proboscis with twenty-two rows of minute papilla?. A short median cirrus occurs 
in the smooth distal region both dorsally and ventrally (Malmgren). 

The species is characterised by the dark brown bristles, the slight development of 
the lamellae, and the wide space between dorsal and ventral divisions. The typical foot 
(Plate LXVII, fig. 4) presents in the anterior region of the body a rounded dorsal 
lamella, as in JSf. longisetosa, (Erst., though it has a different shape, is smaller, and 
extends nearly to the tip, whereas the lobe in the latter does not. At the tip of the 
foot and placed behind the former is the small leaf-like dorsal cirrus, acute at the tip, 
and separated only by a shallow notch from the somewhat massive short branchia, 
which is coiled with the tip outward and forward. The two processes (cirrus and 
branchia) move freely together, and are separated from the dorsal lamella. The 
spinigerous region is comparatively small, and is hidden between the rows of bristles. 
The fillet guarding the base of the barred bristles is developed into a considerable free 
flap superiorly and inferiority, so that it and the dorsal lamella hold the bristles between 
them. 

The inferior division presents a somewhat trifid condition, the posterior flap 
or lobe projecting sufficiently to guard the bases of the bristles, while the fillet 
in front is better developed and projects more superiorly. The conical spinigerous 
region is thus guarded by the rows of bristles and the flaps. The ventral cirrus is 
small and conical. 

In the middle of the body the branchia becomes a short and slender process directed 



40 NEPHTHYS PANSA. 

inward and uncoiled, but the dorsal lamella is distinct. By-and-by only the dorsal cirrus 
is visible, the branchia having disappeared. 

The anterior bristles (Plate LXXVI, fig. 15) are olive coloured, and have com- 
paratively short barred tips, the widest part of the bristle being below the barred region. 
The capillary bristles are short, the blade rather broad, flattened, doubly curved, and the 
edge finely serrated (Plate LXXVI, fig. 14). 

There is some hesitation in identifying this form with Malmgren's, and the 
condition of the specimen leaves room for further investigation. 

Parasitic Infusoria occur on the species both in British and Arctic waters. 

The Nereis incisa of 0. Fabricius x has no connection with this family, being a Nereis. 

So far as the figures and descriptions go, the Nepthys minuta of Theel 2 (1879) would 
appear to be closely related to this form. Not even the presence of ova in the small 
examples dealt with would lead to any other conclusion. 

The Nephthys atlantica of Hansen 3 (1879) is another form which resembles this 
species. 



10. Nephthys pansa, Elders, 1875. 

Specific Characters. — Head as in allied forms (?). Anterior tentacles more slender 
than the posterior. The long, massive, and dark-coloured proboscis has three or four 






Fig. 39.— Extruded proboscis of Nephthys pansa, Ehlers. After Ehlers. Enlarged. 

triangular papilla? in each of the twenty-two rows. Foot short and thick. The dorsal 
division has a very slightly developed lamella. The cirrus and branchia arise from 
the under surface of the division. The cirrus is short and conical; the branchia forms 
a flat conical leaf. The bristles are dark. The inferior division is even shorter than 
the dorsal. 

1 ' Skr. Natnrh. Selsk./ 5, p. 160, Tah. iv, f. 1—3, 1799. 

2 ' Annel. X. Zembla/ p. 28, Tab. ii, f. 18. 

3 < Nyt Mag./ 24, p. 6, pi. iii, fig. 1 and 2, and < Den Norske Nord-Exped./ p. 31, Tab. iv, f. 1—4. 



PHYLLODOCLCLE. 41 



Synonyms. 

1875. Nephthys pansa, Ehlers. Zeitsch. f. w. ZooL, xxv, p. 40, Taf. hi, f. 1 and 2. 
1881. „ „ Horst. Mederl. Arch. Zool. Suppl., Bd. 1, p. 7. 

Habitat— Dredged in 426—458 fathoms on July 30th, 1869, 51° 1' N., 11° 21' W., 
in the ' Porcupine ' Expedition of that year. 



Fig. 40. — Foot of Nephthys pansa, after Ehlers. Enlarged. 

As only a single specimen of this form (about 66 mm. in length) was procured in the 
4 Porcupine ' Expedition of 1869, the minute details of its structure are in need of further 
examination (Figs. 39 and 40). It seems to approach Nephthys incisa and N. nuclipes, as 
Ehlers states, but whether it is a variety of the former or a distinct species appears to be 
an open question. The form of the branchia is certainly peculiar. 



FAMILY VIL— Phtllodooidjb. 

Head bluntly conical, trapezoidal, cordiform, or sub-oval; sub-tentacles two, as 
short subulate organs (Grube) ; tentacles two or three, the unpaired posterior. Eyes 
two, posterior, generally small, rarely four. Large eyes exceptional, and in some probably 
only in the epitokous forms. Body long and vermiform, rounded or depressed, segments 
numerous ; caudal cirri two, short. Buccal segment with one to four pairs of tentacular 
cirri. Proboscis long, bipartite, with short papillse ; no jaws. Blood colourless. Feet 
as a rule simple, with one spine and a fascicle of compound bristles ; capillary bristles in 
certain epitokous forms. Dorsal and ventral cirri foliaceous. 

In the Phylloclocicla3 the body-wall as a rule (Fig. 41) has much less massive muscles 
than in the ISTephthy elicit. The cuticle lies on a thick cellular hypoderm within which 
are the basement-membrane and the circular muscular layer. The nerve-cords are in the 
mid-ventral line and more or less in approximation. The oblique muscles generally pass 
down on each side of the nerve-cords to be inserted into the basement-membrane, though 
these muscles may decussate quite beneath them and thus press the nerve-cords inward. 
No neural canals occur in this family in Britain. The dorsal longitudinal muscles are 

63 



42 PHYLLODOCIDiE. 

separated by a cleft, though in some, as Phyllodoce gramlandica, the separation at the 
dorsal margin seems to be incomplete. On the other hand the separation is pronounced 
in Eteone picta. 

In this family the segmental organs consist of a nephridium, simple or ramified, shut 
off from the coelom and covered with solenocytes. At the moment of reproduction a ciliated 
funnel appears, becomes connected with the nephridial tube, and conducts the sexual 
elements externally (Goodrich and Fage). It was probably the movement of the cilia 
of the segmental organ that Huxley l observed at the bases of the feet of Eulalia viriclis. 

Like other soft marine worms the Phyllodocicls were placed under the Vermes Mollusca 
of Linnaeus ; and subsequent authors, for a considerable time, followed a similar course. 

Savigny (1820) first separated this group as the eleventh genus of his Family 
Nereides, giving a fair account of the principal characters. He was followed by (Erstecl, 
who made it the second division of the Nereids. 




Fig. 41.— Section of the anterior region of Phyllodoce lamelligera, L. d, intestine ; dm, dorsal longitudinal muscles ; 
dc, dorsal cirrus ; dv, dorsal blood-vessel ; nc, nerve-cords ; so, segmental organs ; vc, ventral cirrus ; vm, ventral 
longitudinal muscles. 

De Quatrefages (1865) divided the family into two great groups, viz. the Phyllo- 
dociclae proper, and the Alciopidae, and he pointed out the distinctions between them. 
He arranged the genera of the Phyllodocidae according to the structure of the feet, the 
nature of the tentacles, and the tentacular cirri. 

The Phyllodocea were described by Kinberg (1865) as having the " antennae and 
palpi" equal (for so he diagnosed the anterior tentacles), a single tentacle or none; 
pharynx long, protrusible and devoid of jaws, branchiae foliaceous. He distinguished 
Phyllodoce from Eulalia by the presence of the median tentacle in the latter. He followed 
De Quatrefages in retaining Garobia. 

Ehlers (1868) describes the family as possessing an elongated nearly equally seg- 
mented body. Head with tentacles and eyes. The two or three following segments bearing 
four pairs of tentacular cirri — with or without feet. The rest of the segments distinct. 
Feet not prominent, with jointed bristles ; dorsal and ventral cirri leaf -like. Proboscis 
forming a long, closely papillose tube, and followed by a straight thick- walled stomach. 

1 'Mecl. Times and Gazette/ July 26th, 1856. 



PHYLLODOCIDJE. 43 

He showed, as indeed Claparede had already done, the erroneous nature of the view 
of Dr. Williams x that the ccelomic fluid circulated in the leaf -like cirri. These consist of 
cuticular and hypodermic tissue with glands. The blood is coloured but without cor- 
puscles. Dr. Williams, however, had correctly stated that the cirri are not permeated by 
blood-vessels. 

Grrube 2 gives the following arrangement for the Phyllodocidae : — 

I. — Tentacles five. 

Notophyllvm. 

Eulalia, Sav. 

(a) Eulalia, sans str. 
Eumida. 

Sige. 

(b) Pterocirrus. 

II. — Only two tentacles and two sub-tentacles. 

Myriocyclum. 
Kinbergia. 
Chsetojparia. 
Phyllodoce. 

Langerhans (1879) quotes Grube's classification of the family from the ' Annulata 
Semperiana,' viz. : (1) All the tentacular cirri springing from one segment — -Genet ijllis, 
Mgrn. (2) Three pairs of tentacular cirri from the same segment. A pair of bristle- 
bundles on the second segment — Anaitis, Mgrn. (3) Three pairs of tentacular cirri and 
bristle-bundles on first, or on first and second segments. One pair on the following : — 
Garobia (De Quatref.) Marenz. (4) Both of the first segments bear two pairs of ten- 
tacular cirri, as in Phyllodoce Paretti, P. comiculata, etc. In the second division with 
five " antennas " (tentacles) and four pairs of tentacular cirri he has the following groups : 
a, Proboscis well furnished with papillae : (1) All the tentacular cirri similar — Eulalia; 
(2) Cirrus tentaculus ventralis with a broad border — Pterocirrus. b, Proboscis with a ring 
of papillae in front — Eumida. c, Proboscis smooth, without papillae — Sige. 

Levinsen (1883) followed Malmgren so far as to place the Phylloclociformia after the 
Aphroditiformia, the Nephthydidae being a sub-division. 

Pruvot, after a careful examination of the nervous system of the group, proposed in 
1885 the following classification : — 

I. Five antennae. A. The first three dorsal cirri subulate : Eulalia (including 

Eulalia, Sav. ; Kinbergia, Quatref. ; Eracia, Quatref. ; 
Eumida, Mgrn.). 
B. The first two dorsal cirri subulate {Eulalia incompleta, 
Val.?; E. obtecta, Gr. ?). 

1 'Philos. Trans./ 1852, ii, p. 627. 

2 l Scliles. Gesellscli./ November 19th, 1879. 



44 PHYLLODOOID^E. 

II. Four antennse. a. The first three dorsal cirri subulate : Phyllodoce (Phyllo- 
doce, Sav. ; Lugia, Quatref.; Carobia, Quatref.). 
B. The first two dorsal cirri subulate : 

1. Dorsal cirri on all the segments; 

2. The third segment without a dorsal cirrus. 

c. Only the first dorsal cirrus subulate. The second without 
a dorsal cirrus (Eteone). 

He alludes to the inextricable confusion in the classification of the Phyllodocidas, and 
points out that the innervation of the dorsal and ventral cirri and their form give 
important data. The ventral cirri pass insensibly from the foliaceous to the subulate, but 
the dorsal change their type abruptly at the second or third segment. In other instances 
a segment deprived of dorsal cirri is intercalated between the dorsal subulate and the 
ventral foliaceous. He insists on the fact that the appendages of the buccal segment 
(so-called tentacular cirri) receive their innervation from the first ventral ganglion. 
Moreover, he regrets that the name " buccal segment " should be applied to a double ring 
as in the Eunicidas, or, as in the PhyllodocidaB, to three segments which have distinct 
appendages and nerve-supply, whereas the first feet of the NTephthydidae and Opheliacea3 
receive their nerve-supply from the ganglion of the oesophageal connectives. He would 
therefore arrange the cephalic segments thus : a stomato- gastric with its appendages — the 
palps ; an anterior antennary carrying the two anterior lateral antennse ; a posterior 
antennary bearing the median and the two posterior lateral antennse. The median 
antenna (tentacle) receives a nerve-trunk formed by the union of two nerves, and hence 
he supposes that the median was originally paired. 

Gravier (1900) describes the Family (in which he includes Lopadorhynchus and the 
Alciopidas) as having a head (prostomium) provided generally with four or five tentacles 
(his antennas). First segments of the body, in variable number, with reduced parapodia 
or modified tactile organs. Foot in the other segments uniramous (exceptionally bi- 
ramous). Dorsal and ventral cirri foliaceous. As a rule — compound bristles. Tail 
(pygidium) with two subulate or foliaceous cirri. 

He divides the group into three tribes, the first — or Phyllodocidse proper are 

littoral, the second and third — Lopadorhynchidge and Alciopidse — are pelagic. 

He classifies them after Grube and others according to the number of their tentacles 
and tentacular cirri, thus : — 

Two tentacular cirri (one on each side) . Eteone. 



Four tentacles 



Three „ 

Four 

,Five 

(One tentacular cirrus 

Five tentacles . \ . . rfoot uniramous 

I r our tentacular cirri j 

I I „ biramous 

Development. — Alex. Agassiz l has described the young stages of a form which he 
calls Phyllodoce maculata, from the condition in which the larva has only a few indistinct 



Mystides. 

Phyllodoce. 

Kinbergia. 

Porroa ? 

Eulalia. 

Notophyllum. 



Ann. Lyceum Nat. Hist. New York/ viii, p. 333, pis. x and xi, figs. 46—51, 1 



PHYLLODOCIDJE. 45 

segments behind the enlarged head, to that in which there are about fifteen rings behind 
the prostomium, and the anal cirri are present. 

Claparede and Mecznikow l (1869) give an account of the development of Phyllodoce 
at Naples and Odessa, but it is not known if the same species is referred to. The 
youngest stage is like that of Nephthys — monotrochous, the anterior portion of the larva 
being larger than the posterior, and a characteristic curved tuft (Naples) or tufts (Odessa) 
of cilia occur on the ventral surface anteriorly as Busch noticed. Segmentation begins 
behind the mouth, a pharynx appears in front, bristles form in the feet, tentacles, and 
tentacular and dorsal cirri develop, the alimentary canal becomes deep green, and there 
are short anal cirri — this stage being only a little younger than that procured at St. 
Andrews on June 15th, 1898, and figured (Plate L, fig. 6). 

Claparede (1868) notes that the eggs in the Phyllodocidse penetrate into the extruded 
proboscis. 

The young Phyllodoce maculata, according to Hacker (1896), is on the second day a 
protrochophore, on the fifth day a typical monotrochous trochophore (Mcintosh), tenth 
day an early metatrochophore, and at four weeks with specific characters (Agassiz). 

He further describes a teiotroch trochophore of a Phyllodoce from Naples in spring, 
in which what he calls the umbrella (viz. the prostomium) is about half the length of the 
body. It bears a large tuft of cilia on the ventral surface, a larger and a smaller pigment- 
speck (eye) on each side in front of prototroch, and a broad belt (paratroch) posteriorly. 
The gut (mittelclarmes) is greenish and shows a pair of sac-like appendages. 

Another form is in a more advanced condition (metatrochophore), with eight tentacular 
cirri (rudimentary), and indications of a number of segments posteriorly. The prototroch 
has a brown band of pigment before and behind it. A larger and a smaller eye occur in 
each side of the prostomium, the gut is greenish, and the tail is disc-like. 

He mentions and figures a " Scheitelorgan " of the larva of Phyllodoce, like a ciliated 
pigmented cell in a pit. 

He homologises the anterior pair of antennae with the Scheitelantennen of the larva, 
whilst the unpaired sense-organ on the ventral side of the umbrella (prostomium) he 
thinks the Scheitelorgan. 

The same author 2 describes a Nectochsete stage of Phyllodoce with four tentacular 
cirri, but without eyes or tentacles, and therefore younger than that in Fig. 6, Plate L. 

G-ravier (1900) observed that the Phyllodocidae fed on Terebellidge, Spionidse, Hermel- 
lidas, Sabellidse, Syllicla3, and even their own species (Eidalia viridis), and De St. Joseph 
found another Eidalia in the intestine of Eidalia pallida. 

De St. Joseph mentions the occurrence of a female Copepod on Eidalia pallida ; 
Herpyllobius arcticus, Steenstrup and Lutken, on Pterocirrus macroceros, Grube ; and one 
of the Orthonecticlas, viz. llhopalocera pterocirri in the same species. Gravier met with 
colonies of Vorticella on Phyllodoce mucosa. Gregarines of various kinds are common in 
the intestine — fixed to the epithelial lining, as in the Nemerteans. 

The Phyllodocidae arejto a large extent littoral, frequenting the tidal region, under 
stones, in moist sand near low water, on mussel-beds, in fissures of rocks, as well as in the 

1 c Zeitsch. f. av. Zool./ Bd. xix, sep. abdr. p. 27, Taf. xv, fig. 2. 

2 ' Plankton Expecl. pelag. Polychast./ p. 11, Taf. i, f. 5 and 5a, 1898. 



46 NOTOPHYLLUM FOLIOSUM. 

laminarian region, on oyster-shells and similar ground (Gravier). Genetyllis and Phyllodoce 
Paretti and other forms, however, range to deep water — Genetyllis ocidata of the 
' Challenger ' coming from a depth of 500 fathoms. At the time of reproduction some 
become pelagic, and develop a dorsal series of long natatory bristles. 

The colours of those from the deep sea are of the same character as those from 
shallow water. 



Genus XXX. — jNTotophyllum, Oersted, 1843. Char, emend. 

Body linear, tapered gently at each extremity, dorsum convex, ventral surface 
grooved and flattened. Two considerable eyes. Tentacles five, four anterior and one 
median. Tentacular cirri on two segments (Grube), four on each side, two shorter in 
front and two longer posteriorly. Superior division of the foot little differentiated, yet 
with a spine and a few simple bristles, and supporting a large irregularly reniform 
dorsal cirrus, the series being imbricate and almost covering the back and the feet. The 
inferior division has a much smaller vertical and somewhat reniform lamella, and a spine 
with many bristles, the shafts of which are curved distally and end in a dilated region 
having a series of spikes guarding the base of the long sabre-shaped distal blade, which 
has its edge very minutely serrated. The ventral border of the foot has a prominent 
process or mamilla, to which the inner edge of the lamellar cirrus is attached. Ganglia of 
the ventral cords large, resting on the sheath, with the circular muscular fibres externally, 
the oblique passing down on each side. In the interganglionic areas the insertions of the 
oblique muscles are nearer each other. The dorsal longitudinal muscles are completely 
separated and very broad. The ventral longitudinal muscles are compact. 

Notophyllum is said by Gravier 1 to have a small nerve-cord running above the two 
great cords. The sections of the British forms did not show this clearly, and thus 
differed from those of Nereis diversicolor, in which a third interganglionic cord with 
a neural canal is present. 



1. Xotophyllum foliosum, Sars, 1835. Plate XLV, fig. 1; Plate LXVII, fig. 5— foot ; 

Plate LXXVII, fig. 21— bristles. 

Specific Characters.— -Head forming a rounded cone, with two considerable eyes, 
occasionally with cuticular lenses. A ciliated flap or lappet on each side of the head 
posteriorly. Tentacles, five ; two occur on each side of the snout, are flattened, and 
somewhat pointed and shorter than the median. Two shorter and two longer tentacular 
cirri on each side. All these processes, as well as the head, are greenish-yellow with 
darker grains. Body linear, gently tapered anteriorly, convex dorsally, flattened ventrally, 
with a rectal papilla in the centre posteriorly. Proboscis smooth. Dorsal lobe of the 
foot directed upward and prolonged inward as a narrow process from which the great 
and more or less horizontal and irregularly reniform dorsal lamella (cirrus) springs. A 
slender spine and a few tapering smooth bristles occur towards the ventral edge of the 

1 < Bullet. Sc. Fr. Belg.,' 29, p. 331, pi. xix, f. 19. 



NOTOPHYLLUM FOLIOSUM. 47 

region. Inferior division with a smaller obovate or irregularly reniform lamella, and 
many slightly curved bristles with tapering terminal processes serrated at the edge. The 
tip of the shaft is dilated and fibrillatecl, with a distal series of spikes. Body pale, irides- 
cent, darker on the ventral surface. The dorsal lamellas are bordered with dark brown, 
dotted with white grains, and show a fiery iridescence. 

Synonyms. 

1835. Phyllodoce folios a, Sars. Beskriv. og Iagttag, p. 69, t. 9, f. 26. 

idem. Nyt Mag., 7b, p. 386. 

1843. Notophyllum viride, GErsted. Consp. Annul., p. 26, f. 87. 

,, „ ,, Koren. Nyt Mag*., ix, p. 95. 

1844-45. „ polynoides, GErsted. Kroyer's Nat. Ticls. (Forteg. ved. Drobak), p. 409, pi. ii, 

fig. 12. 

1851. ,, „ Grube. Fam. Annel., pp. 57 and 129. 

„ „ foliosum, idem. Ibid. 

1853. „ „ Sars. (Fauna of Adriatic). Nyt Mag. f. Nat., 7b, p. 386. 

1856. ,, polynoides, Koren. (Norske Fjords). Nyt Mag. f. Nat., 10b, p. 95. 

1865. „ „ Malmgren. Nord. Hafs- Annul., p. 93, T. xiv, f. 33. 

„ ., viride, De Quatrefages. Annel. II, p. 153. 

,, „ polynoides, idem. Ibid., p. 154. 

„ ,, foliosum, idem. Ibid., p. 162. 

1867. ,, polynoides, Malmgren. Annul. Polych., p. 19. 

„ „ ,, Malm. Zool. Obser., 7h, p. 77. 

1869. „ „ Mcintosh. Trans. R. S. Eclinb., xxv, p. 410, pi. xv, figs. 9, 9 a, 

and 9 b. 

1873. Notophyllum polynoides, Kupffer. Jaliresb. Comm. Deutscli. (' Pommerania '), p. 150. 
„ ,, foliosum, Sars. Bidrag. Christ. Fauna, iii, p. 24 (sep. copy). 

1874. „ polynoides, Malm. K. Yet. o. Vitt. Samli. Goteb. Hand., xiv, p. 79. 
„ „ foliosum, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 196. 

1875. „ „ idem. Invert, and Fislies St. Andrews, p. 119. 
1879. „ „ Tauber. Annul. Danic, p. 86. 

1883. „ ,, Levinsen. Yid. Medclel. Foren. Kjobenh., 1883, p. 204. 

„ Trachelophyllum Lutkenii, idem. Ibid., p. 52. 

1893. Notophyllum foliosum, idem. Yidensk. Udbytte. ' Hauchs,' p. 326. 

„ „ ,, Marenzeller. Zool. Ergeb., ii, p. 7, Taf. ii, f. 4. 

1901. ,, polynoides, Mcintosh. Ann. Nat. Hist., ser. 7, vol.viii, p. 223. 

1904. Eulalia obtecta, Allen. Journ. M. B. A., n.s., vol. vii, p. 223. 

Habitat, — Shetland (J. Gr. J.) ; Lamlash Bay, Arran (Dr. Howclen) ; Bay of Gralway 
(Dr. E. P. Wright); St. Andrews Bay, deep-sea fishing boats (E. M.) ; common in 
clreclgings, Plymouth (Allen). 

Norway (GErsted, Sars, Norman, and Koren); Sweden; Adriatic (Sars); Marseilles 
(Marion). 

Head forming a rounded cone with two large dark eyes, which occasionally show a 
central lens-like thickening ; between them is the well-developed and pointed median 
tentacle which generally projects in a slanting direction — forward and upward. On each 
side posteriorly a somewhat flattened lobe or lappet extends backward and slightly outward, 



48 NOTOPHYLLUM FOLIOSUM. 

the median region having darker pigment. The absence of these amongst other points 
in Malmgren's description and figure had probably misled Levinsen. The oral tentacles 
(two on each side) are finely pointed, somewhat flattened, and shorter than the median. 
The buccal segment has two longer and two shorter tentacular cirri on each side, the 
former being frequently carried along the dorsum under the two anterior ]amella3 (cirri), 
and over those behind. All these processes, like the head, are of a pale greenish yellow 
with darker grains. The mouth opens on the under surface, the prostomium in front 
having a ridge on each side with a hollow between, thus forming a channel leading to the 
mouth. The under lip resembles a somewhat broad cushion. 

Body linear, from If in. to 2 -J in., and from 80 — 110 segments, gently tapered at head 
and tail, and for the most part overlapped dorsally by the large imbricate cirri. The dorsal 
surface is convex, and each segment is crossed by several impressed lines, arranged 
symmetrically. The ventral surface is flattened, and frequently grooved in the centre. 
The rather small anterior region of the proboscis in protrusion forms an ovoid or 
somewhat elliptical mass, which, under a lens, is apparently smooth, though its surface is 
covered with minute granules under a higher power. It is thus practically smooth. The 
tail is terminated by a blunt papilla indicating the vent. 

The dorsal cirri are bordered with brown which increases in depth in those kept in 
confinement, and are speckled with white grains. The body is pale, iridescent dorsally, 
darker on the ventral surface, which is marked by minute dark grains on each segment. 
The lobes of the feet have blackish tips, the inferior papillae forming a row of rounded 
granular spots on each side. During the various motions of the animal the dorsal cirri 
present very fine iridescence of a fiery hue, as if from the reflection of red-hot coal. 
The softness and beauty of the species is accurately indicated in the coloured figure. 

The dorsal lobe of the foot (Plate LXVII, fig. 5) is directed upward, with a pointed 
prolongation inward, to which the great lamella is attached. The spinigerous papilla is 
at its outer border, but its translucent, tapering bristles are few (about two) and smooth. 
This lobe bears the large, more or less horizontal, dorsal cirrus which is described by 
Malmgren as elliptico-subrectangular or unequally reniform. The new or regenerated 
lamellae are more or less reniform, especially posteriorly, whilst the older cirri incline to 
an elliptico-subrectangular shape. The new lamella are pale, smooth at the edges, and 
minutely speckled with white grains. When fully developed the edge presents slight 
frills, and the coloured border is rich blackish-brown, which glistens now and then with 
the purplish-red iridescence of the surface of the lamella, enlivened here and there by 
groups of white grains. The first lamellae are smaller, and more elongated than those 
behind. 

The inferior division has a trunk not much less than the dorsal, and it carries a small 
obovate or irregularly reniform, more or less vertical, cirrus, the free border of which is 
marked by blackish brown pigment. Internal to the lamella, and connected with it, is a 
large lobe, or papilla, which is also marked by black grains, and thus a conspicuous 
moniliform row occurs along each side of the ventral surface. The first pair of ventral 
lamellae are nearly lanceolate, and are borne at a different angle from their successors, 
forming a pair of ventral processes, which stretch behind the labial cushion. This 
modification of the ventral division is common in the Annelids. 



NOTOPHYLLUM FOLIOSUM. 49 

The ventral bristles (Plate LXXVII, fig. 21 a, b, c) spring from the tip of the lobe, 
and they are seldom visible from the dorsum, even during the movements of the animal. 
The shafts of the bristles have a distinct curve, and strise pass from the base of the 
spikes for a short distance obliquely downward. On each side of the dilated end of the 
shaft a series of short spikes occur, thus, as it were, embracing the base of the terminal 
sabre, which is of moderate length and finely serrated at the edge. 

Reproduction. — Two female specimens, procured by my late friend Dr. Howden 
of Montrose, at Lamlash, Arran, in August, were laden with mature eggs. 

The species is more sluggish than the ordinary examples of the Phyllodocidae, 
and, when irritated, coils its body in a somewhat stiff manner. The colours, after 
seven months' confinement in Perthshire, were considerably altered, the dorsal cirri 
becoming much darker — brownish green, and the burnished-red iridescence was not 
visible. 

Louis Page 1 shows that the nephridia proper of this form resemble generally those 
of Eidcdia viridis, but that the solenocytes are studded with structures like palpocils so 
that they are hirsute. 

A figure given in Montagu's MS. drawings (1808) in the library of the Linnean 
Society (Plate LXIV, fig. 5) appears to refer to this species. Only three tentacular 
cirri are observed on each side. Though thus early indicated, this form was not 
recognized recently at the Plymouth Laboratory, being confounded with Eulalia. 

The early description of Sars (1835), as given in the ' Beskrivalser,' leaves little 
doubt that this was the species before him, the only ambiguous feature, indeed, being the 
outline of the inferior lamella (cirrus), a condition, however, probably due to the state of 
the specimen. Even the ventral papilla internal to the latter is clearly shown. It would 
be difficult to find any other form to answer the clear description of the author. 

CErstecl (1843), in his ' Conspectus,' apparently without a knowledge of the previous 
paper of M. Sars, correctly separated the genus from Phyllodoce since it has a bifid foot. 
He overlooked the fifth (median) tentacle, but he described the ventral mamilla of the 
foot. There is little to separate his descriptions of Notophyllum viride and N longum, 
and the somewhat elementary figures do not give much aid in this respect. His fig. 87, 
Plate Y, however, clearly refers to N foliosum. 

In his earlier publication (' JNTordiska Hafs-Annul.,' 1865) Malmgren gave a good 
description of the genus, improving on the brief account of (Ersted. He, however, did 
not associate the N polynoides of the latter with the N foliosum of Sars, and, while in his 
later publication (' Annel. Polychset./ 1867) he grouped the N longum of (Ersted with the 
present form, he appeared to have doubts about the species originally described by Sars. 
He had reasons for excluding Schuarda's Notophyllum 2 from his generic description. 

The Notophyllum japonicum of Marenzeller 3 appears to approach the northern species 
very closely. 

The Trachelophyllum Lutheni of Levinsen (1883) is this species. 

1 c Ann. Sc. Nat./ 9 e ser. iii, p. 281, figs. 6 and 7, 1906. 

3 ' Wirb. Thiere/ ii, p. 87. 

3 ' Sudjap. Annel/ (op. cit.), p. 126, Taf. iii, fig. 1, 1879. 

64 



50 EULALIA BILINEATA. 

Marenzeller 1 describes and figures two ciliated lappets on the head (one on each 
side). They are single in front, bifid behind. He therefore considers the' A", alatum of 
Langerhans to be the same form, especially as Levinsen mentions them in the northern 
species. 

Allen finds this species common in clredgings off Plymouth, but all the examples are 
small, though adult. Like certain northern forms the size diminishes in southern waters. 



Genus XXXL— Eulalia, (Erstecl, 1843. 

Head furnished with five tentacles, the median arising in front of the eyes. 2 First 
segment free from the head. Four pairs of tentacular cirri with or without feet beneath 
(first pair attached to first segment, (Erst.). Dorsal cirri (QErsted's branchiae) arranged 
in various ways. Ventral cirri nearly horizontal. Cerebral lobes with a special 
ophthalmic lobe ; median tentacle supplied by two nerves which unite at its base ; 
s tomato -gastric branches arising from oesophageal connectives (Pruvot). 

Claparede includes Eumida, Malmgren, and Eracia, Be Quatrefages (= Eumida) 
under this genus. He rather approves of the vagueness of the descriptions of the 
segments bearing tentacular cirri since it is impossible to fix hard and fast limits 
to these. 

Grube (1879) defines Eulalia as having the dorsal cirri lancet-shaped or pointed, 
more or less constricted at the base. 



Sub-genus Eulalia, sans str., Grube, 1879. 
Dorsal cirri oval- elongate, not pointed. 



1. Eulalia bilineata, Johnston, 1840. Plate XLIII, fig. 5 (var.) ; Plate L — fig. 4, head; 
Plate LXVII, figs. 6 and 7— foot, etc. ; Plate LXXVII, figs. 23 and 24— bristles. 

Specific Characters. — Head somewhat longer than broad; eyes two, black, placed 
towards the posterior border of the head. Anterior tentacles subulate, rather short. 
Median tentacle very short, arising just in front of eyes. Tentacular cirri likewise some- 
what short — only a little longer than the diameter of the body. Length from one and a 
half to three inches, the body being tapered anteriorly, and diminished more distinctly 
posteriorly, where it terminates in two subulate or sometimes clavate cirri. Colour dull 
yellowish, greenish-yellow, or straw, the proboscidian region of the body being paler. 
Ventral surface slightly greenish, a pale greenish band, most distinct anteriorly, passing 

1 "Polychast. des Grundes.," c Zool. Ergebn./ ii, p. 7, Taf. ii, fig. 4. 

3 The exceptional position of the median tentacle, viz. at the extreme posterior border of the 
head, in Ehlers' Eulalia imbricata, is noteworthy. 



EULALIA BILINEATA. 51 

along the base of the feet, and in some a faint, interrupted double line occurs in the 
middle. Ripe females are of a fine pea-green, a faint, interrupted double line extending 
along the middle. On the other hand, the ripe males are whitish-yellow and the posterior 
region of the body is enlarged. At the base of each foot dorsally is a dark speck. The 
dorsal cirrus is ovato-lanceolate, and the ventral has a similar shape. It (ventral) is not 
so prominent as the setigerous process, which bears bristles having the dilated ends of the 
shafts distinctly spinous. The shape of the foot approaches that in E. trilineata. 



Synonyms. 

1840. Phyllodoce bilineata, Johnston. Ann. Nat. Hist., iv, p. 227, pi. vi, f. 7 — 10. 

„ Grnbe. Earn. Anne]., pp. 56 and 129. 

„ Johnston. Cat. Brit. Mns., p. 177, pi. xvi, f. 7 — 10. 

„ De Qnatrefages. Annel., ii, p. 137. 

„ Malmgren. Nord. Hafs-Annul., p. 99, Tab. xiii, f. 26. 

„ idem. Annul. Polych., p. 25. 

„ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 197. 

„' Malm. K. Yet. o. Yitt. Samli. Goteb. Handl., xiv, p. 80. 

,, Mcintosh. Invert, and Fishes, St. Andrews, p. 120. 

„ Tauber. Annul. Danic, p. 89. 

„ Langerhans. Canar. Annel., Nov. Act. Leop., xli, p. 113. 

„ Levinsen. Yidensk. Meddel. Foren. Kjoben., p. 205. 

Habitat. — On oysters near Prestonpans, in the Firth of Forth ; Berwick Bay, rare 
(Dr. Johnston) ; in fissures of shale and sandstone north of the pier, St. Andrews ; under 
stones at the East Rocks, St. Andrews, where it is abundant (B.M. and R.M.), and 
between tide-marks, St. Peter Port, Guernsey; dredged also in the Sound of Harris in 
6 — 8 fathoms ; under the crust of Corallina officinalis in company with Polydora, near low- 
water mark, Lochmaddy, North Uist. 

Finmark (Malmgren). Canaries (Langerhans). 

Head (Plate L, fig. 4) somewhat ovoid, longer than broad. Eyes two, black, 
placed towards the posterior border of the head and larger than in Malmgren's figure. 
Anterior tentacles subulate, rather short ; median tentacles very small, arising just in 
front of the eyes. The tentacular cirri are likewise somewhat short, being only a little 
longer than the diameter of the body, and therefore Dr. Johnston's form — with these 
organs three times the diameter of the body — differs. 

Body from one and a half to three inches in length, tapered anteriorly and more 
distinctly diminished posteriorly, where it terminates in two subulate cirri. The general 
colour is dull yellowish, greenish-yellow, or straw-coloured, with a distinct greenish- olive 
or brownish line along each side, rather within the bases of the feet, and often presenting 
a crenated appearance from the folding of the segments — especially anteriorly. The 
proboscidian region of the body is paler from the hue of the proboscis. The ventral 
surface is slightly greenish, a pale-greenish band passing along the bases of the feet to 
the posterior end, but being most distinct anteriorly. In some examples a faint, inter- 



1851. 


)? 


1865. 


)) 


?j 


)j 


?> 


Eidalia 


1867. 


>) 


1874. 


» 


» 


» 


1875. 


)> 


1879. 


)> 


1881. 


>) 


1883. 


35 



52 EULALIA BILINEATA. 

rupted double line extends along the middle. When filled with ova, e. g. in May, they 
are of a fine pea-green hue with the two dark-brownish lines. On the other hand, the 
ripe males (July) are whitish-yellow from the contained sperms, the posterior region of 
the body especially being enlarged. In spirit-preparations a dark speck is observed on 
the dorsum at the base of each foot ; whilst in life a dark spot occurs on each side of the 
segments ventrally. 

The typical foot somewhat resembles that in Eulalia trijnmctata, and it is doubtful if 
the species could be distinguished readily by the outlines of the feet when mounted. The 
bristles (Plate LXXVII, fig. 23 a and b) are also very similar, the only feature being the 
presence in this species (E. bilineata) of a more distinctly spinous region at the enlarged 
distal end of the shaft, the slightly longer bristles and their more evident curvature. 

Both dorsal and ventral cirri are pale and ovato-lanceolate. The ventral is not so 
prominent as the tip of the setigerous process. 

Reproduction. — Both males and females are ripe in the summer months from May to 
July, the females in this condition being pea-green, and the males whitish-yellow — 
the posterior half of the body being much enlarged. The sperms (Plate LXVII, fig. 7) 
are comparatively large, have a globular head and a long tail. Their vitality seems to be 
considerable. 

Moreover the feet of the males have long simple bristles (Plate LXXVII, fig. 24) 
dorsally in addition to the ordinary forms, so that an epitokous condition is present. 
They also swim readily : even the separate fragments pass swiftly through the water with 
a wriggling screw-motion. In the ripe forms the muscular walls of the body are thinner. 
Habits. — A restless species — constantly pushing its snout here and there and throw- 
ing its mobile body into various folds and loops. It is common in the tidal region, 
lurking in crevices of the fissured rocks along with Spio, in cavities on the under surface 
of stones, and under empty shells to which it adheres. It is slender and graceful, 
moving in captivity about the vessel swiftly and with somewhat regular undulations. 

In his original description in 1840 Dr. Johnston omitted the short median tentacle, 
but in the ' Catalogue of Worms in the British Museum ' he adds that Mr. Dyster 
observed it. 

The Eulalia saxicola of De Quatrefages l appears to be most nearly related to this 
form. 

Such species as the Eulalia gracilis of Verrill 2 approach this form in the closest 
manner, and further examination may render separation unnecessary. 

The Eulalia quadrilineata of De St. Joseph 3 from Cape Coz is an allied species. 
An Eulalia from deep water off St. Andrews Bay, which has only been seen in spirit, 
differs from the foregoing species in its larger size, greater development of the dorsal 
lamella, and the slightly broader tips of the bristles. The head is somewhat ovate, with 
the long diameter antero-posterior, and with a pair of small black eyes situated near the 
posterior border. The anterior end is small, and the body gently dilates from before 

1 < Annel./ ii, p. 119, 1865. 

2 ' Invert. Vineyard Sound, U.S. Comm. P. and F./ 1878. ? 

3 ' Ann. Sc. Nat./ 8 e ser., t. v, p. 327, pi. xviii, figs. 127—130, 1898. 



EULALIA NEBULOSA. 53 

backwards, and probably again diminishes towards the tail, but the posterior region 
is absent. At the tenth foot the dorsal lamella is ovate with a somewhat blunt tip. The 
setigerous process is bifid, and the ovate or renif orm ventral lamella extends beyond its tip . 
The dorsal lamella considerably increases in size at the fortieth foot, but the ventral remains 
nearly the same. The shafts of the bristles (Plate LXXXIV, fig. 5) are somewhat slender, 
with a slight curve distally, the enlargement and its spines resembling the parts in Eulalia 
bilineata. The terminal process begins with a broad base, and rapidly diminishes to a 
hair-like tip. The edge is finely serrated, and the serrations appear to be continued on the 
shaft a considerable distance downwards. 

This may be a large example of E. bilineata, and is a male laden with sperms. 



2. Eulalia nebulosa, Montagu, MS. (1808). Plate XLIII, fig. 6 ; Plate LVIII, figs. 1 
and 2— head; Plate LXVII, figs. 8 and 9— feet ; Plate LXXVII, fig. 1— bristle. 

Head broadly conical, the median tentacle arising a little in front of the eyes, which 
are large and blackish-brown. Tentacles pale, with a dark speck at the base of the 
median, one at the base of the anterior pair of tentacular cirri, and another on the enlarged 
part of the latter pair. Body five to ten inches long, somewhat soft in texture, tapered 
considerably anteriorly and more evidently so posteriorly, where it terminates in an 
attenuated tail with the anus at the tip. Only one or two examples — out of dozens — 
presented two small and somewhat clavate anal cirri. The finely tapered tail is charac- 
teristic. It is of a dull yellowish or pinkish hue, minutely speckled on the dorsum with 
dark green touches which are symmetrically arranged — a long lozenge occurring in the 
middle line in front and a bar behind — stretching between the feet of opposite sides. 
Montagu figures a row of six spots at the segment-junction, and four on the space behind. 
Ventral surface pale. Proboscis very long, with a rasp-like series of minute papillaa. 
Dorsal cirrus large and cordate, the margin dull green, and some have a V-shaped mark 
in the centre of the process and one or two rows of dark specks above it. The pedicle is 
large. Setigerous lobe bifid ; bristles pale^ with an enlarged and bevelled tip, while the 
terminal blade is moderately long, and with a finely serrate edge. The ventral cirrus is 
o vate-acumm ate . 

Synonyms. 

(1808). Eulalia nebulosa, Montagu. MS., Linnean Soc. 

1860. Phyllodoce (Eulalia) punctifera, Grube. Arch. f. Naturges., t. i, p. 83, Taf. hi, f . 5. 

1867. „ nebulosa, Parfitt. Trans. Devon Assoc, ii, pt. 1, p. 233 (near G-riffithsiee, fide 

Parfitt). 
1885. Eulalia punctifera, Cams. Fauna Medit., ii, p. 242. 

1888. „ „ De St. Joseph. Ann. Sc. Nat., 7 e ser., t. v, p. 289, pi. xii, f. 155—157. 

1897. „ „ G-ravier. Bull. Sc. Fr. Belg., t. xxix, p. 308, pi. xvi, f. 17 ; pi. xvii, f. 1, 11, 

12; pi. xviii, f. 5; pi. xix, f. 1—5, 20; pi. xx, f. 10, 11; 

pi. xxiii, f. 2. 
1904. „ „ Allen. Journ. M. B. A., n.s., vol. vii, p. 223. 



54 EULALIA NEBULOSA. 

Habitat. — Abundant at St. Peter Port, Guernsey, and at Herm, under stones between 
tide-marks amongst muddy gravel, and occasionally in mud under dead limpet- shells. 
Col. Montagu dredged specimens about four inches long at Torcross in 1813. Plymouth 
(Allen). Mediterranean (Grube). 

Head (Plate LVIII, figs. 1 and 2) broadly conical; the median tentacle springing 
from the surface a little in front of the eyes, which are two, large, blackish-brown. 
Tentacles pale, with a dark speck at the base of the median, one at the base of the posterior 
pair near the eyes, and another at the enlarged part of the latter pair. The tentacles and 
cirri generally are of moderate length and somewhat thick. In many preserved specimens 
a dark brown speck of pigment remains on each side just in front of the median tentacle. 

Body five to ten inches long, somewhat soft in texture, tapered considerably anteriorly 
but much more so posteriorly, where it ends in a long slender tail, having two small 
cirri at the tip, which, however, are rarely seen. It is of a dull yellowish or pinkish hue 
minutely speckled on the dorsum with dark green touches, which are symmetrically 
arranged — a long lozenge occurring in the middle line in front, and a bar behind — 
stretching between the feet of opposite sides. Montagu's figure shows one row of six 
spots at the segment-junction, and another series of four on the space behind. Sometimes 
the hue of the dorsum is less pale orange and more of a dull yellowish tint. The ventral 
surface is pale. After preservation in spirit a narrow transverse bar (dorsally and 
ventrally) characterizes each segment-junction. It is wider in the middle and tapers off 
at each margin. 

In the coloured figure (which I owe to the courtesy of Dr. Allen of the Plymouth 
Marine Laboratory) each segment is marked by a transverse row of brownish clots, the 
outer on each side being the larger, and behind is a row of smaller dots. 

The proboscis (Plate LVIII, f[g. 2) is remarkable for its great length, narrow cylin- 
drical form, and dense series of minute papilla?, so that its surface is rasp-like. In some the 
entire alimentary apparatus seems to be ejected (in spirit) through a rupture of the 
body-wall. The fixed end of the proboscis is firm and muscular, and is succeeded by a 
short glandular region with thinner walls ; then the firm proboscis increases in diameter 
to its junction with the alimentary canal proper, a well-marked constriction occurring at 
this point. The first part of the region is firm, but it soon becomes thin and glandular. 
The specimens are somewhat softened. 

The contents of the alimentary canal were sandy mud containing foraminifers, 
masses of debris of Neplithys with bristles, bristles of other annelids, and sponge-spicules. 

The typical foot (Plate LXVII, figs. 8 and 9) presents dorsally a large and somewhat 
cordate cirrus, the colouration of which is subject to some variation. Thus many have a 
dull green border dorsally, whereas in those from Herm the centre of the lamella had a 
dark V-shaped mark, with one or two rows of dark specks above it. The lamella? 
spring from a prominent peduncle (ceratophore), which in the preserved specimens forms 
a feature of the dorsum — especially as in many they are devoid of lamella?. The lamella? 
vary a little in shape at the base, from the nature of the supporting peduncle, the anterior 
(at the tenth foot) presenting a truly cordate base, whereas at the ninetieth foot the broad 
base is nearly flat. The hypodermic tissue is arranged in a fan-like manner from the base 



EI7LALIA VIRIDIS. 55 

outward on both sides in each papilla. The setigerous process is deeply bifid, the upper 
limb being the larger. The pale bristles have a slightly bent shaft with an enlarged, 
bevelled, and slightly bifid tip, which is more or less smooth, for articulating with the 
terminal whip, which has a moderately elongate, flattened blade with a finely serrate 
edge (Plate LXXVII, fig. 1). 

Attached somewhat behind the inferior border of the setigerous process is the ovate 
acuminate inferior cirrus, the edge of which is rounded internally and pointed externally. 
The hypodermic streaks run from the base outward to the free edge. The tip does not 
extend so far outwards as that of the setigerous process. 

Reproduction. — In the posterior region of a large example in July, masses of eggs, 
apparently ripe, occurred. When distended with ova, as in July and August, the body- 
wall in section is very thin, the dorsal and ventral longitudinal muscles forming narrow 
bands, and even the nerve-cords join in the general flattening. The wall of the 
alimentary canal, however, remains of considerable thickness. 

The species is characteristic of the southern shores, and is soft and delicate. Slight 
interference results in rupture and disintegration — especially in warm weather. Thus in 
few examples is the tail furnished with cirri, and on the long, tapered region the feet 
are very minute, leaving the terminal anus prominent. 



3. Bulalia viridis, 0. F. Midler, 1776. Plate XLIII, fig. 7, var. aurea ; Plate XLIV, 
fig. 1; Plate XLIV, fig. 5, var. ornate; Plate LVIII, fig. 3— head ; Plate LXVII, 
figs. 10 and 11— feet; Plate LXXVII, figs. 2 and 2 a— bristles. 

Specific Characters. — Head bluntly conical and pale throughout ; eyes two, black, 
and occasionally an additional speck occurs external to each. Tentacles subulate, slightly 
tapered. First pair of tentacular cirri somewhat lanceolate, tapered, and attached to the 
first segment on each side ; the second and third are longer, and are attached to the 
succeeding segment, whilst the fourth springs from the third segment. Body linear, one 
to four inches long, slightly tapered anteriorly and more so posteriorly, and terminating 
in two lanceolate caudal cirri. Colour of a rich deep green, or grass-green, paler in the 
region of the proboscis, and dorsally each segment has two dark belts. Ventral surface 
marked by a median moniliform line of dark olive, and posteriorly with a dark touch at 
each foot. Proboscis with twenty-two or twenty-three papillae terminally in extrusion, 
but a variety has only fourteen. 

Dorsal cirrus lanceolate with a tapering tip ; setigerous region short ; bristles in two 
groups pale, the enlarged ends of the shafts minutely spinous, the terminal blade short, 
rapidly tapered, and with bold serrations on the edge. 

Ventral cirrus ovate and slightly acuminate, projecting a little beyond the tip of the 
setigerous lobe. 

Synonyms. 

1767. Nereis viridis, Linn. Syst. Nat., ed. 12, i, pt. 2, p. 1086. 
1776. „ „ 0. F. Miiller. Zool. Danic. Prodr., Xo. 2636. 



56 EULALIA VIRIDIS. 

1870. Nereis viridis, Fabricius. Fauna Groenl., p. 297. 

1789. „ „ Bruguiere. Encycl. Meth., Vers, i, p. 134, Tab. lvii, f. 7—1 1. 

1791. „ „ Linn, amelin's edit. (13), i, pt. 6, p. 3117. 

1799. „ „ Adams. Trans. Linn. Soc, v, p. 8. 

1800. Die grime Nereide, 0. F. Miiller. Naturges. einiger Wurm-Arten, p. 162, Taf. ii, f. 1 
1802. Nereis viridis, Bosc. Vers, i, p. 171. 

1806. „ „ Turton's G-melin, p. 88. 

1807. „ „ Ibid. Brit. Fauna, p. 135. 
1812. „ „ Pennant. Brit. Zool., iv, p. 94. 

1825. „ „ De Blainville. Diet. Sc. Nat., t. xxxiv, p. 445. 

1828. Nereiphylla viridis, idem. Diet. Sc. Nat., t. Ivii, p. 466, pi. xiii, f. 2. 

1833. Phyllodoce clavigera, Aud. & Ed. Ann. Sc. Nat., t. xxix, p. 248, pi. xvi, f. 9 — 13. 
„ „ Gervillei, idem. Ibid., p. 250. 

1834. „ clavigera, idem. AnneL, p. 226, figs. 9 — 13. 

1840. „ viridis, Johnston. Ann. Nat. Hist., iv, p. 228, pi. vi, f. 11 — 14. 

1842. Eulalia viridis, (Ersted. Kroyer's Nat. Tids., p. 122. 

1843. „ „ idem. Grronl. Annul. Dorsib., p. 188. 

„ „ „ idem. Consp. Annul. Danic., p. 27, f. 22, 30, 85, 86, 88. 

„ Phyllodoce clavigera, H. Rathke. Beitr. Fauna Norweg., p. 170. 

1844. „ „ De Quatrefages. Ann. Sc. Nat., 3 e ser., 14, p. 357, pi. ix, f. 9. 
„ Nereis viridis, Thompson. Rep. Brit. Assoc, for 1843, p. 273. 

1844-5. Eulalia viridis, CErsted. Kroyer's Nat. Tids., p. 410. 

1849. Phyllodoce clavigera, Cuvier. Reg. Anim. Illust., p], xiii, fig. 3. 

„ „ viridis, Leuckart. Archiv f. Naturges., p. 202. 

1851. „ [Eulalia) viridis, Grube. Fam. Annel., pp. 56 and 129. 

„ Eulalia viridis, Sars. Nyt Mag., vi, p. 209. 

1864. „ virens, Ehlers. Borst. i, p. 159, Taf. vii, f. 1—5. 
,, Phyllodoce [Eulalia) viridis, Grube. Insel Lussin, p. 81. 

1865. „ viridis, Johnston. Cat. Brit. Mus., p. 178, pi. 16, f. 11—15, and p. 342. 
„ Eulalia ,, Malmgren. Nord. Hafs- Annul., p. 98, Tab. xv, f. 39. 

„ ,, ,, De Quatrefages. Annel. ii, p. 122. 

„ „ clavigera, idem. Ibid., p. 118. 

,, Phyllodoce Gervillei, idem. Ibid., p. 139. 

1867. „ viridis, Parfitt. Trans. Devon. Assoc, ii, pt. 1, p. 232. 

„ Eulalia ,, Malmgren. Annul. Polycheet., p. 25. 

1873. „ „ Sars. Bidrag. Christ. Fauna, iii, p. 24. 

1874. „ „ Malm. Op. cit,, p. 80. 

„ „ „ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 196. 

„ „ .j Marenzeller. Sitz. Akad. Wien, sep. abdr., p. 18. 

1875. ,, „ Mcintosh. Invert, and Fishes, St. Andrews, p. 120. 

1878. „ „ idem. Trans. Linn. Soc. Zool., i. p. 502. 

1879. „ „ Tauber. Annul. Danic, p. 88. 

„ „ ,, Langerhans. Zeitsch. f. w. Zool., xxxiii, p. 309. 

1881. „ „ idem. Canar. Annel., Nov. Act. Leop., xli, p. 113. 

1883. „ „ Levinsen. Yidensk. Meddel. Foren. Kjoben., p. 205. 

1885. ,, clavigera, Pruvot. Arch. Zool. Exper., 2 e ser., t. iii, p. 291, pi. xiv, f. 9 11. 

1886. ,, viridis, Harvey Gibson. Proc Lit. Phil. Soc. Liverp., xi, p. 152. 

1888. „ „ De St. Joseph. Ann. Sc. Nat., 7 e ser., t. v, p. 283, pi. xii, f. 154. 

„ „ ornata, idem. Ibid., p. 291, pi. xii, f. 158 — 161. 



EITLALIA VIRIDIS. $7 

1890. Eulalia viridis, Malaquin. Annel. Boulon., p. 43. 

„ ,, ornata, idem. Ibid., Rev. Biol. Nord., Tav. iii, p. 99. 

1891. ,, viridis, Hornell. Trans. Liverp. Biol. Soc, vol. v, p. 246. 
1893. ,, ornata, Levinsen. Yidensk. Ud. 'Hanchs/ p. 327. 

1897. „ aurea, Gravier. Bull. Sc. Fr. Belg., t. xxix, p. 309, pi. xvi, f. 2—6. 

„ ,-, viridis, idem. Ibid., p. 308, pi. xvi, f. 16; pi. xviii, f. 13 — 15; pi. xxi, f . 2, 3 ; 

pi. xxii, f. 3 — 6. 

1903. „ „ Mcintosh. Mar. Invert. S. Africa, vol. iii, p. 34. 

1904. „ ornata, Allen. Journ. M. B. A., n.s. vol. vii, p. 223. 
„ ,, aurea, idem. Ibid., p. 223. 

1905. „ viridis (= E. virens, Ehlers, and E. guttata, Clap.), Grseffe. Arbeit. Zool. Stat. Triest, 

xv, p. 325. 

1906. „ „ De St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 224. 

Habitat. — Abundant under stones in rock-pools and other places near low water- 
mark and in the fissures of shelving rocks about half tide level, where groups of five or 
six may be found, and also under tangle-roots at St. Andrews, as well as all round the 
British shores, from Shetland to the Channel Islands. It also ranges to deeper water, 
as off the Bell Bock (E. M.), and in Galway Bay (Dr. E. P. "Wright). 

Shores of France (De St. Joseph) ; Faroe (Willemoes-Suhm) ; Greenland (O. F. 
Miiller and CErsted) ; Madeira and Canaries (Langerhans) ; South Africa, col. Gilchrist 
(Mcintosh); Behring's Sea (Marenzeller) . 

Head bluntly conical and pale throughout. Eyes two, of moderate size, black, and 
occasionally an additional speck occurs external to either, probably due to brownish- 
green pigment deposits. The latter can with difficulty be detected in spirit-preparations. 
Tentacles subulate, slightly tapered ; median tentacle similar ; tentacular cirri greenish, 
four on each side. The first pair is short, somewhat lanceolate, tapered, and attached 
to the buccal segment (one on each side) ; the second and third which are longer are 
fixed to the succeeding segment, whilst the fourth springs from the third body-segment, 
and beneath is a modified foot with a few bristles, and an ovate ventral cirrus. 

Body linear, If to 4 in. in length, rounded dorsally and flattened ventrally, slightly 
tapered anteriorly and more distinctly so posteriorly, and terminating in two lanceolate 
and somewhat darker caudal cirri. The inner curve of the caudal styles is longer than the 
outer. Colour of a rich deep green or grass green, paler in the region of the proboscis. 
The lamellae are tinted of the same rich green, with a darker portion near the base, and 
in some males are pale olive. In others each segment shows dorsally two dark belts — 
narrower in the middle, broader externally — each being interrupted by a dark spot. 
When ova are present in these varieties the posterior half has the ground-colour of a 
deep green. The ventral surface is marked by a median moniliform line of dark olive, 
supplemented posteriorly by a dark touch at each foot. A variety from the tangle-roots 
of Bressay Sound is deep yellow, duskier on the dorsum, which, as in the previous form, 
has two brownish lines, on each side of a central stripe. A lateral border of brown 
also occurs on each side. The lamellae and under surface are yellow. Others from the 
Outer Hebrides are greenish-yellow, yellowish-orange, or grass-green dotted with minute 
black grains, and with a darker line at each segment-junction, while a dull madder-brown 

65 



58 EULALIA VIRIDIS. 

variety also is met with. The ventral surface is somewhat pale, with a pale central line. 
The head is shorter and more rounded than in the green forms. 

The variety ornata (the Eulalia ornata of De St. Joseph) has a double bar in each 
segment in three sections, the middle touches being most distinct, as seen in the 
figure (Plate XLIV, fig. 5) courteously given by Dr. Allen of the Marine Biological 
Association at Plymouth. 

The Eulalia aurea of Gravier appears to be a variety closely resembling that from 
the tangle-roots of Bressay Sound, and the figure of which (Plate XLIII, &g. 7) is also 
given by Dr. Allen. 

The green pigment of this species has been examined by Dr. Marion Newbigin, 1 
and it appears to be similar to that found in other Invertebrates, e. g. Chastopterus. 

Externally the proboscis (Plate LVIII, fig. 3) presents anteriorly (in situ) a massive 
greenish portion with ribs, and posteriorly an elongated cylindrical region of a paler 
green. The anterior massive region has a series of greenish glandular papillae on its 
inner surface. The narrower portion succeeding is, on the other hand, lined only by 
rounded cells, the outer layer being pale and firm. 

Two varieties in regard to the row of terminal papillae in extrusion occur, viz. one 
with twenty-two or twenty-three papillae, the other with only fourteen. The firmness 
of the body in those showing the latter is marked. 

The first foot, as indicated, occurs in the form of a short setigerous process with a 
few bristles and a small ovoid ventral cirrus under the fourth tentacular cirrus. The 
next foot is normal, though somewhat small, having a lanceolate dorsal cirrus with its 
ceratophore, a bifid setigerous region with bristles, and a somewhat ovate ventral cirrus. 

The tenth foot (Plate LXVII, fig. 10) has a pointed lanceolate dorsal cirrus (lamella), 
the inner edge of which is nearly straight, or has only a slight curve at apex and base, 
whilst the outer edge has a convex curve which reaches its maximum about a third from 
the base and then is sloped inward. The short setigerous region is bifid, with the typical 
bristles, and the ventral cirrus is ovate with a blunt tip. 

The dorsal division of the typical foot (Plate LXVII, fig. 11) bears a lanceolate 
cirrus with a tapering tip, the midrib being nearly median, with the pinnae stretching 
obliquely outward and forward superiorly, and nearly at right angles posteriorly. The 
bifid setigerous process is somewhat short and bears a series of pale bristles, the shafts 
of which are slightly curved distally and the ends enlarged and minutely but distinctly 
spinous (Plate LXXVII, figs. 2 and 2 a) . The terminal spear is comparatively short, rapidly 
tapered, with rather bold serrations inferiorly, slanting outwards and upwards, that is, 
towards the delicate tip. The serrations are more evident than those of Eumida 
sanguined, though the terminal blade in the latter is longer. The pale spine separates 
the two groups of bristles in each foot. The inferior cirrus is ovate and slightly 
acuminate, and projects a little beyond the tip of the setigerous process, though in life 
it appears scarcely to do so. These lamellae vary a little in acuteness in specimens from 
different localities. 

Reproduction.— -Many ripe males and females are found in July. The females are 



i c 



Quart. Journ. Micr. So./ n.s., vol. xli, p. 427. 



EULALIA VIRIDIS. 59 

greenish inferiorly, whereas the ripe males are whitish on the ventral surface — from the 
masses of sperms. The latter have a globular head and a long tail like those of Nereis or 
Lepidonotus, and are much larger, for instance, than those of Girratulus tentaculatus. 

Lo Bianco 1 gives November as the month in which the sexual products are present 
in the Neapolitan examples. 

Louis Fage 2 has recently described the segmental organs of this form, the nephridia 
rarely having more than three divisions, fan-like in outline, the border being formed 
of the solenocytes from which small tubes lead to the duct, whilst the " pavilion " or 
genital duct is funnel-shaped and leads to the exterior, becoming specially enlarged at 
the epoch of reproduction, and transmitting the sexual products, as in other members 
of the family. 

Habits. — In the groups of adults procured in the fissures of the shelving rocks, 
amidst muddy debris, only occasionally are a few small forms, about If in. long, pro- 
cured. In confinement they remain at the side of the vessel, while rapid undulatory 
motions take place from head to tail, probably in connection with respiration. No 
ciliated band, however, was observed on the lamellae. They also seek shelter under 
Ascidians and sponges, and if prevented, sometimes coil themselves into a rounded knotted 
mass. Some take shelter both in the sea and in confinement in tubes of Serpula and 
Sabellaria, to which they cling with tenacity. The movements of this species have been 
recently described by Gr. Bolm 3 as largely due to what he terms " thigmotactisme." 
Bright light causes disordered movements. 

They tinge the spirit at first of a pink colour, and become themselves of various 
shades of straw colour or pale brown. 

They are chiefly littoral, but occasionally come from deeper water. 

0. F. Miiller's description (1771) of this, as of other Annelids, is remarkable for its 
general accuracy, and the same may be said of his figures in the " Nat. einiger Wurm- 
Arten." 

Audouin and Milne- Edwards (1834) described a Phyllodoce Gervitteii, which only 
differs from their P. clavigera in the absence of the median tentacle and the brevity of the 
tentacular cirri. It is possible that this absence may have been accidental, as no other 
species so closely approaches it. De Quatrefages followed, so that he represented this 
form by three species. 

The Eulalia virens of Ehlers (1864), from the Adriatic, comes so close to this species 
that it is doubtful if it can be separated. 

It is difficult to identify the Phyllodoce ellipsis of Dalyell, or the P. Griffdhsii and 
P. cordifolia mentioned by Johnston in the Catalogue of the Annelids in the British 
Museum (1865). Grube thought P. Griffithsii resembled his Eulalia punctata from St. 
Vaast la Hougue. 

This species seems to be subject to considerable variation in colour, for so far as 
can be observed, the E. ornata of De St. Joseph (1888) differs only in this feature. At 
any rate, those examples forwarded from Plymouth by Dr. Allen did not show sufficient 
1 'Mitth. Zool. Stat. Neapel/ xiii, p. 486 (1899). 

3 'Ann. Sc. Nat./ 9 e ser., iii, p. 274, figs. 1, 2, and 3 ; pi. vi, fig. 1-3. 
3 'Ann. Sc. Nat./ 9 e ser., iii, p. 67, 1906. 



60 EITLALIA (PTEROCIRRITS) MACROCEROS. 

distinction in the buccal segment, in the structure of the feet, or other characters, to 
warrant specific separation. The colouration, as described by De St. Joseph, is certainly 
striking, consisting of a brownish ground colour, with two parallel greenish bands on each 
side of a central spot in each segment. The ventral surface is slightly violet. 

The form with fourteen papillse on the proboscis is that found at Dinard by De 
St. Joseph (1888), who considered the following as varieties, viz., Phyllodoce clavigera 
(Aud. & Ed.), Eulalia viridis ((Erst.), P. viridis (Johnston), E. vixens (Ehlers), and E. 
guttata (Claparede) . 

Hornel (1891) thought the green colour was not protective. 

Gravier 1 shows a strong transverse band of muscle passing from side to side 
below the gut and above the nerve-cords in this form. Between it and the outer portion 
of the oblique muscle above it, lies the segmental organ, the internal ciliated funnel 
and the external opening of which (just within the base of the ventral cirrus) are 
also figured. 

A closely allied form is Eulalia aurea (Gravier), 2 which is of a bright golden hue, 
with a violet band along the middle line of the dorsum, and a dark line on each side, or 
rather a kind of arch in each segment, this alone being continued in front. The eyes 
are large and circular; the paired tentacles short, the median longer, and inserted in 
front of the eyes. The first segment bears a pair of tentacular cirri ; the second two 
pairs, with a minute bundle of bristles between. The dorsal (tentacular) cirrus extends to 
the twelfth segment. The third segment bears the fourth pair of tentacular cirri, accom- 
panied by a bundle of bristles and a minute ventral cirrus. The dorsal cirri are elongate, 
with a blunt point. The ventral cirri are a little less, and relatively large. Anal cirri 
are foliaceous, and double the length of the dorsal cirri of the region. The bristles have 
a hook at the end of the shaft which is covered with minute spines. The terminal 
blade is very short, and shows oblique striation. Proboscis is entirely covered with 
minute papillse. The end has a ring of twenty papillse. 

It belongs to the type of E. viridis, with elongate, lancet-like, dorsal cirri. Gravier 
says it is distinguished by its colouration, by the remarkable length of the proboscis, by 
its dorsal cirri being less acuminate, and also by its bristles — the terminal piece of which 
is short. It has not been thought necessary to separate this form from E. viridis. 



4. Eulalia (Pterocierus) macroceros, Grube, 1860. Plate LVIII, fio-. 4 head ■ Plate 

LXVII, fig. 12— foot; Plate LXXVII, fig. 3— bristle. 

Specific Characters. — Head rather large and long, with two large circular black eyes, 
in front of which arises the large, subulate, and long median tentacle. Occasionally a 
dark line occurs behind each. The other tentacles are also comparatively large and long. 
The first pair of tentacular cirri is normal, but somewhat short. The dorsal of the 
second pair is of considerable length, whilst the ventral forms a long, lanceolate, and 

1 'Bull. Sc. Fr. Belg./ t. xxix, pi. xxii, fig. 3. 
3 Op. cit., t. xxix, 1896, p. 309, pi. xvi, fig. 2—6. 



EULALIA (PTEPOCIPRUS) MACROCEROS. 61 

foliaceous process. The dorsal of the third pair is shorter than the preceding, and the 
ventral is small (De St. Joseph). 

Body abont an inch or two in length, slightly tapered anteriorly, and more so 
posteriorly, where the great elongation of the feet give a character to the region. Colour 
straw-yellow to pale-brown, with a dorsal bar going to the bases of the feet. Dorsal 
cirrus lanceolate, with a pointed tip. Setigerous lobe tapered a little towards the bifid 
extremity. Bristles with the end of the shaft slightly dilated and bevelled, with a minute 
spike or two at the tip, and a single oblique line. The somewhat long terminal piece 
tapers to a fine tip, is obliquely striated, and distinctly serrated. The ventral cirrus is 
slipper-shaped, lanceolate at its tip, and extends a little beyond the setigerous region. 



Synonyms. 

] 860. Phyllodoce (Eulalia) macroceros, Grube. Arch, f . Nat., Bd. xxvi, p. 82, Taf . iii, f . 4. 

„ „ „ „ idem. Ausflug nach Triest, p. 141, Taf. iii, f. 4. 

1864. „ „ „ Bhlers. Borst., i, p. 165, Taf. vii, f. 6 -10. 

1868. „ (Pterocirrus) velifera, Claparede. Annel. Neap., p. 250, pi. xvii, f. 2. 

1874. „ „ macroceros, Marenzeller. Sitz. Akad. der Wiss. Wien, Bd. lxix, p. 424, 

sep. abdr., p. 18. 

1875. „ „ „ Marion & Bobretzky. Ann. So. Nat., 6 e ser., t. ii, p. 63. 
„ „ „ velifera, idem. Ibid. 

1888. „ „ „ De St. Joseph. Ibid., 7 ser., t. v, p. 300, pi. xii, f. 170—174. 

1897. Pterocirrus macroceros, Gravier. Bull. Sc. Fr. Belg., t. xxix, p. 308, pi. xvi, f. 15 ; pi. xx, 

f. 12; pi. xxii, f. 8, 9. 

1904. „ „ Allen. Journ. M. B. A., n.s., vol. vii, p. 223. 

1905. Eulalia „ ( — E. volucris, Ehlers, and Eracia volucris, Clap.), Grgeffe. Arbeit. Zool. 

St. Triest., xv, p. 325. 

1906. „ (Pterocirrus) macroceros, De St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 225. 

Habitat. — Dredged off St. Peter Port, Guernsey, in 15 fathoms on shelly ground in 
1868. Plymouth (Dr. Allen). 

Quarnero (Grube). Amongst nullipores and Bryozoa, at Triest (Marenzeller). 
Common in the Gulf of Marseilles (Marion and Bobretzky), Mediterranean, Atlantic. 

Read (Plate LVIII, fig. 4) large and long in life; bluntly conical in spirit, with 
two large circular black eyes, with or without lenses. The subulate tentacles are 
comparatively large and long, the median (the longest) arising in front of the eyes. 
They thus correspond with Claparede's Eulalia velifera, and differ from his E. marginata. 
The first pair of tentacular cirri is normal in outline, but somewhat short. The second 
has dorsally a subulate process of considerable length (extending to twelve segments, 
De St. Joseph), whilst the ventral is foliaceous and lanceolate. This shows no striated 
border as in E. limbata, Claparede, neither is the tentacular differentiation visible at 
the posterior border. The whole forms, in spirit, a leaf-like process, a little more 
opaque towards the posterior border. 

Body of a straw-yellow colour, or with brownish dorsal bars, slightly tapered 



62 EITLALIA (PTEEOCIRRUS) MACROCEROS. 

anteriorly, more distinctly so posteriorly, where the great elongation of the feet and the 
lamellae give a character to the region. Posteriorly it terminates in two thick cylindrical 
cirri (De St. Joseph). 

The proboscis has fine conical papillae at the basal region in extrusion, but they 
increase in size and become coarse towards the tip, which has a series (12 — 15 ?) of 
papillae. Forty-eight large blunt papillae surround the entrance to the stomach (De St. 
Joseph). The last-named author found fragments of Fuci and often gregarines in the 
alimentary canal. 

The foot has a prominent lanceolate dorsal cirrus (Plate LXVII, fig. 12) with a 
pointed extremity. The setigerous process beneath tapers towards the tip, which is 
slightly bifid, the superior division being the more prominent and acute. The bristles 
are pale and comparatively short with a very slight curvature. The distal end of the 
shaft is a little dilated and bevelled, but only shows traces of spikes at the end, the 
translucent region being otherwise structureless, except for a single oblique striation. 
The somewhat long terminal blade tapers rather abruptly to a fine tip, and has delicate 
oblique lines and serrations (Plate LXXVII, fig. 3) which slope distally. The process 
at the tip of the bristle is certainly longer than Claparede shows in his E. limbata. 
Unfortunately, he has figured neither feet nor bristles in the other species. He describes 
the dilated end of the shaft as having asperities, and the terminal piece in his figure l 
is poorly represented. The serrations, though not easily seen, come considerably further 
downward, beyond the split at the articulation, and the tip is longer and more tapered. 
This author gives an interesting account of the mucus-glands and bacilliporous follicles 
of the dorsal and anal cirri. The inferior lamella (cirrus) is also somewhat lanceolate, 
the tip extending a little beyond that of the setigerous region. 

Reproduction. — Marenzeller (1874) found that the females had in August dark bluish 
green eggs — occupying all the segments except a few anteriorly and posteriorly. De 
St. Joseph describes the eggs as green. 

Grube's original description and figure (1860) are, as De St. Joseph observes, fairly 
accurate, and could scarcely refer to any other form, though the reniform outline of the 
eyes has not since been observed. The Eulalia volucris of Ehlers is a synonym for the 
same form according to De St. Joseph, though Ehlers describes the third pair of 
tentacular cirri as having the ventral cirrus leaf -like as well as that in front. Ehlers (1864) 
gives a careful account of the structure of the proboscis, which has about forty short 
papillae at the opening into the stomach. He also describes the leaf -like anal cirri which 
were absent in Grube's example. 

Claparede in 1868 distinguished the group to which this form belongs, charac- 
terizing it by the foliaceous condition of the ventral tentacular cirrus of the second 
segment, and making it a sub-genus under the name of Pterocirrus. 2 He described three 
species, but he had apparently overlooked the description of Grube, so that his Eulalia 
(Pterocirrus) velifera is synonymous with Grube's E. macroceros. It may be questioned, 
indeed, whether the other species — viz. Eulalia (Pterocirrus) limbata and E. marginata — 
have not a closer connection with his velifera than at first appears. Marenzeller points 

1 Op. cit., pi. xii, fig. 170. 
3 < Annel. Nap.,' 248, and Suppl., p. 98. 



EULALIA TRIPUNCTATA. 63 

out that the dark line shown by Grube and Claparede behind each eye is not constant, 
and it was not present in the small example from the Channel Islands (1868), and 
that the ventral leaf of the second pair of tentacular cirri has a spine. 

De St. Joseph, at Dinard, met with comparatively large examples — viz. 5 centimetres 
in length, 6 mm. across the dorsal cirri, and with 116 segments. He found the parasitic 
crustacean, Herpyllobius arcticus (Steenstrup) fixed to the side of one specimen. 



5. Eulalia tripunctata, Mcintosh, 1874. Plate XLIV, fig. 3; Plate LVIII, fig. 23— head; 
Plate LXVII, fig. 13— foot; Plate LXXVII, fig. 4— bristle. 

Specific Characters. — Head bluntly conical, flattened and pale, with four rather 
short tentacles at the tip, and a median between the well-marked black eyes, which are 
posterior in position. The first pair of tentacular cirri occurs on the buccal segment, the 
second segment bears two pairs. The long posterior tentacular cirri are only once and a 
half the breadth of the body. . The latter is about two and a half inches long, narrow, 
gently dilating behind the head, and again diminishing gradually towards the attenuated 
tail, which has two short, baton-like or fusiform cirri, flattened dorsally and ventrally. 
A slender papilla occurs in the centre between them. Of a pale yellow colour, with 
three rows of black spots on the dorsum, the lateral commencing on the first segment and 
behind the attachment of the foot, and the central on the intermediate fold at the segment- 
junction, so that they form a triangle. The ventral surface is pale yellow, with a faint 
lateral row of dark grains at each side in the middle of the body. The dorsal cirrus is 
ovato-lanceolate — probably nearly lanceolate in life. The setigerous lobe is compara- 
tively short, indistinctly bilabiate (De St. Joseph), with short pale bristles, the shafts of 
which in lateral view are bevelled at the enlarged end. The terminal blade is short, and 
rapidly tapers to a point. The serrations on the edge are very minute. The ventral 
cirrus is ovato-lanceolate, the tip extending fully as far as the setigerous process. 

Synonyms. 

1874. Eulalia tripunctata, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 197. 

1875. „ „ idem. Invert, and Fishes, St. Andrews, p. 120. 

1888. „ Claparedii, De St. Joseph. Ann Sc. Nat,, 7 e ser., t. v, p. 285, pi. xi, f. 144 — 148. 
1904. „ „ Allen. Journ. M. B. A., n.s., vol. vii, p. 223. 

Habitat. — From the debris of the deep-sea fishing-boats, St. Andrews, May, 1868 
(R.M.) ; between tide marks, Herm ; dredged amongst dead shells off St. Peter Port, 
Guernsey. 

Plymouth (Allen). Shores of France (De St. Joseph). 

Head, — Pale throughout, flattened, bluntly conical, with four rather short tentacles 
at the tip, and a median between the well-marked black eyes, which are situated 
posteriorly. There are three tentacular cirri, the posterior the longest, yet the latter are 
only once and a half as broad as the body. 

Body about two and a half inches in length, much elongated proportionally, narrow, 



64 EULALIA IMBKICATA. 

gently dilating behind the head, continuing for a long distance of nearly equal calibre, 
and again diminishing gently towards the attenuated tail. The segments are distinctly 
marked, and are flanked by a series of lanceolate lamellae (cirri) of a deeper yellow 
than the rest of the body, and which become ovato-lanceolate in spirit. The tail is 
terminated by two rather short baton-like cirri, which are flattened dorsally and 
ventrally. In preservation the body is rounded dorsally, and slightly flattened 
anteriorly on the ventral surface. In life it is coloured throughout of a pale yellow, 
with three rows of black (or greenish, De St. Joseph) spots on the dorsum of the body. 
The lateral rows commence on the first segment, and the central about the fourth. 
These spots are arranged in a regular manner, the central being on the intermediate 
narrow fold, or pseudo-segment, at the junction, and the lateral just behind the attach- 
ment of the foot on each side, so that they form a triangle. The under- surface is 
pale yellow, without spots anteriorly and posteriorly, but towards the middle it is 
marked with a faint row of dark grains at each side. 

The proboscis is irregularly covered anteriorly by obtusely conical papillae, and has 
a crown of fourteen papillae at the " stomach" (De St. Joseph). 

The foot (Plate LXVII, fig. 13) has dorsally an ovato-lanceolate cirrus, probably 
nearly lanceolate in life. The setigerous process is comparatively short, slightly bifid, 
and bears a series of short, pale bristles. The shaft has only a slight curvature, chiefly 
marked just below the enlarged end, which is rounded when viewed antero-posteriorly, 
bevelled and hoof -like when seen laterally. The terminal process is short, translucent, 
and rapidly tapers to a point (Plate LXXVII, fig. 4). The serrations on the edge are 
so minute that they are observed with difficulty. 

The inferior cirrus is also ovato-lanceolate, the tip extending fully as far as the 
setigerous process. 

The muscular fibres are firm and tough in the preparations, presenting in some the 
appearance of the glandular secretion of Panthalis. 

Reproduction. — De St. Joseph found that the females (in spring?) were distended 
Avith greenish eggs ; whereas the males, when filled with sperms, have the two posterior 
thirds of the body dull reddish. 

This seems to be a widely distributed species, but is not common in the north. 

Dr. Allen kindly sent me a drawing of a specimen — his " Eulalia ? brick-red " — of 
a general pale pink, with rather more than the middle third of the body of a reddish 
hue (Plate XLIII, fig. 8). No specks were visible. Nothing distinctive occurred about 
the head, which resembled that of Ewnida, and the caudal cirri were lanceolate and 
narrow, inclining to subulate, as in Eumida. The precise relationships of this form, 
therefore, are in need of further investigation. 



6. Eulalia tmbeicata, Elders, 1875. Fig. 42, head. 

Specific Characters. — Head somewhat massive, pentagonal, eyeless. Median tentacle 
in the angle at the nuchal border of the head. 



EULALIA (?) IMBRICATA. 



65 



Body flattened, tapered for a short distance anteriorly, and terminating posteriorly 
in two elongate-ovoid caudal cirri. Foot with a large and broadly ovate dorsal cirrus 
(lamella), an elongate-ovoid ventral cirrus, and a bilabiate setigerous region with bristles 
having the ends of the shafts dilated and furnished with two long spines, whilst the 
tapering terminal pieces are serrated (Ehlers). 

Synonym. 
1875. Eulalia imbricata, Ehlers. Zeitsch. f. w. Zool., Bd. xxv ; p. 43, Taf. ii, f. 15. 







Fig. 42. — Head of Eulalia imbricata (after Ehlers). 



Eulalia (?) imbricata, Ehlers, 1875. 

Habitat. — Dredged in the 'Porcupine' Expedition of 1869 in 664 fathoms, on 
July 2nd, at Station 23, 56° 9' K, 14° 10' W. 

Head (fig. 42, after Ehlers) somewhat large, pentagonal, with the greatest breadth 
posteriorly. The four anterior tentacles arise further from the tip than usual, whilst the 
small median tentacle springs from the extreme central process of the head posteriorly. 
No eyes. The head thus differs from the general type of the genus. 

Body about 13 mm. in length, 2 mm. broad, and has seventy segments. It is 
flattened, tapered anteriorly, and yellowish- white in spirit. The first three segments 
bear four pairs of tentacular cirri, the longest reaching from the third to the tenth 
segment. The last segment has two elongate-ovoid anal cirri. 

The average foot has a broadly ovoid dorsal lamella (cirrus). The setigerous 
process is two-lipped, the lower being larger than the upper. The distal end of the shaft 
of the bristles is enlarged and two-spined, whilst the edge of the terminal sabre is 
serrated. The ventral lamella (cirrus) is elongate-ovoid, and extends about the length 
of the setigerous lobe. 

66 



66 EITMIDA SAXGUINEA. 

This interesting species is remarkable for the structure of its head and the foliaceous 
condition of the dorsal cirri. In the position of the median tentacle it differs from all 
other forms in the genus, and further investigation of the type is necessary. 

Besides the foregoing, several varieties of Eulalia occur from Gal way, deep water 
off St. Andrews Bay, and elsewhere. In these the eyes vary considerably in size, some 
being larger, others smaller than in the type, and the dorsal cirri are shorter or longer. 



Genus XXXII. — Eumida, Malmgren, 1865. 

Eumida, Claparede (1868) states, is distinguished by the presence of a small but 
normal lamellar ventral cirrus below the last tentacular cirrus. He would, therefore, for 
the moment make Eumida (Eracia, Be Quatref.) a sub-genus of Eulalia. Five tentacles, 
median on the vertex; four pairs of tentacular cirri. First segment soldered to the 
head-lobe (Levinsen), and first pair of tentacular cirri spring from it. The arrangement of 
the dorsal and ventral longitudinal muscles and other features as in Phyllodoee and Eulalia. 



1. Eumida sanguinea, (Ersted, 1843. Plate XLIII, figs. 9 and 10; Plate XLIV, fig. 4; 
Plate XL VIII, figs. 1 and 2 ; Plate LVIII, fig. 22— head ; Plate LXVII, figs. 14 
and 15— feet; Plate LXXVII, fig. 5— bristle. 

Specific Characters. — Head rounded, cordate, broader than long, the large black or 
brownish-black eyes thus being prominent. Anterior tentacles of moderate length, 
subulate; median, somewhat shorter, arising in front of the eyes. Tentacular cirri of 
moderate length, the dorsal in each pair being the longer, and all taper from the base to 
the filiform tip, which in the posterior pair is finely attenuate ; the ventral of the second 
pair is slightly flattened and approaches a narrow lanceolate form. Body one and a half 
to three inches in length, of considerable proportional bulk, soft, and mobile ; somewhat 
narrowed in front, and tapered posteriorly, where it ends in two fusiform or subulate 
styles. Colour light greenish-brown, yellowish, straw-coloured, or almost translucent 
anteriorly, with a faint straw-coloured line along the centre and a dusting of brownish 
grains. The first segment has a white bar or white grains. In many a narrow white 
or yellowish bar occurs at each segment-junction. In spirit it is generally madder 
brown. Proboscis long, transversely rugose behind the tip, which has twenty papilla? 
(Malmgren). In the typical foot (sixtieth) the dorsal cirri are sub -vertical, broadly 
ovate, with an acuminate tip, while the base is sub-cordate. It is borne on a long 
pedicle (ceratophore). Spinigerous process comparatively long and bifid. The translucent 
shafts of the bristles have a distinct shoulder, a bevelled tip with spinigerous ridges, 
and a minutely serrated terminal blade, longer than in the ordinary Eulalia, and tapering 
to a fine tip. The ventral cirrus is broadly lanceolate, with a somewhat acuminate tip, 
which does not quite reach that of the spinigerous lobe, and thus is shorter than in the 
tenth foot. 



EUMIDA SANGUINEA. 67 



Synonyms. 



)} 


Eulalia 


,, 


1867. 


?J 


>> 


1873. 


}) 




1874. 


Eumida 


a 


)} 


}) 


}} 


1875. 


}) 


>) 


)> 


Eulalia 


>> 


1879. 


)) 


jj 


1883. 


Eumida 


;y 



1843. Eulalia sanguinea, (Ersted. Annul. Danic Consp., p. 28, f. 80 — 82. 
1851. Phyllodoce (Eulalia) sanguinea, Grube. Fam. Annel., pp. 56 and 129. 
1865 Eumida sanguinea, Malmgren. Nord. Hafs- Annul., p. 97, Tab. xiv, f. 28. 

De Quatrefages. Annel., ii, p. 123. 

Malmgrem. Annul. Polychaet., p. 25. 

Kupffer. Jahresb. Com. deutsch. (' Pommerania'), p. 150. 

Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 196. 

Malm. Op. cit. Goteb., p. 80. 

Mcintosh. Invert, and Fishes St. Andrews, p. 119. 

Mobius. Jahresb. Com. deutsch. ('Pommerania'), p. 170. 

Tauber. Annul. Danic, p. 88. 

Levinsen. Vid.-Med. Foren. Kjoben., p. 205. 
1888. Eulalia pallida, De St. Joseph. Ann. Sc. Nat., 7 e ser., t. v, p. 294. 
1893. „ „ Levinsen. Vidensk. Ud. ' Hauchs/ p. 327. 

1897. Eumida communis, Grravier. Bull. Sc. Fr. Belg., t. xxix, p. 310, pi. xvi, f. 7 — 10. 
1901. „ sanguinea, Mcintosh. Ann. Nat. Hist., ser. 7, vol. viii, p. 223. 

1904. Eulalia pallida, Allen. Journ. Mar. Biol. Assoc, n.s., vol. vii, p. 223. 

1905. „ „ Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 325. 

1906. „ „ De St. Joseph. Ann. Sc Nat., 9 e ser., t. iii, p. 224. 

Habitat. — Under stones near low water-mark amongst the rocks, and from the debris 
of the lines from deep water, St. Andrews (E.M.) ; Outer Hebrides (Lochmaddy), both 
between tide-marks and in the Laminarian region, and Sound of Harris ; on shells from 
deep water off the Channel Islands ; as well as between layers of rock near low- water 
mark (W. C. M.). In 80 fathoms, St. Magnus Bay, Shetland (J. G. J.). On greyish 
sand, stones, and coral at 164 fathoms in the ' Porcupine ' Expedition of 1869. A species 
abundant in both tidal and deep waters all round the British shores. 

It extends to the Baltic, Norway, and Finmark. 

Head (Plate XLIII, fig. 9) rounded, cordate, broader than long ; pale, or with a slightly 
pinkish hue, straw-coloured or greenish, the large black or brownish black eyes thus being 
prominent, and they apparently enlarge in ripe examples. Anterior tentacles of moderate 
length and subulate ; the median, somewhat shorter, arises in front of the eyes. The 
tentacular cirri are also of moderate length, the dorsal in each pair being the longer, 
and all taper from the base to the filiform tip, which in the posterior pair is more finely 
attenuate. The ventral of the second pair is broader and slightly flattened, approaching 
a narrow, lanceolate form. As in Eulalia the first pair springs from what (Ersted termed 
the base of the head, but which probably represents a fused peristomial region, the 
second and third from the following segment, which is united with the former, and the 
fourth from the succeeding segment, a rudimentary foot (as in Eulalia viridis) being 
beneath it. In some a pale area intervenes between the eyes, and occasionally a few 
white grains are found in the same region. In several from St. Peter Port, Guernsey, 
the tentacular cirri, opposite the white post-cephalic belt, were white. 

Body 1^ to 2-| inches in length, of considerable proportional bulk, soft and 



68 EUMIDA SANGUINEA. 

mobile. It is rounded dorsally, flattened ventrally, somewhat narrowed in front, and 
tapered posteriorly, where it terminates in two fusiform or subulate cirri. In colour it is 
light greenish brown, or straw-colour, occasionally almost translucent anteriorly, with a 
faint, straw-coloured, or yellowish line along the centre, greenish white, or pale brownish, 
with a dusting of brownish grains. The posterior region is more opaque. The first 
segment is marked by a pure white bar, or by white grains, and in many a narrow white 
or yellowish bar occurs at each segment-junction. In one from St. Peter Port, 
Guernsey, the body was of a fine light purple or mauve hue, with a white silvery bar 
across each segment and whitish specks between the eyes. Some are pale greenish 
brown, faintly marked by transverse lines on the dorsum and by a dull yellowish or 
brownish line in the centre from the intestine, while a variety is very prettily banded with 
dark green in a large bar anteriorly in each segment and a smaller behind, the effect 
being heightened by the pallor of the lamellae and the tentacles. The ventral surface is 
pale or marked by the salmon-tint of the viscera. 

In section the oblique muscles pass below the nerve-cords on each side, and meet 
at their insertion in the middle line. The arrangement of the dorsal and ventral 
longitudinal muscles is typical. 

Proboscis (Plate LVIII, fig. 22) long, transversely rugose behind the tip, which has 
twenty papillae. 

It is seldom that the organ is everted by the mouth in the preparations. It is 
generally detached anteriorly and thrust out by a rupture of the body-wall. 

The coelomic fluid is mixed with many large flattened cells (August) measuring fully 
5~Jq of an inch in their longer diameter, while the ordinary corpuscles are less than -^too 
of an inch. 

The wall of the alimentary canal is richly glandular. 

Dr. Allen forwarded a drawing of a form from Plymouth, which he termed Eulalia 
sp. B, of a pale greenish hue anteriorly, slightly pinkish posteriorly, the general aspect 
agreeing with that of Eumida, the white bar behind the head being conspicuous. In the 
coloured drawing, however, the median tentacle is considerably longer than the anterior 
or paired tentacles — a feature less marked in the preserved form with the proboscis 
extruded. The shape of the foot and the general aspect of the proboscis appear to agree 
with Eumida sanguined, of which this appears to be a fairly characteristic example. 

In the typical foot (Plate LXVII, figs. 14a — tenth, and 15 — sixtieth) the dorsal cirri 
are borne subvertically, and are broadly ovate with an acuminate tip, while the base is sub- 
cordate, and the hypodermic granules are arranged in very regular and close streaks, so 
that the appearance is characteristic. The streaks proceed from a central region more 
opaque and areolar than the marginal. The papilla bearing the process is elevated, 
so that the space between the lamella and the spinigerous lobe is marked. The 
spinigerous process is comparatively long, and bifid, and the inferior broadly lanceolate 
cirrus arises from a distinct shoulder inferiorly and posteriorly. Its tip is also somewhat 
acuminate, and does not quite reach that of the spinigerous region, and therefore is 
proportionally shorter than at the tenth foot. When seen antero-posteriorly a single 
large median spine occupies the centre, and a series of smaller spines trend off on each 
side. The translucent bristles have a distinct shoulder and a spinigerous bevelled tip 



EUMIDA SANGUINEA. 69 

in lateral view (Plate LXXVII, fig. 5), in which position the very slight curvature of 
the tip of the shaft is visible. The terminal blade is longer than in the ordinary 
examples of Eulalia, tapers to a fine tip, and is minutely serrated along the edge. 

Coiled vessels occur at the base of the dorsal cirrus, but none have been traced into 
it. Cilia seemed to be arranged along the basal part of the foot. 

Reproduction. — Ripe ova occur towards the end of May, and it is probable that the 
breeding period extends to August. Ripe males came from four fathoms in Valencia 
Harbour on May loth, 1870 (J. Gr. J.), and from Lochmaddy in August. A ripe 
female occurred in August under the spreading crust of Gorallina officinalis at Herm. 
Beyond the enlargement of the body no change of external appearance takes place in this 
condition, but in section the whole coelomic space and the cavities of the feet in the males 
are distended with sperms, and the wall of the gut appears to be thinner. In the females 
the early ova occur in masses in the feet. Allen observes that at Plymouth it (his 
Eulalia pallida, and also his greenish Eulalia B, with white bar) breeds from May to 
July, and that the eggs are green. 

Young examples of about sixteen segments were procured in the bottom tow-net 
on June 16th, and others of seventeen bristled segments on October 22nd. A white 
band occurs behind the head, the rest of the body being of a pale straw-colour. Eyes 
comparatively large, black. Some of the young forms at a later stage, from deep water, 
are of a madder-brown colour. 

Habits. — The body is very mobile, constrictions taking place at various points 
(Plate XLIII, fig. 9) and again disappearing. It is an active and restless species, moving 
backward and forward, or in confinement swimming with considerable speed in a 
serpentine manner at the surface or through the water of a vessel, and evading capture 
with adroitness. A vermicular contraction and dilatation often occur posteriorly — as if 
water entered the rectum. The vitality of fragments, e. g. the posterior end, is great. 

It is difficult to say whether the Nereis ellipsis of Dalyell 1 is this form or a Eulalia. 

The Eulalia punctifera, Grube, 2 from Cherso in the Adriatic, approaches this form 
very closely. 

De Quatrefages 3 (1843) described and figured a Phyllodoce saxicola which had five 
tentacles, and which accordingly in the 6 Anneles ' was termed Eulalia saxicola — a form 
which appears to be one of the varieties of Eumida sanguinea. 

It is doubtful if such species as Claparede's Eulalia (Eumida) pallida 4 merit specific 
distinction. The form mentioned would appear to be a variety. His Eulalia (Eumida) 
guttata ° also is a closely allied form. Eumida sanguinea is variable in colour, and the 
pale examples from Plymouth (=Eulalia pallida, Allen) differ in no material way from 
the type (which is absent from the list of the local annelids). De St. Joseph found a 
female parasitic copepod on this form at Dinard. 

1 < Pow. Creat./ ii, p. 152, pi. xx, figs. 7—10, 1853. 
3 ' Archiv f. Naturges./ 1860, p. 83. 

3 f Magasin de Zoologie/ 1843, p. 1 (sep. copy), pi. i, figs. 1 — 6. 

4 ' Annel. Nap./ p. 246, pi. xvi, fig. 6, 1868. 

5 Ibid., SuppL, p. 97, pi. ix, fig. 2. 



70 ANAITIS. 

The differences of Webster's Eumida maculosa, 1 dredged on shells in 5 — 10 fathoms 
off the Virginian coast, seem to be very slight — indeed only of coloration. 

A variety with a larger ventral cirrus was procured in Gal way in 1868 by Professor 
B. P. Wright, of Dublin, the main feature being the size of the lanceolate ventral cirrus, 
the pointed tip of which projected considerably beyond the setigerous process. The 
minute structure of the bristles agreed with that in E. sanguinea. Malmgren gives Sige 
fusigera a lanceolate and pointed ventral cirrus like the foregoing, but there is no reason 
for separation in the present case. 

The points of distinction claimed by Gravier (1897) for his Eumida communis, viz., 
coloration, rounded form of the dorsal cirri, great development of the setigerous lobe 
next the ventral cirrus, and the condition of the bristles, do not seem to form stable 
grounds on which to base separation from E. sanguinea. 

The foot of Eidalia (Ptero cirrus) microcephala, Claparede, 2 as subsequently shown 
by De St. Joseph, 8 would readily be taken for that of Eumida sanguinea or one of 
its many varieties. 

Several varieties of Eumida sanguinea occur in the deeper water of St. Andrews 
Bay, the eyes being larger in one, the setigerous lobe of the foot more prominent in 
others, the dorsal cirri smaller than usual and the ventral varying in length, but the 
median tentacle remains shorter than the lateral and the bristles do not differ from the 
type. One specially soft form shows slight differences in the structure of the end of the 
shaft of the bristles, which are more abruptly truncated. 



Genus XXXIII. — Anaitis, Malmgren, 1865. 

Head broad, rounded anteriorly, slightly sinuous posteriorly, somewhat indistinctly 
separated from the first segment. Four short tentacles. Two eyes of moderate size. 
Four tentacular cirri on each side, the first, second, and third fixed to the first segment, 
the fourth to the second segment. Superior cirri of the feet large and imbricate back- 
wards. Ventral cirri compressed, a little longer than the setigerous part of the foot. 
The latter is ovate, diminishing at the tip, and slightly bifid. Bristles numerous, 
compound, terminal sabre long and tapered, minutely serrated on the edge. Two 
sub -globose anal cirri. 

I have some hesitation in adopting this genus — the species included in it requiring 
more exhaustive examination. 

Grube (1879) described the posterior border of the head in Anaitis as arising from a 
median arch, whilst Levinsen thought it was soldered with the first segment. There are, 
it is stated, three pairs of tentacular cirri. Grube places Phyllodoce Paretti, Blv., here. 
Anaitis rosea, McL, he ranges under the group b 2 , in which segment 2 is smaller. 

Claparede (1868) notes that the essential distinction between this and Phyllodoce is 

1 ' Trans. Albany Instit./ ix, p. 15, pi. iv, figs. 38—41, 1879. 

2 ' Annel. Nap., Suppl./ 1870, p. 98, pi. ix, fig. 3. 

3 c Ann. Sc. Nat./ 9 e ser., t. iii, p. 225, pi. v, figs. 99—103, 1906. 



ANAITIS ROSEA. 71 

the occurrence of the four pairs of tentacular cirri on three segments in the latter and on 
two in the former. He, however, observes that it is still uncertain whether the buccal 
segment does not represent two fused segments. In some a minute ventral cirrus occurs 
under the last pair of tentacular cirri (e.g. Anaitis cephalotes, Claparede). 



1. Anaitis rosea, Mcintosh, 1877. Plate XLIII, fig. 4; Plate LXVIII, fig. 13— foot; 

PL LXXVII, fig. 10— bristle. 

Specific Characters. — Head somewhat ovoid, slightly tapered in front, little diffe- 
rentiated from the first segment. Eyes comparatively small; situated at the posterior 
third. In front of them is a bright rose-red band slanting downward and backward, and 
curving to join the larger rose-red area. A pinkish band from the tip of the snout joins 
the latter behind the eyes, thus isolating a pale area around each eye. A broad belt of 
rose-red occurs in rear of the eyes, and a pale wedge impinges behind into its centre. 
Tentacular cirri as in Phyllodoce. Body tapered in front, and more distinctly so 
posteriorly, rose-red dorsally, marked along the centre by a yellow band which is flecked 
with reddish grains. Surface generally is pale buff. Foot with a prominent pedicle 
bearing a lanceolate dorsal cirrus, and a slightly bifid setigerous process. The enlarged 
ends of the gently curved bristles are spinulose along the terminal curves, and the blade 
is elongate, tapering, and finely serrated. The ventral cirri are lanceolate, acutely 
pointed, and project beyond the tip of the bristle-bearing process. Tail terminated by 
two short curved cirri. 

Synonym. 
1877. Anaitis rosea, Mcintosh. Journ. Linn. Soc, xiii, p. 161. 

Habitat. — In the sand near low-water mark, East Sands, St. Andrews, September, 
1876. In 8 fathoms amongst sand in the Bight of Vatsland, north of Bressay Sound, 
Shetland. 

Head. — Somewhat ovoid in outline, slightly tapered in front, while posteriorly it is 
hardly differentiated from the succeeding segment. The two circular eyes are situated 
at the posterior third and are comparatively small. It is pale anteriorly as likewise are 
the tentacles, which are subulate and tapering. In front of each eye is a bright rose-red 
band, slanting downward and backward and then curving to join the larger rose-red area. 
Moreover a pinkish band from the tip of the snout joins the latter behind the eyes, thus 
mapping off the pale area around each eye. A broad belt of rose-red occurs in rear of 
the eyes, into the centre of which impinges a pale wedge from behind, so that the lateral 
roseate regions as seen from the dorsum are somewhat triangular. The same rose-red 
hue tints the bases of the tentacular cirri, which have the normal arrangement, viz. two 
shorter in front and two longer behind. 

Body 1\ inches long, somewhat tapered in front and more distinctly so posteriorly, 
of a fine rose-red dorsally, marked along the centre by a yellow band — from the fourth 
segment to the tip of the tail. For the first three segments the band is flecked with 
reddish grains, but thereafter it is bright yellow. On each side the surface is minutely 



72 ANAITIS KOSTERIENSIS. 

dotted with red grains, and throughout these have a tendency to be arranged in transverse 
bars or streaks. The general tone of the rest of the body and of the lamellas is pale buff. 
Besides the latter hue the ventral surface shows a rose-red band from the line of the first 
tentacular cirrus to that of the first foot, so that when the animal rests on the side of the 
glass vessel it closely resembles the abundant Nemertean Amphiporus lactifioreus, Johnston 
— with its reddish ganglia. 

The superior division of the foot (Plate LXVIII, fig. 10) has a prominent pedicle 
carrying a lanceolate cirrus which thus projects from the body. 

The inferior division is slightly emarginate, and has a spine and a series of pale 
bristles with rather slender shafts and enlarged distal ends — spinulose along the terminal 
curves (Plate LXXVII, fig. 10). The terminal tapering blade is elongate and finely 
serrated along the edge, and the stria3 on it slope from the point and edge downward 
and backward. The ventral cirri are also lanceolate, but much more acutely pointed, 
and project beyond the tip of the bristle-bearing process. 

That so beautiful a species should for a considerable part of its life inhabit the sand 
is one of those curious features so difficult to explain on any modern theory of coloration. 

The Anaitis lineata of Claparede l (1870) from Naples has certain resemblances to 
this species in general structure if not in coloration. 



2. Anaitis kosteeiensis, Malmgren, 1867. Plate LVIII, fig. 6 — head; Plate LXVIII, 
figs. 14 and 15— feet; Plate LXXVII, fig. 11— bristle. 

Specific Characters. — Head somewhat rounded in spirit; eyes of considerable size. 
Tentacular cirri comparatively short and with finely tapered tips. Body apparently rather 
short and broad as well as soft, tapered more posteriorly than anteriorly. Foot with a 
broad and massive pedicle which bears the cordate and foliaceous cirri. The setigerous 
region forms a short cone with a bifid tip. Bristles with a comparatively small distal 
enlargement, which is split by the insertion of the terminal blade — with a spine on each 
side. The ventral cirrus is elliptico-subacuminate, and its tip projects considerably 
beyond the spinigerous regions. 

Synonyms. 
1867. Anaitis ~kosteriensis , Malmgren. Annul. Polych., p. 142. 
1869. „ „ (?) Mcintosh. Rep. Brit. Assoc. (1868), p. 337. 

1874. „ „ Malm. Op. cit., G-oteb., p. 79. 

Habitat. — Dredged during the 'Porcupine' Expedition of 1869 off Bundoran in 
Donegal Bay in 20 — 35 fathoms. 

Malmgren's sole specimen was procured from the Island of Koster, Sweden. 

Head (Plate LVIII, fig. 6) somewhat rounded in spirit, with two eyes of consider- 
able -size. Tentacles absent. Tentacular cirri comparatively short and with finely 
tapered, almost capillary, tips, the anterior apparently having the usual proportion to 
the posterior. 

1 c Annel. Nap., Suppl./ p. 94, pi. ix, f. 4. 



ANAITIS JEFFKEYSII. ^ 

Body about 2 inches in length, apparently rather short and broad as . well as soft. 
The dorsum is almost bare, for the large dorsal cirri do not much encroach on it. The 
tail is absent, but the body tapers much more in this region than anteriorly, though, as 
already indicated, the proportional breadth throughout is considerable. The bristles 
freely project on each side beyond the lamella?. The segments have a prominent fold 
dorsally at their posterior border, a pit on each side at the base of the lamellae indicating 
the commencement of the succeeding ring. 

The foot (Plate LXVIII, figs. 14 [10 th ] and 15 [35 th ] ) shows a broad and massive 
dorsal division which is directed upward and bears the large, foliaceous, and somewhat 
cordate cirrus which is more or less oblique in position — that is, is not quite horizontal. 

The inferior division forms a short cone with a bifid tip and a pale spine. The 
translucent bristles (Plate LXXVII, fig. 11) are of considerable length, the shaft being 
slightly curved and the distal enlargement comparatively small and of peculiar formation. 
The front of the shaft ends in a somewhat conical process with a bifid tip. The pro- 
cesses slope from behind to the serrated edge of the blade. The posterior border again 
shows a marked step at the base of the cone, and from this the long and very finely 
attenuate distal blade takes its origin (as if articulated), its front edge, which is finely 
serrated, trending from the spinous tip of the cone. When the terminal blade snaps, the 
distal end of the shaft thus presents a somewhat trifid appearance. The structure of 
such a bristle differs materially from that in Anaitis rosea, and shows with other features 
the necessity for the relegation of that species to another genus. Malmgren's artist had 
caught the peculiarity, though somewhat indistinctly. 

The inferior cirrus is more or less elliptico-subacuminate, and its tip projects 
considerably beyond the bifid apex of the setigerous region. 

Claparede's Anaitis pusilla 1 (1870) is an allied form. 

3. Anaitis Jeffreysii, 3 n.s. Plate LVIII, fig. 7; Plate LXVIII, figs. 16 and 17— feet; 

Plate LXXVII, fig. 12— bristle. 

Specific Characters. — Head ovoid, narrowed anteriorly, and cordate posteriorly ; two 
eyes near the posterior border. Tentacles subulate with delicately tapered tips, and the 
second pair of tentacular cirri is unusually long and also finely tapered. Body iridescent. 
Dorsal cirrus large and irregularly renif orm ; setigerous process elongate and bifid. Shafts 
of the bristles have dilated ends with spinous curves on each side of the base of the long 
finely tapered and serrated terminal blade. Ventral cirrus more or less lanceolate. 

Habitat. — Dredged off Valencia Harbour, west coast of Ireland, May 31st, 1870 
(J. G-. J.). 

Head (Plate LVIII, fig. 7) somewhat ovoid, narrowed anteriorly, cordate posteriorly, 
and with two moderately large eyes towards its posterior border. Tentacles subulate 
with delicately tapered tips. The second pair of tentacular cirri appears to be more 
slender and elongated than usual, and the tips are finely tapered. 

1 ' Annel. Nap., Suppl./ p. 96, pi. ix, f . 5. 

2 Named after my old friend and veteran explorer of the British seas. The type represented by 
Dr. Grwyn Jeffreys is now almost extinct. 

67 



74 PHYLLODOGE. 

Body linear, rounded dorsally, and flattened ventrally — both surfaces being iridescent. 
The ventral surface has a broad median groove which has a slightly monilif orm appearance 
from the depressions at each segment- junction. The large dorsal cirri cover a con- 
siderable portion of the lateral regions, notwithstanding the length of the feet 
posteriorly. 

The dorsal division of the foot (Plate LXVIII, figs. 16 [10 th ] and 17 [40 th ] ) is short 
anteriorly, but is considerably longer posteriorly. It bears a large irregularly reniform 
cirrus, the hilus of which is situated nearer the lower than the upper border, and which, 
as a transparent object, presents a radiated arrangement of its hypoderm from the 
region of the hilus. These organs appear to be borne obliquely, and overlap each other 
along the sides of the body. 

The tip of the inferior division presents an elongated and sharply conical process 
superiorly, then a dimple, and a slightly receding margin inferiorly. The spine seems to 
issue just beneath the conical process. The shafts of the bristles (Plate LXXVII, fig. 12) 
have a slight curvature with a terminal dilatation, the distal edges of which have minute 
spines, and they slope downward to a step from which the long, finely tapered, and 
distinctly serrated blade springs. The proximal end of the latter is thus bevelled. The 
inferior cirrus resembles the greater part of a lanceolate leaf, with an acute tip, but one 
edge is attached. The tip is more acute in the posterior part of the body, as likewise is 
the superior conical process. 

In the Arteritis peremptoria of Claparede, 1 from Naples, the ventral of the first pair of 
tentacular cirri is very long, the dorsal and the succeeding pairs being shorter. 



Genus XXXLV.—Phyllodoce, Savigny (1820), Char, emend. 

Head elongated, often longer than broad, oval or egg-shaped, or cordate posteriorly. 
One pair of circular eyes. Occasionally two rudimentary nuchal organs; four short 
tentacles, four pairs of tentacular cirri, which Pruvot considers as modified dorsal cirri 
of the ordinary kind, the first pair under the cephalic lobe, second and third on the 
peristomial segment, fourth on the second segment (Malmgren). Body elongated, 
flattened; segments two-ringed; dorsal cirri large, lamelliform, sometimes almost 
rectangular, carried vertically; ventral cirrus much less, fixed to a horizontal pinna. 

Distinguished from Meone by the number of the tentacular cirri, from Eulalia by 
the absence of the fifth tentacle, and from both by the form of the cirri and their 
insertion (CErsted), whilst Anaitis is closely allied. 

Vascular system consists of two longitudinal vessels which communicate with each 
other anteriorly, and which debouch into the posterior part of the body-cavity 
(Gravier). 

Segmental organs.— A pair of ciliated tubes in each segment, the trumpet-shaped 
atrial opening being on the anterior face of each dissepiment, the external aperture 
on the ventral face at the base of the parapodium. Goodrich has recently described 

1 < Annel. Nap., Suppl./ p. 95, f. 6, 1870. 



PHYLLODOCE. 75 

the ciliated genital funnel as open in front and closed behind, and intimately con- 
nected with the segmental organ, which is closed internally, branched, and furnished 
with solenocytes. 1 At maturity an opening is found at the point of fusion of the two 
structures, so that the ripe products escape ventrally. 

Nervous system. — Cephalic ganglia with antero-superior and postero-inferior lobes, 
the latter, the antennary centre, most bulky. The antero-superior centres give origin 
to the stomato-gastric which receive an increment from the oesophageal connectives. 
Ventral chain with fused ganglia, the connecting trunks being separate, and the first pair 
larger than the others, and apparently formed by the fusion of three. 

Reproductive organs developed from ccelomic epithelium, and shed into the perivisceral 
space. Goodrich and Jourdan think that the ova escape by the segmental pores. 

Gravier 2 describes foot-glancls debouching near the segmental aperture ventrally. 

Ehlers (1 868) characterises Phyllodoce, Savigny, thus : " Body elongated, generally 
flattened, distinctly segmented ; head with four tentacles, the first two segments with four 
pairs of tentacular cirri, and often with a rudimentary foot ; the remaining segments 
distinct, with simple feet bearing a fan-like group of jointed bristles and leaf -like dorsal 
and ventral cirri." 

Grube (1879) made two groups of Phyllodoce, viz. : (1) those with the head more or 
less heart-shaped, having a deep dimple posteriorly (P. laminosa and P. groenlandica) ; 
and (2) those with the head heart-shaped or rounded, but with the posterior border only 
a little notched (P. maculata and P. citrina). 

De St. Joseph (1888), after a critical examination of the classification of the 
" Phyllodoces," adopts the following genera : (1) Genetyllis, Malmgren. Achetous buccal 
segment with four pairs of tentacular cirri ; (2) Phyllodoce sans str., Czerniavsky. 
Achetous buccal segment with two pairs of tentacular cirri, other two pairs in the following 
setigerous segment; (3) Anaitis, Malmgren, nee Claparede. Buccal segment achetous, with 
three pairs of tentacular cirri, second segment with one pair, and with bristles; (4) 
Garobia, De Quatrefages, Marenzeller, rev. {Anaitis, Clap., Phyllodoce, Mgrm., Anaitides, 
Czerniavsky). Achetous buccal segment with one pair of tentacular cirri, fused or not 
with the following segment, which has two pairs of tentacular cirri and bristles ; or the 
achetous buccal segment with two pairs of tentacular cirri, fused or not with the following 
segment, which has one pair of tentacular cirri and bristles ; third segment with one pair 
of tentacular cirri, a setigerous lobe more or less developed, and a foliaceous ventral cirrus. 

Cunningham 3 says: "In Phyllodoce no well-marked canals can be distinguished. 
The (nerve) cords are widely separated from the epidermis." 

Leschke 4 procured trochophores of Phyllodocidse in the Bay of Kiel in July, and 
older forms in September, October, and November. 

1 c Q. Journ. Micros. Sci/ vol. 43, n.s., p. 706. 

2 An interesting account of the general structure of the group is given by the author in his 
thesis, "Kecherches sur les Phyllodociens," Lille, 1896. 

3 Op. cit., p. 270, 1888. 

4 ' Wiss. Meer. Biol. Anstalt, Helgol./ n.f., 7 Bd., p. 117, 1903. 



76 PHYLLOD'OCE LAMELLIGERA. 

1. Phyllodoce lamelligera, Gmelin sec. Pallas, 1788. Plate XLV; Plate XL VII, fig. 2; 
Plate XLIX, figs. 1 and 2 variety; Plate LVIII, fig. 20— head; Plate LXVIII, 
figs. i_3_f eet; Plate LXXVIII, fig. 6— bristle. 

Specific characters. — Head somewhat conical, bluntly rounded anteriorly, and cordate 
at the base from two projecting processes. The ground colour is olive green, with a black 
touch in front, and shows pinkish iridescence. Eyes two, large, placed about the middle. 
Tentacles subulate and short ; tentacular cirri with a ringed basal region (ceratophore), 
three pairs on the first segment and a rudimentary ventral cirrus on the second. 
(De St. Joseph says : " First pair under the head on buccal segment ; second and third 
pairs on the second segment ; fourth pair on third segment." Pruvot gives a 
similar arrangement.) Both appendages have the same dull, greenish hue as the dorsal 
cirri. A papilla or boss occurs at each side posteriorly. Body linear, much elongated 
(14 — 18 in.), convex dorsally and flattened ventrally, tapered a little anteriorly 
and more distinctly so posteriorly, where it ends in two rather stout caudal cirri. 
Segments transversely ringed, shorter and broader anteriorly than posteriorly. The 
cuticle is remarkably iridescent, and there is a median series of blackish specks, which, 
with the dull greenish lamellae, make three touches in each segment. A conspicuous dark 
belt on the upper and anterior edge of the pedicle of the foot. Ventral surface with 
dusky pigment and three rows of specks, the median being pale. The proboscis in 
extrusion is long, dark, and clavate, the basal section being covered with small papillae, 
while the distal part has numerous low transverse papillae, and the terminal, firm disc has 
about twenty-one papillae. The dorsal cirri anteriorly are reniform or ovate, at the 
thirtieth elongate-ovoid, at the sixtieth foot the npper part is wide, so that the whole 
process forms a broad flap. Posteriorly, they again diminish so as to resemble an acumi- 
nate leaf. A ciliated ridge behind the dorsal border of the lamella. Setigerous process 
somewhat clavate, Avith a bifid tip. Bristles pale, slightly curved at the distal end of the 
shaft which has minute spines on the terminal ridges and over the surface. The terminal 
blade is of moderate length, and has its edge boldly serrated and with striae sloping 
dorsalward and backward. The ventral lamella is almost reniform in front, becomes 
ovato-acuminate behind and longer than the setigerous region; anal cirri subulate. 

Proboscis with six rows of papillae at the base in extrusion, and sixteen to twenty-one 
papillae at the opening into the stomach. 

Cerebral mass bilobate with a distinct ophthalmic lobe, and the oesophageal connectives 
give off the stomato-gastric nerves. The ventral chain has more or less fused ganglia, 
but the cords are distinct (Pruvot). 

Synonyms. 
1767. Nereis maculata, Linnseus. Syst. Nat., ed. 12, i, pt. 2, p. 1086. 
1788. „ lamellifera, Pallas. Nova Acta Petrop., ii, p. 232, tab. v, figs. 11 — 17. 
1791. „ lamelligera, Grmelin. Linn. Syst. Nat., ed. 13, i, pt. 6, p. 3120. 
1797. „ „ Shaw and Nodder. Naturalist's Miscell., ix, tab. 311 (fide aut.). 

1806. „ „ Turton. Linn. Syst. Nat., iv, p. 90. 

1807. „ „ idem. Brit. Fauna, p. 135. 



PHYLLODOCE LAMELLIGERA. 77 

1808. Nereis foliosa, Montagu. MS. vol. Linn. Soc, pi. i., fig. 1. 
„ „ ,, Leach. Coll. Brit. Mus. (fide Johnston). 

1811. „ lamelligera, Jameson. Mem. Wern. Soc, i, p. 557. 

1812. „ „ Pennant. Brit. Zool., iv, p. 96. 
1818. Phyllodoce laminosa, Lamarck. Anim. s. Vert., v, p. 317. 
1820. „ „ Savigny. Syst. Annel., p. 43. 

1825. Nereis lamelligera, De Blainville. Diet. Sc. Nat., xxxiv, p. 444. 

1826. Phyllodoce lamellosa, Kisso. Hist. nat. Europ., iv, p. 419 (?). 

1828. Nereiphylla laminosa, De Blainville. Diet. Sc. Nat., lvii, p. 467. 

1829. Phyllodoce gigantea, Johnston. Zool. Journ., iv, p. 53. 

1830. „ laminosa, Bosc. Vers, i, p. 173. 

1833. „ „ Audouin and Edwards. Ann. Sc. Nat., t. 29, p. 244, pi. xvi, figs. 1 — 8. 

1834. „ „ idem. Annel. (Lit. France, ii), p. 222, pi. 5a, f. 1—8. 

1840. „ lamelligera, Johnston. Ann. Nat. Hist., iv, p. 225, pi. 6, f. 1 — 6. and xiii, 

p. 438. 

1843. „ laminosa, Bathke. Beitr. Fauna Norweg., Nov. Act., xx, p. 169 (?). 

1844. „ lamelligera, Thompson. Report on the Fauna of Ireland (Bept. Brit. Assoc. 1843), 

p. 273. 
1849. „ laminosa, (M. Ed.) Cuvier. Reg. Anim. Illust., pi. xiii, fig. 2. 

1851. „ „ Grrube. Fam. Annel., pp. 55, 129. 

1853. Nereis remex (Phyllodoce laminosa), Dalyell. Pow. Creat., ii, p. 148, pi. 23, f. 1 — 7. 

1864. Phyllodoce laminosa, Grrube. Insel Lussin, p. 81. 

1865. „ lamelligera, Johnston. Cat. Brit. Mus., p. 175, pi. xvi, f. 1 — 6, and p. 342. 
„ „ laminosa, De Quatrefages. Annel., ii., p. 133. 

1867. „ „ Malmgren. Annul. Polych., p. 24, Tab. iii, fig. 17. 

„ „ lamelligera, Parfitt. Trans. Devon. Assoc, ii, pt. 1, p. 233. 

1874. „ laminosa, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 196. 

1875. „ „ idem. Invert, and Fishes St. Andrews, p. 119. 

„ „ Paretti, Marion and Bobretzky. Ann. Sc. Nat., 6 ser., t. ii, p. 61. 

„ „ lamelligera, idem. Ibid., p. 62. 

1885. „ laminosa, Pruvot. Arch. Zool. Exper., 2 e ser. t. iii, p. 287, pi. xi, f. 6, 7, and pi. 

xiv, f. 4—8. 
1888. „ (Garobia) laminosa, De St. Joseph. Ann. Sc. Nat., 7 e ser., t. v, p. 274, pi. xi, fig. 

133—136. 

1890. „ laminosa, Malaquin. Annel. Boulon, p. 42 (Rev. Biol. Nord. Fr., t. ii, p. 437). 
„ „ „ Griard. Bull. Sc. Fr. Belg., t. xxii, p. 77. 

„ „ lamelligera, idem. Ibid. 

1891. „ laminosa, Hornel. Trans. Liverp. Biol. Soc, vol. v, p. 246. 

1897. „ lamelligera, Grravier. Bull. Sc. Fr. Belg., t. xxix, p. 308, pi. xvii, f. 14 — 16, pi. xviii, 

f. 1—4, pi. xix, f. 6—18, pi. xx, f . 1—6, pi. xxi, f. 4—9, pi. 
xxii, f. 1, pi. xxiii, f. 3 — 5. 

1900. „ „ Fischli. Polych. Ternate, p. 120. 

1902. „ „ Marenzeller. Polych. Grrund, p. 14. 

1904. „ laminosa, Allen. Journ. M. B. A., n. s. vol. vii, p. 223. 

1905. „ lamelligera (=P. Ehlersii, De Quatrefages), Grasife. Arbeit. Zool. Stat. Triest., xv, 

p. 325. 

Habitat. — Under stones in pools near low water, from Shetland to the Channel 
Islands; tossed on shore after storms. It is by no means confined to the Laminarian 



78 PHYLLODOOE LAMELLIGERA. 

zone as Benham 1 seems to think, though it is long since Prof. Jameson received from 
Mr. Neill examples dredged on oyster-shells in the Forth. 

Stretches to the Adriatic (Quarnero) — Ehlers ; shores of France (And. and Ed., 
De St. Joseph, etc.). Moluccas (Ternate) — Fischli. 

Head somewhat cordate, with the large eyes placed near its middle, and of a pinkish 
iridescent aspect, the ground colour being olive green, with a dark touch in front. 
Tentacles subulate. Tentacular cirri with a ringed ceratophore. Both appendages have 
the same dull greenish hue as the dorsal cirri. At the posterior and outer angle of the 
head, and just in front of the upper tentacular cirri, is a considerable papilla — probably 
sensory. A minute papilla also occurs in the centre of the posterior sinus of the head. 
The two longer tentacular cirri are often borne straight backward and adpressed to the 
smooth dorsum between the lamellse. 

Body linear, much elongated, from 14 to 18 inches or even 2 feet (Johnston and 
Dalyell), and with more than 400 segments. Convex dorsally, flattened ventrally, tapered 
a little anteriorly and more distinctly so posteriorly where it ends in two rather stout 
caudal cirri. The dorsum is of a fine iridescent bluish hue marked by regularly arranged 
transverse furrows. Each segment in the middle of the body has a narrow belt in front, 
succeeded by three others gradually extending in a transverse direction, a fifth twice as 
deep, and then a part not definitely bounded except by the segment-junction. In the 
middle line of the dorsum is a series of blackish specks, which, with the dark lamella on 
each side, make three in each segment. A conspicuous dark belt also occurs-on the upper 
and anterior edge of the pedicle of the foot. The ventral surface is likewise iridescent, 
but the bluish sheen is less developed. In life and in certain lights in the preparations 
three rows of spots are visible on this surface, the central consisting of a pale speck in 
the interrupted median band. The dusky pigment is arranged in transverse bands as on 
the dorsum. The remarkable iridescence of the cuticle of this species is conspicuous in 
softened preparations. 

In general the colour of the dorsum varies considerably, perhaps according to 
habitat, those most exposed to sunlight being most deeply tinted. In one fully two feet 
long, which was procured under a large stone about half tide mark, the bluish sheen of 
the dorsum is most marked in front, yet it does not conceal the two dark bars which cross 
the segment between the feet and the median dark patches which lie in front and behind 
them. The segments, moreover, are ringed. Posteriorly the hue is olive with a tinge of 
russet, and the dark pigment on the dorsum is arranged in two lateral touches and 
a median a little behind them. A trace of the bluish sheen is noticeable even to the tip 
of the tail. The general hue of the body is thus darker in front, paler (olive) posteriorly. 
In the same way the olive green pigment of the cirri is darker in the anterior, paler in the 
posterior region, the margin of the dorsal cirri being paler. The under surface is pale 
with a tinge of iridescent pink. 

The proboscis in extrusion forms a long, dark clavate organ, the basal section being 
covered with small papilla (Plate LVIII, fig. 20). The wall of the organ beneath the 
basement-membrane is marked by intersecting fibres. The distal region has numerous 
low transverse papillas which are not so distinctly arranged in a hexagonal form as in 

1 ' Camb. Nat. Hist./ ii, p. 314. 



PHYLLODOCE LAMELLIGERA. 79 

P. groenlandica, though they form rows. The organ terminates in a firm disc with about 
21 blackish papillae at its border. The distinction drawn between P. laminosa, with 
less regular papillse, and P. lamelligera, with the papillae in rows, does not seem to be of 
moment. 1 I regard them as synonymous. Gravier shows six nerves running longitudinally 
in a section of the organ. 

The proboscis is followed by a pale, elongated, and tough membranous stomachal 
region without papillae, after which comes the gut with its glandular walls. 

The perivisceral fluid readily coagulates in sea-water after extrusion. The cut edge 
of a lamella also exudes a coagulable fluid, and this forms minute vermiform processes at 
the wounds. 

This form is carnivorous, the fasces being loaded with bristles and cuticle of annelids, 
such as the Spionidae. 

The dorsal cirri are dusky green, paler at the edges and with small pale specks. In 
some specimens they are brownish green with a pale green margin, thus simulating 
the condition in Phyllodoce Paretti, Aud. and Ed., which has a bright yellow border to its 
leaflets. In a series of examples from various localities the cirri vary considerably, the 
Neapolitan examples, perhaps, being most pointed — both dorsally and ventrally. Fine 
examples from deep water off Montrose had them bluntly rounded in the middle of the 
body. 

The smaller specimens are usually paler than the larger, the bluish iridescence of the 
dorsum being only faintly marked anteriorly. The majority of those from the Channel 
Islands are small, with a brownish body and pale green lamellae. A similar condition 
would appear to occur at Naples. A variety also was obtained in the West Voe of 
Scalloway, in which the dorsum was dull yellowish, with the usual central line and cross- 
bars. The cirri were brownish olive with a dull yellow margin. Some large forms, again, 
have more of the greenish iridescence than the blue. 

The dorsal cirrus (Plate LXVIII, fig. 1) in the anterior segments (tenth) is 
somewhat reniform or ovate, at the thirtieth it becomes greatly enlarged superiorly so as 
to assume an elongate ovoid form, rounded and broad inferiorly and diminishing a little 
superiorly to a rounded tip. At the sixtieth foot (Plate LXVIII, fig. 2) the upper part 
is wide, so that the whole process forms a broad flap. Towards the tail, the lamella 
again becomes diminished dorsally, so that it resembles an acuminate leaf. The hypo- 
dermic streaks and vessels trend from the margins obliquely downward and inward to a 
central axis, and thus resemble the midrib of a leaf and its veins. A pale ridge just 
external to the inner edge of the lamella is conspicuous in the preparations. 

In the living form these lamellae have a band of powerful cilia (visible under a lens), 
commencing about a third from the apex of the process and coursing to the middle of the 
pedicle at the base. The cilia are placed just behind the dorsal border of the lamella, 
and do not follow the boldly marked curve proceeding downward and inward. The 
lamellae are highly sensitive, shrinking with a rapid jerk when interfered with. 

Parasitic Infusoria are common on the basal portion of the lamellae. Each can 
shorten and elongate its stalk, expand and contract its disc, which has a vestibule in the 

1 Vide Dr. Marion Newbigin, ' Millport Biol. Station Communications/ i, p. 3, 1900. 



80 PHYLLODOCE LAMELLIGERA. 

centre and a band of cilia around its edge. Free forms — -of the greenish hue of the 
borders of the cirri — become entangled in the mucus surrounding the annelid. 

The ventral division of the foot consists of a somewhat clavate process with a bifid 
tip, from which the spine issues, and beyond which the lamellge project considerably. 
The bristles (Plate LXXVIII, fig. 6) have pale shafts slightly curved towards their 
dilated extremities, which have a series of minute spines, forming a hispid margin on 
each side of the peak in some, and they are not confined to the edge only, but pass 
some distance downward on the obliquely striated bulbous region. In certain views 
the spines, which are slightly curved, appear to increase in size from the lower part of 
the dilated region to its apex. This explains the appearances frequently observed at the 
tip. The tapering terminal blade has its edge boldly serrated, and is of moderate length. 
The serrated edge, is split inferiorly, and each limb is spinous. Considerable variation 
occurs in the length of the spines, which are very long in small examples from Naples. 
The inferior cirrus is almost reniform in front (tenth), but becomes ovate-acuminate as 
the fiftieth foot is approached, and continues of this shape to the posterior end. It is 
fixed to the posterior and ventral border of the inferior division for fully a third of its 
length and is borne vertically. The variations in the feet may be estimated by comparing 
fig. 2, Plate LXVIII, with fig. 3, from Naples. 

Reproduction. — Some deposited green ova in a gelatinous mass in May. The animal 
readily secretes a large quantity of mucus and in this the eggs very probably are 
immersed. A ripe male, which is distinguished by the paler-yellowish hue of the body and 
the bases of the feet — caused by the masses of white sperms — occurred in June (19th). 
The sperms have a comparatively large head and long slender tail. Ehlers observes that 
the female is larger than the male, and Dalyell mentions that young specimens are 
usually green, but it is doubtful if he had always the same type before him. 

A recently developed tail is quite pale, and is thus in contrast with the fine blue 
segments in front. The dorsum of the pale region has a median row of dark specks, 
whilst a less distinct speck occurs at the base of each lamella. 1 

When immersed in clean sea- water the animal is for some time very restless, moving 
its elongated and complex body with ease and grace, first in one direction and then in 
another. The lamellge (cirri) reverse their feather-like motions in whole or in part as 
occasion requires, and apparently act in conjunction with the bristles, and thus the most 
varied movements are performed as easily by this animal with its feet in hundreds as by 
one with a few. After it secures a suitable position, no motion of any organ is noticed 
for considerable periods, the cilia on the lamella? sufficing for aeration. In this condition 
the lamellae are stretched outwards from the body, except at the curves, or when com- 
pressed against the glass ; while the tentacular cirri are laid straight backwards. By the 
aid of gelatinous mucus secreted in considerable quantity, it can also swim near the 
surface of the water, and appears generally to relish the gentle movement of the 
lamella?. A regular series of wave-like contractions pass from before backward — 
about five or six per minute — in some otherwise at rest. 

1 Vide A. Michel, " Recherches sur les Regenerations chez les Annel.," f Bull. Sc. Fr. et Belg./ 
31, ser. 4, 1893, pp. 245—417, pis. 13—19. 



PHYLLODOCE LAMELLIGERA. 81 

It is interesting to note how an animal of such extreme length moves so gracefully 
in the water, emerging from its folds and coils with ease and dexterity, and making 
progress over the bottom with considerable rapidity. 

The specimens of this species from Naples are but pigmy representatives of the 
gorgeous Scotch examples, a condition, perhaps, not unexpected in view of the small size 
of those from the Channel Islands. The head is more elongated, but this may arise from 
the mode of preparation, and it is noteworthy that whilst these small southern examples 
were diagnosed as P. lamelligera, the large, brightly coloured forms, were termed 
P. Paretti. The lateral papillae posteriorly, and the median in the sinus of the same 
region, are not distinguishable. The dorsal lamellae (Plate LXVIII, fig. 3), moreover, are 
more rounded, and the posterior ciliated band is very distinct. The tip of the ventral 
lamella is also more pointed. Judging from the spirit-preparations the pigment would 
seem to be much less developed than in the large northern examples. 

The specimens collected in Shetland were also of comparatively small size, but this 
may have been accidental. 

A variety apparently of this species was sent by Dr. Allen from Rum Bay, 
Plymouth, where it occurred in sand. The general hue was yellowish brown with white 
touches on the head, body, and tentacular cirri, the latter indeed being almost white. A 
white patch — bifid in front — ran forward to the inner side of each eye, and both were 
separated from the pale area bearing the tentacles by a brown belt — the bifid part being 
thus prominent. On the dorsum of the body the pale madder-brown pigment was cut 
into three rows by a touch of white on each side of the median stripe (Plate XLIX, fig. 1). 
On the ventral surface (Ibid. fig. 2) the median sulcus had a stripe of madder-brown, and 
a yellowish brown touch, fading externally on each side of it. The dorsal cirri had a 
central brown touch and a pale margin, and their shape seemed to be more distinctly 
triangular (or heart-shaped) than in the ordinary form (Plate L, fig. 7). The shape of 
the head, which agreed with those from the Channel Islands in being more elongate, and 
its processes, correspond with P. lamelligera, and so far as observed the proportions and 
minute structure of the bristles do not differ in any respect, and the same with regard to 
the setigerous region and the other parts of the foot. The specimen was much softened 
and injured before it was minutely examined by other than the artist. 

In transverse section of the body-wall of P. lamelligera the great depth of the 
hypoderm and its glandular nature are marked, as also is the strength of the circular 
muscular coat, and the great extent of the dorsal longitudinal sheets which come down on 
each side of the oblique muscles. The ventral longitudinal muscles are considerably less. 

In this species the nephridial region of the segmental organ is stated by Louis 
Fage 1 to be four-lobed, but the genital funnel resembles that in allied forms, e.g. that of 
P. Paretti. 

Col. Montagu (1808), in the MS. volume of drawings by Miss Dorville, represents a 
comparatively small and pale specimen of this species, and a foot. It is such southern 
variations which subsequently have given rise to confusion in species. Montagu himself 
probably referred to one of these variations, as Nereis argentea, in Plate XXXII, fig. 2, 
but whether of this or another species is uncertain. 

1 ' Ann. Sc. Nat./ 9 e ser. ; in, p. 263, fig. 10. 

68 



82 PHYLLODOCE PAEETTI. 

Eisso's genus Eunomia (1826) probably refers to Eulalia or Phyllodoce, and his 
E. tympana is a reddish form perhaps akin to Phyllodoce rubiginosa. 

The Phyllodoce mucosa of (Ersted 1 may have some relation to a young form of this 
species. 

In the illustrated edition of Cuvier (1849), Milne Edwards shows a variety with a 
dark spot at the segment-junction near the base of the cirrus on each side. 

Ehlers (1864) gives an extended account of the structure of this form which he 
considered different from Savigny's P. laminosa. He points out that the colour of the 
latter is a beautiful green, that it appears to differ in the form of its bristles, the arrange- 
ment of the papillaB of the proboscis, and other details. In alluding to this author's views 
Miss Newbigin leaves the question undecided. 3 

The Phyllodoce Kinbergi of De Quatrefages (1865) would seem to come near this 
species, if it is not identical therewith. His P. BathJcii s appears also to be a well-marked 
pale variety. 

Malmgren (1867) considered that Rathke's form though similar did not pertain to this 
species, which had not yet been found in Scandinavia. 

This seems to be the Phyllodoce marginata of Gosse from Torbay. 

Tauber (1888) is of opinion that P. metadata, Mull, and (Erst., P. Millleri, Leuck., 
P. teres, pulchella, badia, Rinhii, citrina, groenlandica, incisa, mucosa, and assimilis all 
belong to Ph. lamelligera, Johnston. Further study may lead to a union of such forms as 
Phyllodoce lamelligera and P. groenlandica, but it is advisable to leave them separate at 
present. Ehlers separates P. laminosa, Sav., from P. lamelligera, Johnst. Cznerniavski 
ranges P. laminosa under the sub-genus Genetyllis. . 

Giard (1890), like Ehlers, separates the Phyllodoce laminosa of Savigny from the 
P. lamelligera of Johnston, which is rare at Wimereaux, and is, he thinks, near the type 
of P. Pancerina and P. splendens. So far as British experience goes, the latter suggestion 
would seem to have slender foundation, but the experienced zoologist of Wimereaux may 
have reasons — not yet disclosed — for his view. 

Goodrich 4 finds a distinction between the solenocytes of Phyllodoce Paretti, where 
they are grouped only at the tips of the branches, and those of P. laminosa, in which the 
cell-bodies are bent round so as to rest on the segmental organs, the latter rather 
resembling those of Glycera unicornis than those of its nearer allies. In the latter species, 
also, he says cilia are developed on the coelomic surface between the rows of solenocytes. 

2. Phyllodoce Paretti, De Blainville, 1828. Plate XLIV, figs. 6, 6a to 6c; 

PI. LXXIX, fig. 27— bristle. 

Specific Characters. — Head comparatively small, rounded, with two large eyes, which 
are blackish with a dull yellow rim, the colour being continued posteriorly as a broad band 
to the edge of the buccal segment. The rest of the head is pale brown, especially 

i 'Annul. Dame. Consp./ p. 31, figs. 25, 79, 85 and 89. 

2 ' Communic. Millport Marine Lab./ i, November, 1900, p. 3. 

3 c Annel./ ii, p. 131, pi. ix, figs. 12—14. 

4 'Quart. Journ. Micros. Sc./ n. s., vol. xliii, p. 707, fig. 13, 1901. 



PHYLLODOCE PARETTI. 83 

between and in front of the eyes. The terminal tentacles are large and dull yellow. 
Posterior border of the head entire. Tentacular cirri long, comparatively thick, 
tapered at the tip and of the same yellowish colour as the tentacles. Body 6 — 10 inches 
in length and 4 — 5 mm. in diameter, slightly tapered anteriorly and more distinctly 
so posteriorly. Iridescent bluish, paler in some, with a shade of pink mesially. 
Ventrally the same bluish iridescent colour prevails. Dorsal cirri of a rich deep green 
with a border of yellow or greenish yellow. Posteriorly the body terminates in 
two rather thick anal cirri of the shape and colour of the tentacular cirri. Foot 
with foliaceous dorsal cirrus which is bluntly cordiform in front, nearly regularly 
cordiform in the middle, and more or less lanceolate posteriorly. Setigerous region 
a blunt cone, bifid at the tip, and having a single powerful spine and a fan of nearly 
straight (the distal curve being slight) translucent bristles, the ends of the shafts 
being bluntly clavate, so that the region is short and minutely spinous. The terminal 
process is characteristically small and tapers to a fine point with a serrated 
anterior edge. Ventral cirrus irregularly reniform or ovato-lanceolate in front, ovato- 
lanceoktte in the middle of the body and more pointed (that is more distinctly 
lanceolate) posteriorly. 

Synonyms. 

1828. Nereiphylle de Paretto, De Blainville, Diet. Sc. Nat., lvi, 476, et Atlas, pi. xiii, fig. 1. 

1829. Nereis Paretti, Delle Chiaje. Mem., iv, p. 195, Tav. lxiv, f. 3—5. ? 

1833. Phyllodoce Paretti, M. Edwards. Ann. Sc. Nat., xxix, p. 248. 

1834. „ „ Audouin et Edwards. Annel., p. 229. 

1840. „ Bathhii, Grube. Actin. Ech. u. Wiirm., p. 78. 

1841. „ Parettiana, Delle Chiaje. Discriz., iii, p. 98, et P. Paretti, v, p. 104, Tav. 166, f. 3. 
1849. „ Paretti, M. Edwards. Reg. Anim. Illust., pi. xiii, f. 1. 

1851. „ Rathhii, Grube. Fam. Annel., pp. 55 and 129. 

1865. „ Paretti, De Quatrefages. Annel., ii, p. 130. 

1870. „ Pancerina, Claparede. Annel. Nap. Suppl., p. 92, pi. ix, fig. 1. 

1888. „ Paretti, De St. Joseph. Ann. Sc. Nat., 7 e ser., torn, v, p. 279. 

„ „ (Carobia) splendens, idem. Ibid., p. 278, pi. xi, fig. 138. (?) 

1904. „ Paretti, Allen. Journ. M.B.A., n. s., vol. vii, p. 224. 

1905. „ „ G-raeffe. Arbeit. Zool. Stat. Triest., xv, p. 325. 

1906. „ splendens, De St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii. (?) 

Habitat. — Guernsey, between tide-marks (Ray Lankester) ; offshore in depth of 
10—15 fathoms, Plymouth (E. J. Allen). 

Shores of France (Milne- Edwards, De Saint Joseph). Mediterranean (Delle Chiaje, 
Grube, Claparede). 

Head rounded or ovoid, comparatively small, with two large black eyes surrounded 
by a yellowish rim, and passing as a band from each to the posterior border of the region. 
They look both dorsally and laterally. The terminal tentacles are large, fusiform, and with 
a short slender tip. The upper pair are carried forward slightly sloping from each other. 
A speck of dark pigment occurs behind the base of each. The ventral pair are usually 
borne transversely or with the tips pointed downward. Their shape is similar to that 



84 PHYLLODOCE PARETTI. 

of the dorsal pair. The ventral surface of the head is — like the median dorsal — pale 
brownish. No distinct depression occurs at the posterior border. 

Tentacular cirri vary in length in different examples. In that from Plymouth 
(Plate XLIV, fig. 6) they are somewhat short, thick but tapered terminally, and have 
the same yellowish colour as the tentacles. The first or most anterior of the series in life 
springs from a ceratophore which projects forward at the side of the head, is fusiform in 
outline and ventral in position. Its yellow is slightly tinged with brown. The next is 
dorsal but in rear of the foregoing, is carried more or less erect, usually sloping 
obliquely outward, and forms a long flattened lanceolate leaf, its yellowish hue also having 
a few brownish touches, sometimes as a band on each side of a midrib of dull yellow on 
its upper or anterior surface. The third lies below the former towards the ventral border 
of the buccal segment, is fusiform in outline, slightly flattened and about the same length 
as the first. The fourth arises from a ceratophore which projects forward from the second 
ring of the body, and passes horizontally outward, being less prominent than the con- 
spicuous second pair. It is next the latter in size and has a flattened lanceolate outline 
with a pointed tip — its general yellow being tinged with brown. 

Body 6 — 10 inches in length and 4 — 5 mm. in diameter, distinctly tapered anteriorly, 
and more so posteriorly where it ends in two rather thick anal cirri of the shape and 
colour of the tentacular cirri. The coloration is most striking, the dorsum being of a 
rich iridescent blue throughout, or in some with a tinge of pink mesially, probably from 
iridescence, whilst the ventral surface is similarly tinted — only slightly paler. In front 
the mesial part of each segment — between the feet — is darker, the narrower band at the 
segment-junction being paler, but throughout the rest of the body the segment-junction is 
discriminated by a glistening transverse line on each side of a narrow belt. Ventrally 
the segment-junctions are marked by a straight line anteriorly and by two flattened 
spindles or crescents behind, so that there is a forward sinus in the middle lin#. 

The lamellae (dorsal cirri) are of a rich deep green or blackish-green with a border 
of yellow or greenish yellow, and they have a tendency to be curled and twisted, especially 
in specimens in confinement, a feature greatly increasing the complexity of the dorsum. 
The ventral lamellse which lie behind the setigerous lobes are considerably paler, being 
light green with bold touches or a belt of brownish pigment within the border, which has 
a margin of yellow. In some the buccal and the second segments are marked off by the 
deep madder-brown hue of the two following ventral and dorsal cirri, this colour thus 
bringing into relief the yellowish tentacular cirri, and the yellow margin of the dorsal 
cirri. 

Few animals — marine, fresh- water, or land — can excel this Phyllodoce in the chaste 
beauty of its coloration, whether viewed in whole or in part. Though the hues are 
similar throughout the cirri, yet the soft folds and curvatures of each present endless 
variations, so that whilst the general plan is carried out there is no monotony. The 
dark iridescent blue of the body gleams as the annelid moves with an ever changing 
lustre which relieves the dark green and yellow of the foliate cirri. 

In one with a reproduced tail, the new segments with their lanceolate dorsal cirri 
are yellowish, just a tinge of green or bluish-green appearing in the centre of the 
anterior cirri so as to differentiate the yellowish border. The ventral cirri are yellowish 



PHYLLODOCE PAEETTI. 85 

throughout. The body of the new region is yellowish with a slight shade of green 
dorsally, a somewhat opaque linear dorsal streak indicating the intestine. Yellow would 
thus appear to be the fundamental colour, the greens and browns being subsequently 
formed. 

The proboscis has not been extruded in any example. De Saint Joseph found thirty 
papillse at the entrance to the stomach. 

The foot has dorsally the foliaceous cirrus, which springs from a long pedicle 
and anteriorly has a bluntly cordiform outline, a regularly cordiform shape in the middle 
and following regions of the body, and a more or less lanceolate outline posteriorly. In 
the anterior and middle regions it is flecked with dark pigment in Neapolitan specimens, 
and in one from Plymouth the dark pigment is even more conspicuous. The setigerous 
region forms a somewhat long blunt cone with a horizontally bifid tip. 

The shafts of the bristles (Plate LXXIX, fig. 27, male) have a very slight distal curve, 
and the hue is brownish. Moreover, the dilated terminal region is more or less truncate, 
and thus the spinous area is shortened by the absence or abrasion of the conical end with 
its larger curved hook-like spines. The contrast with the typical Phyllodoce lamelligera is 
thus marked. The truncate end of the shaft in Paretti has a much shorter spinous 
region, but the trend of the spines is the same. The distal sabre, moreover, is shorter, more 
filiform towards the tip, and the serrations on the edge are more minute. If the effect of 
age and special surroundings affected the bristles as just described, then the connection of 
this form with Phyllodocera lamelligera would be clear. In an example from Plymouth the 
free parts of the bristles were tinted of a brownish hue, the same pigment being greatly 
developed in the cirri and setigerous region. 

Reproduction. — Goodrich and Louis Fage * describe the segmental organs as having a 
largely ramified nephridial lobe, each of the divisions with its solenocytes and tubercles. 
The funnel is large and crenate. 

The Phyllodoce Paretti, De Blainville, 1 is a species about which diverse opinions may 
be taken, more especially as no connected series from the young to the adult condition ' 
has been studied. So far as known all those procured in the Channel Islands and in 
South England have been adult, though, if the Phyllodoce sjplendens is the same form, two 
young examples, of 7 and 16 mm. respectively, about which, however, there maybe doubt, 
were found by De Saint Joseph. 

Grube's 2 original description (1840) of Phyllodoce Rathldi agrees with the structure 
of P. Paretti, for he notes the form of the tentacular and dorsal cirri, the nature of the 
bristles, and the coloration. He found it at Palermo. 

Grube's 3 diagnosis is : Mouth-segment scarcely visible from the dorsum; dorsal cirri 
heart-shaped ; the longest tentacular cirrus as long as the fifth segment ; on each side only 
four tentacular cirri, whereas P. laminosa has five. 4 

De Quatrefages appears to have made two new species in connection with this form, 
for his P. Kinbergi can scarcely apply to any other European species — notwithstanding 

1 f Ann. Sc. Nat./ 9 C ser. hi, p. 282, figs. 8 and 9. 

2 ' Actin. Ecliin./ etc., p. 78. 

3 <Fam. Annel./p. 129. 

4 The author had misinterpreted the condition in this form. 



86 PHYLLODOCE GRCENLANDICA. 

his figure of the elongated dorsal cirri anteriorly — a feature which may have been due to 
imperfect preservation. He makes the remark under P. Paretti that the appendages of 
the bristles are very short, and for this reason he emphasises the distinction between it 
and the former. It may be doubted, however, whether the structure of the bristles was 
studied with sufficient minuteness, and the figures bear out this view. 

Claparede (1870) places much reliance on the relative length of the tentacular cirri 
as distinguishing his P. Pancerina from P. Paretti, the former having them very long 
and thick. It is doubtful, however, if much weight can be placed on this point. The 
tentacular cirri arise from the same transverse line as in P. Rathkii of Grube, whereas 
Claparede thought such did not exist in the P. Paretti of Edwards and Delle Chiaje, nor 
in the P. Kinbergi, De Quatrefages, a species closely approaching P. Pancerina. He 
thought the bristles differed from those of P. Paretti in having the terminal piece 
undulated and somewhat longer — distinctions which may be found in the same specimen. 

Marion and Bobretzky (1875) consider that Phyllodoce Pancerina (Claparede) is the 
same as P. Paretti, and they separate it from P. lamelligera (Johnston). Claparede's form 
may be a well-marked southern variety, with swollen tentacles and enlarged lamellae. 
Delle Chiaje does not show a yellow border to the dorsal cirri in his figure in the 
6 Memoire,' but mentions it in his description. 1 

The Phyllodoce splendens 2 (De St. Joseph), which the author links closely to 
P. Kinbergi (De Quatrefages), P. Paretti, De Blainville, and P. Pancerina (Claparede), 
seems to be the same form. It agrees in colour, the small size and shape of the head, 
and other features. He places the tentacular cirri thus : Two pairs on the buccal, the third 
pair on the second segment, and the fourth on the third segment, with a rudiment of a foot 
without a ventral cirrus. The dorsal cirri are cordate at the base, and the eggs green. 
Fragments of Synapta inhserens occur in the stomach. He only found two young ones 
(16 mm., 82 seg.; and 7 mm., 41 seg.). " Les soies ont la hampe tres renflee en avant," 
and the terminal process is shorter than in P. Pancerina, and thus agrees with P. Paretti, 
of which it is probably a variety. The author's P. papulosa 3 from Dinard, also comes 
very near the present form. 



3. Phyllodoce gecenlandioa, (Ersted, 1843. Plate LVIII, fig. 5— head; Plate LXVIII, 
figs. 4, 5, 6— feet ; Plate LXXVII, fig. 7— bristle. 

Specific characters. — Head somewhat ovate, cordate posteriorly ; two eyes, tentacles 
short, tentacular cirri normal. Body of considerable length, four to eight inches ; greyish 
green ;. segments short. Proboscis in extrusion with six rows of large tubercles 
anteriorly, numerous rows of smaller conical papilla? posteriorly. Dorsal cirri much 
developed superiorly, so as to be almost rectangular ; ventral cirri ovate-elliptical, and in 
the middle segments turned upwards ((Ersted) ; setigerous process bifid, shafts of bristles 
with enlarged spinigerous tips and a long serrated blade. 

1 p. 176. 

2 ' Ann. des Sc. Nat./ 7 e ser., torn, v, p. 278, pi. xi, fig. 138, 1888. 

3 Ibid., 8 e ser.,, t. v, p. 320, pi. xviii, figs. 117—120, 1898. 



PHYLLODOCE GRCENLANDICA. 



87 



Body-wall with a comparatively thick coat of hypoderm under the cuticle, and 
the circular muscular coat is well developed. The dorsal and ventral longitudinal 
muscles are large, the former being almost separated by a median fissure, their outer 
borders in section being very massive, whilst the ventral are somewhat ovoid. The 
fasciculi in the ventral are flattened in transverse section, and almost all have their ends 
placed externally and internally, the flat faces being parallel to each other. 

The oblique muscles meet or decussate beneath the nerve-cords, the circular coat, 
the basement membrane, the hypoderm and cuticle being external. In certain sections 
their attached ends widen into a broad fan. 



Synonyms. 

1842-3. Phyllodoce groenlandica, OErsted. Kroyer's Nat. Tids., p. 121. 

1843. „ idem. Gronl. Annul. Dors., p. 192, f. 19—20, 22, 29, 30, 31, 32. 

idem. Annul. Dan. Consp., p. 32. 
Sars. Beskriv., Tab. ii, figs. 19, 21, 22, 29—32. 
1851. ,, ,, Grube. Fam. Annel., pp. 56 and 129. 

Sars. Nyt Mag., vi, p. 209. 
„ „ mucosa, (Ersted. Consp. Annul., p. 31, f. 25, 79, 83, and 89. 

1853. „ groenlandica, Stimpson. Invert. Grand Manan, p. 33. 

1863. „ „ idem. Annel. Greenland, Proc. N.S. Philad., p. 140. 

1865. „ „ Malmgren. Nord. Hafs- Annul., pp. 96, 129. 

,, „ „ De Quatrefages. Annel., ii, p. 141. 

„ „ lamelligera, Carrington. Annel. Southport (Proceed. Lit. and Philos. Soc. 

Manch., iv), p. 5 (180). 
1866-9. „ groenlandica, Packard. Mem. Bost. Soc. Nat. Hist., i, p. 294. 

1867. „ „ Malmgren. Annul. Polych., p. 21. 

1871. „ „ Ehlers. Sitz. phys. med. Soc. Erlangen, iii, p. 79. 

1873. „ ,, Sars. Bidrag Christ. Fauna, iii, p. p. 223 — 4. 

1874. „ „ Malm. Op. cit. Goteb., p. 79. 

„ „ „ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 196. 

1875. „ „ idem. Invert, and Fishes St. Andrews, p. 119. 

1878. „ „ Marenzeller (Nordpol. Exped.) . Denksch. d. k. Akad. Wien, 35 Bd., 

p. 395. 
„ „ „ Mcintosh. Trans. Linn. Soc, Ser. 2, p. 502. 

1879. „ „ Tauber. Annul. Danic, p. 87. 

Theel. K. vet. Akad. Forhl., Bd. 16, 3, p. 34. 

1880. „ „ Grube. Jahresb. Schles. Ges., 1879, p. 214. 

1881. „ „ Horst. Niedel. Arch. Zool., Suppl. Bd. i {« Willem Barent" Exped., 

1878), p. 11. 

1883. „ „ Wiren. Cheetop. "Vega " Exped., p. 400. 

1884. „ „ Webster and Benedict. Rep. U.S. Comm. F. and F., p. 703. 
1896. „ „ Michaelsen. Polych. deutsch. Meere (Biol. Anst. Helgol., ii Th.), 

p. 32. 
1898. „ „ De St. Joseph. Ann. Sc. Nat., 8 e ser., t. v., p. 326, pi. xviii, figs. 

124—126. 
1901. ,, „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. viii, p. 223 (bis). 

„ „ „ Whiteaves. Geol. Surv. Canada, No. 722, p. 82. 



88 PHYLLODOCE GROEXLANDICA. 

Habitat.— Tossed on shore in great numbers after storms on the West Sands, St. 
Andrews (E. M.), and probably inhabits the sand inshore. Stomach of cod (E. M.). 
Montrose Bay (Dr. Howden). Connemara, Co. Gal way (A. Gr. More). 

The species is common in the seas of Spitzbergen, Nova Zembla, and Greenland, but 
less frequent in Norway, Finmark, and Sweden. Extends to Canada (W. C. M.) and the 
American coast. Siberia and Behrings Strait (Marenzeller, Wiren). 

Head (Plate LVIII, fig. 5) somewhat ovate with a cordate posterior border, two 
acute peaks being separated by a deep fissure. It is longer than broad. The eyes are 
well marked, and occur about the commencement of the posterior third. The tentacles 
are short and subulate. The tentacular cirri have the usual proportions. 

Body slightly tapered anteriorly and much more so posteriorly, rounded dorsally and 
somewhat flattened ventrally where a median groove occurs. The segments are distinctly 
marked on both surfaces by deep grooves. The mouth forms a transverse fissure 
ventrally with various furrows. The slender tail bears two caudal cirri. The animal is 
of a sandy or greyish-green hue with darker pigment on the edges of the lamellse. It 
tinges spirit greenish and becomes so itself. The exserted proboscis is sub-cylindrical, 
but is often clavate. Proximally twelve rows of elongated papillae (thirteen to seventeen 
in each) are ranged in two groups of six, a blank occurring in the mid-dorsal and the 
mid-ventral line. The distal region (in extrusion) presents a somewhat hexagonal 
appearance in transverse section, from the disposition of the large tubercles or blunt 
papillas which form six rows. Seventeen papillae surround the aperture, the furrows 
between them trending inward so as to make a long and slightly tapering process to 
each papilla, and as the aperture rapidly diminishes, the whole constitutes a fluted basin 
with a central aperture. 

The dorsal division of the foot (Plate LXVIII, figs. 4, 5, and 6) forms a short and 
massive pedicle bearing the more or less vertical cirrus, which in the anterior part of the 
body, as at the tenth foot, has a short conical form, whilst at the sixtieth, its dorsal part 
has become so much developed as to give it almost a rectangular outline, the dorsal 
margin being nearly straight. Towards the posterior part of the body it again becomes 
rounded dorsally. The hypoderm of the cirrus has a somewhat radiate arrangement. 

The inferior division consists of a bifid setigerous lobe 1 — the spine issuing from the 
terminal pit. The shafts of the bristles (Plate L XXVII, fig. 7) have a terminal 
curvature and a dilated end, the sides of which are minutely serrated. The terminal 
blade is long, finely tapered, and minutely serrated. The ventral cirrus is ovate, 
elliptical, with an acuminate tip, and is carried nearly vertically at the posterior border 
of the foot. It is shorter and broader anteriorly, more elongated and more pointed near 
the tail. 

The original description and figures of (Ersted (1843) leave no doubt concerning this 
species, though perhaps the outlines might be improved. His specimens had from 
300—350 segments. He considered that the bristles agreed with those of P. lamelligem. 

The Phyllodoce fragilis of Webster 2 from the Virginian coast seems to be a closely 
allied form, which occurs elsewhere on the shores of America. 

1 The setigerous process is more elongated in specimens from Southport. 

2 ' Trans. Albany Inst./ ix, p. 14, pi. iii, figs. 32—37, 1879. 



PHYLLODOCE MACULATA. 



89 



4. Phyllodooe maculata, Linnseus, 1767? (Johnston, 1842). Plate XLV, fig. 2; 
PL XL VII, fig. 3; PL LVIII, figs. 21 and 24; PL LXVII, figs. 7 and 8— feet; 
PL LXXVIII, figs. 23 and 24— bristles. 

Specific Characters. — Head longer than broad, speckled with yellow, cordate 
posteriorly. Eyes two, of moderate size. A median and two lateral yellow spots on the 
dorsum of the first segment. Body long, linear with three dorsal rows of dark spots, 
the lateral from the lamellae. Posteriorly a yellowish- white spot appears between the 
central rows. Proboscis subcylindrical with six rows of small papillae on each side in 
extrusion, distally six rows of large tubercles or folds, and terminating in a circle of 16 — 17 
papillae. Dorsal cirrus of the foot irregularly ovate, the dorsal edge being rounded, or 
subrectangular throughout the greater part (middle) of the body. Ventral cirrus broadly 
lanceolate, the pointed tip projecting beyond the setigerous lobe. Spinigerous region 
rather short, bifid. Bristles with a dilated end to the shaft, and spinigerous curves. 
Terminal blade of moderate length and minutely serrated. 



1767. 
1776. 
1780. 
1789. 
1791. 
1800. 
1806. 
1825. 
1829. 
1830. 
1842-3. 

1843. 
1849. 
1851. 



1865. 

1867. 

i) 
1874. 
1875. 

)) 
1879. 
1883. 
1886. 
1890. 
1891. 
1893. 



Synonyms. 
Nereis maculata, Linnseus. Syst. Nat., ed. 12, i, pt. 2, p. 1086. 

O. F. Miiller. Zool. Dan. Prodr., p. 217, nr. 2635. 
,, ,, O. Fabricius. Fauna GrcenL, p. 298. 

Bruguiere. Encycl. Meth., Vers, i, p. 134, Tab. 57, f. 1—6. 
„ „ Linn. Grmelin, Syst. Nat., ed. 13, i, pt. 6, p. 3118. 

Die gefleckte Nereide, O. F. Miiller. Naturges. einiger Wiirm-Arten, p. 156, Tab. x, f. 1 
Nereis maculata, Turton's Grmelin, iv, p. 88. 

n „ De Blainville. Diet. Sc. Nat., xxxiv, p. 446. 

Phyllodoce pulchra, Johnst. Zool. Journ., iv, p. 54. 
Nereis maculata, Bosc. Vers, i, p. 171. 
Phyllodoce maculata, (Ersted. Kroyer's Nat. Tids., p. 121. (?) 

» 99 Johnston. Ann. Nat. Hist., iv, p. 227, pi. vii, f. 1 — 3. 

(Ersted. Gronl. Annul. Dorsib., p. 191, figs. 46 and 48. (?) 
Mulleri, B. Leuckart. Archiv f. Nat., p. 204, Tab. iii, f. 13. 
„ maculata, Grube. Fam. Annel., p. 55. 

„ Mulleri, idem. Ibid., p. 56. 

„ maculata, idem. Ibid., pp. 55 and 129. 

Johnston. Cat. Brit. Mus., pp. 177 and 342. 

De Quatrefages. Annel., ii, p. 138. 

Malmgren. Annul. Polych., p. 144, Tab. iv, f. 16. 

Parfitt. Trans. Devon. Assoc., ii, pt. 1, p. 232. 

Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 196. 

Idem. Invert, and Fishes St. Andrews, p. 119. 

Mobius. Jahresb. Com. deutsch., p. 170. 

Tauber. Annul. Danic, p. 87. 

Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 205. 

Idem. Kara Hav. Ledorme., p. 7. 

Griard. Bull. Sc. Fr. Belg., t. xxii. 

Hornel. Trans. Liverp. Biol. Soc, vol. v, p. 246. 

Levinsen. Yidensk. Ud. "Hauchs.," p. 326. 

69 



90 PHYLLODOCE MACULATA. 

Habitat. — Not uncommon amongst the rocks between tide-marks at St. Andrews, 
under stones (E. M.), and at Lochmaddy dredged on the surface of sandy mud amongst 
alga3 in the laminarian region. Also in sand near low water-mark, St. Andrews. 
Iceland (Malmgren). Faroe (Willemoes-Suhm). 

Head longer than broad, 1 pale, speckled with yellow, cordate posteriorly. Eyes two, of 
moderate size. Tentacles subulate with finely tapered tips. Tentacular cirri normal. Imme- 
diately behind the eyes (apparently on the first segment) are three bright yellow spots, a 
central and two anterior. In some the tentacular cirri have a dark spot a little below the tip. 
Body from 3 to 4 inches in length, linear, tapering gently anteriorly, with three rows 
of dark spots, the central being most conspicuous. Posteriorly a yellowish- white spot 
appears between the dark, central ones. Many have a narrow yellowish bar between each 
of the brown bars, and as the centre is somewhat imperfect, it has thus the aspect of two 
specks. The lateral brown spots are due to the dorsal lamellse. In some the body is of 
a deep greenish hue on the dorsum with a median dark stripe and a dumb-bell shaped 
transverse bar (Plate XLVII, fig. 3). Such may be due to the development of ova in 
March. A dark variety in which the colours are beautifully marked is common amongst 
the tangles in Bressay Sound and on a sandy bottom in Vatsland Bight. The under 
surface is pale. The body tapers posteriorly and ends in two lanceolate cirri. 

The proboscis in extrusion (Plate LVIII, fig. 21) presents proximally two groups of 
six rows of papillas, which are fewer (about ten in each row) than in P. groenlandica. The 
distal region has a somewhat hexagonal outline in section from the six rows of large 
tubercles or folds. The aperture presents the same fluted condition as in the former 
species, with about sixteen papillas. 

When the alimentary canal has been largely distended before section, the cells, 
especially dorsal, arrange themselves in two series, the outer with large granular nuclei 
which stain deeply. 

The muscular system of this species conforms to the type, though the scale is smaller. 
The muscular fasciculi are similarly arranged in the dorsal and ventral longitudinal 
muscles. The oblique muscles pass below the proportionally large nerve-cords to the 
basement-membrane, which — with the hypoderm and cuticle — are external. A considerable 
area of granular tissue lies between the cords ventrally. 

The dorsal division of the foot (Plate LXVIII, figs. 7 [tenth] and 8 [sixtieth]) bears 
a cirrus which is irregularly ovate, the dorsal edge being rounded, so that irrespective of 
size it differs from the typical lamella of P. groenlandica. In the anterior region the 
lamella has the form of a conical or ovate-acuminate leaf, and posteriorly a similar outline 
is assumed. In the intermediate region it is sub-rectangular. The bases of these lamella? 
are ciliated, and stalked Infusoria are common on them. 

The inferior division consists of a rather short bifid spinigerous lobe with the bristles 
(Plate LXXVII, fig. 23) which have slightly curved shafts with dilated tips finely 
serrated. The dilated region of such a bristle is grooved so as to guard the origin of the 
tapering terminal blade, which is of moderate length and minutely serrated. The ventral 
cirrus is broadly lanceolate, the pointed tip projecting beyond the setigerous lobe. 

The dorsal cirri have very little of the greenish pigment anteriorly, so that they are 
1 Malmgren ; s "latitudine longitudinem superans/' referring to spirit preparations. 



PHYLLODOCE MACULATA. 91 

pale, but towards the latter half of the body the increase of the pigment renders them 
more conspicuous, especially from the dorsum, as dull greenish processes. A slight change 
occurs after immersion in spirit, and the minute structure of the dorsal cirrus is rendered 
indistinct ; but the alteration is not so great as might be anticipated. Besides, to remove 
a foot from a living specimen seriously cripples it and leads to its destruction, and the 
operation (by no means simple in an entire form) usually results in crushing the foot. 

An interesting abnormality was found at St. Andrews, viz., the presence of five anal 
cirri (Plate LVIII, fig. 24). On the left are a small pair of cirri, posteriorly two larger, 
whilst a single very large cirrus occurs on the right. So far as appearances go, such 
would seem to indicate a tendency to form buds. It was observed by my sister 
(Mrs. Grunther) while collecting a number for her coloured drawing. 

Reproduction. — The ripe females (June 18th) undergo a considerable change in 
colour due to the development of the green ova internally. The early development of 
the ova, which are of a fine green colour, and which are deposited in a somewhat bulky 
gelatinous mass, and the early larvas, have been described and figured 1 to the advanced 
trochophore stage, and by Alexander Agassiz 2 to a somewhat later stage. 

Garstang gives January and February as the breeding period of this form at 
Plymouth, so that if correct it is much earlier than at St. Andrews. 

It is very active, and hides in fissures and crevices into which the long, flattened, 
mobile body fits easily. The flattened snout is pushed hither and thither, while the 
pliable tentacles and tentacular cirri are readily doubled with safety. 

This is the Nereis maculata of Montagu (1808) in the MS. volume of drawings in the 
possession of the Linnean Society, Plate XIX, fig. 3. 

Malmgren considers that Johnston's P. maculata belongs to a different species, but 
there is no reason to believe that the British author referred to any other form than the 
present. The specimen in the British Museum, from Berwick Bay, has been dried, and 
therefore the characters are indistinct, but it approaches the ordinary form most closely. 
So far as can be observed, there is nothing in the original description and figures of 
0. F. Miiller 3 to show that he alluded to another species. CErsted considered that Johnston 
referred to his form, and therefore the views of De Quatrefages, who instituted the 
separate species, Phyllodoce OErstedii* for (Ersted's annelid, and those of De St. Joseph, 
who thought that (Ersted's form, while agreeing with the P. citrina of Malmgren, differed 
from Johnston's, appear to be doubtful. 

It is possible that the Phyllodoce attenuata of Carrington 5 (1865) is to be referred to 
this species. It is more difficult to say to what form his P. clava is to be relegated. 

Grube (Schles. Gresell, 1879) notes that perhaps Johnston's species and the Eidalia 
quadricornis of (Ersted 6 (which may have only four tentacles) are forms pertaining to 
Anaitis, but it is more probable that such is a misapprehension. 

1 ' Ann. Nat. Hist./ ser. 4, vol. iv, p. 104, 1869. 

2 'Ann. Nat. Hist./ ser. 3, vol. xix, p. 240, 1867. 

3 'Naturges. Wurm-Arten/ p. 156, tab. x. 

4 'Annel./ ii, p. 139. 

5 'Annel. of Southport/ 1865, p. 5. 

6 'Annul. Dan. Consp./ p. 28. 



92 PHYLLODOCE RUBIGIXOSA. 

Tauber 1 (1879) adopts the opinion that the Phyllodoce teres, P. pulchella, P. badia, 
and P. BinJcii of Malmgren are only varieties of this species ; and, still further, that the 
Phyllodoce laminosa of Savigny and Aud. and Edwards, the P. lamelligera of Johnston, the 
P. remex, Dalyell, P. groenlandica, GErsted, P. citrina, Malmgren, P. mucosa, (Ersted, and 
P. assimilis, (Ersted, are all one and the same species. There is no doubt that several of 
these, such as P. citrina, P. mucosa, and P. groenlandica, are very closely allied. 

5. Phyllodoce eubiginosa, De St. Joseph, 1888. Plate XL VII, figs. 4—5; Plate LVIII, 
fig. 25— tail; Plate LXVIII fig. 9— foot; Plate LXXVII, figs. 8 and 8a— bristles. 

Specific Characters. — Head small, rounded, with two large eyes. Anteriorly are 
four proportionally large tentacles. Two vibratile grooves (nuchal organs) posteriorly. 
Buccal segment with a pair of tentacular cirri; second segment has two pairs with 
bristles ; third segment carries the fourth pair, and a rudiment of a foot with bristles and 
a minute ventral cirrus. 

Body four inches long and upwards, terminating in two fusiform cirri. From the third 
to the last segment the middle of the dorsum is marked by green or blue bands, parallel 
and longitudinal, agreeing the one with the other. Proboscis papillose anteriorly and 
with eight large papillae at the entrance to the stomach. The large cordiform and 
imbricate dorsal cirri are throughout of a fine red colour. These organs are rounded, 
cordate anteriorly, and carried on a short ceratophore, broadly cordate and pointed in the 
middle of the body, cordate and still more pointed as well as smaller posteriorly. From 
a median streak extending from the ceratophore parallel lines radiate to the circumference, 
and many have touches of dark pigment. Setigerous region a short blunt and bifid 
cone supported by a single translucent spine, and the bristles are slightly curved towards 
the end of the shaft, which is truncated and dilated, a little bevelled, and has traces 
of spinous strias ; terminal piece short or of moderate length, finely serrated, tapered and 
often bent at the tip. Ventral cirrus about a fourth the size of the dorsal, reniform, 
somewhat more pointed posteriorly. 

Synonyms. 
1888. Phyllodoce (Carobia) rubiginosa, De St. Joseph. Ann. Sc. Nat., 7 e ser., t. v, p. 282, pi. xi, figs. 

141—143. 
1904. „ rubiginosa, Allen. Journ. M.B.A., n.s., vol. vii, p. 223. 

Habitat— This finely coloured species was first found by De St. Joseph in dredged 
material at Dinard, and subsequently by Allen in dredgings from various grounds near 
Plymouth. 

Head (Plate XL VII, fig. 4) small, rounded, inserted on the buccal segment, 
not indented posteriorly, with two large eyes. Anteriorly are four proportionally large 
tentacles. De St. Joseph describes two vibratile grooves posteriorly, as in other 
Phyllodocidas, and which he thinks analogous with the nuchal organs of Syllids and 
Eunicids. The buccal segment bears a pair of tentacular cirri; the second segment has 

1 ' Annulata Danica/ i. Kjobenhavn, 1879. 



PHYLLODOCE VITTATA. 93 

two pairs with bristles ; the third segment carries the fourth pair, accompanied by a 
rudiment of a foot with bristles and a minute ventral cirrus. 

Body about four or more inches in length, little tapered anteriorly ; much more 
distinctly posteriorly where it terminates in two fusiform cirri (Plate LVIII, fig. 25) 
which are broad at the base — tapered distally. In this case, however, the tail appears 
to be undergoing reproduction. The cirri are not compressed, and from the third to the 
last segment the middle of the back is marked by green or blue bands, parallel and 
longitudinal, agreeing the one with the other. 

The proboscis is papillose anteriorly, and has eight large papillas at the entrance to 
the stomach. 

The dorsal cirri (Plate LXVIII, fig. 9) are throughout of a fine red colour, large, 
cordiform, and imbricate. From a median streak, which extends from the short conical 
pedicle, parallel lines radiate to the circumference. The setigerous region forms a 
short blunt bifid cone supported by a single translucent spine, and the translucent 
bristles (Plate LXXVII, figs. 8 and 8 a) which project from it are slightly curved towards 
the tip of the shaft, which is somewhat abruptly dilated, truncated, a little bevelled, and 
shows only traces of spinous striae. The terminal piece is short or of moderate length, 
finely serrated, tapered, and often bent at the tip. The ventral cirrus is about a fourth 
as large as the dorsal, and is more or less renif orm with a tendency to be more distinctly 
pointed in the posterior part of the body. 

This is a southern form of great beauty, and may be a young stage or a variety of a 
better known form, such as Phyllodoce Paretti. 

The only British examples of this form are those forwarded from Plymouth by 
Dr. Allen. So far as shown by an examination of the living specimen, from which the 
figure (Plate XLVII, fig. 4) was taken, it would seem to approach a variety of Phyllodoce 
Paretti, especially as the dorsal cirri, the ventral cirri, and the bristles resemble each 
other closely. Both have the tips of the shafts of the bristles somewhat truncated and 
spinous, and the terminal pieces short, and while the bristles in the larger examples of 
P. Paretti are more deeply tinted brownish those in P. rubiginosa are also slightly tinted. 
Unfortunately the enlarged drawing of the bristle given by De St. Joseph is not sufficiently 
detailed for accurate diagnosis. Besides, all the examples of the latter sent from Plymouth 
have been small, so it is probable that slight differences may occur during growth. The 
shape of the head and eyes, the large size of the tentacles, and the shape of the tentacular 
cirri more or less correspond. Further examination of the life-history of each form is 
therefore very desirable. 



6. Phyllodoce vittata, Ehlers, 1864. Plate LXVII, figs. 10— 12— feet; PI. LXXVII, 

fig. 9 — bristle. 

Specific Characters. — Head short and rounded, with rather short and thick tentacles 
and tentacular cirri. Eyes two, comparatively large. Body 1J inches long, dotted with 
brownish pigment, which also occurs on the anterior lamellae. Dorsal division of the foot 
carrying an ovate cirrus of considerable size with smoothly rounded margins. Spinigerous 



94 GENETYLLIS. 

process large, obliquely truncated below the notch, and extending beyond the ovato- 
lanceolate ventral cirrus. Bristles comparatively long, slightly enlarged at the tip, and 
with one or two curved and prominent spikes. 

Synonyms. 

1864. Phyllodoce vittata, Ehlers. Borstenw., i, p. 150, Taf. vi, figs. 7 — 14. 

1865. ,, „ Carrington. Annel. Southport., p. 5. 

Habitat. — Dredged in 100 fathoms in St. Magnus Bay, 1869 (Dr. J. Gr. Jeffreys). 

Head short and rounded in spirit ; eyes comparatively large ; tentacles and tentacular 
cirri somewhat short. 

Body about 1 J inches in length, dotted with brownish pigment-grains on the dorsum 
and on the adjacent anterior lamellae. Ventral surface pale. 

The typical foot (Plate LXVIII, figs. 10, 11 and 12) bears dorsally a considerable 
broadly-ovate cirrus, its upper border never showing traces of angularity or truncation. 

The spinigerous process is large, obliquely truncated below the notch, and carries a 
series of rather long slender bristles (Plate LXXVII, fig. 9) with a slight curvature 
towards the distal end of the shaft, which is less dilated than usual and provided with one 
or two large and various smaller spines on each side. The terminal blade is long, curved, 
finely tapered, and serrated. 

The ventral cirrus is ovato-lanceolate, the tip remaining more or less blunt through- 
out. It is shorter than the spinigerous process of the foot. 

There is some doubt as to the identity of this with Elders' form, which came from 
Quarnero in the Adriatic, but until other and more complete examples are obtained it 
may be provisionally placed under that species. 

This appears to differ from any other British Phyllodoce. 

The eyes are peculiarly large, head short and rounded, and the tentacular processes 
short and thick. The tentacles at the tip of the snout are absent, so that this descrip- 
tion applies to the tentacular cirri. Dorsum dotted over with brownish pigment-grains, 
ventral surface pale. The anterior cirri also have some brownish grains. 

The dorsal cirri are all modifications of the leaf-shape, not showing any angular or 
truncated border. The ventral cirrus is somewhat lanceolate, and rather shorter than the 
bristle-papilla, which has a series of long and strong bristles, with very long and minutely 
serrated tips, and two conspicuous spikes at the swollen end of the shaft. 

Grube states (1879) that the group to which this form (his B 2) belongs has on the 
peristomial segment one tentacular cirrus ; on segment 2 — two tentacular cirri ; and on 
segment 3 — one. 

Genus XXXV. — Genetyllis, Malmgren, 1865. 

Head rounded ovate, with two eyes of considerable size (as in Notophyllum) . The 
four tentacles are about the same length as in the latter, and finely pointed. The 
tentacular cirri have a similar shape, the two anterior being shorter than the posterior. 

Body flattened, linear, slightly narrowed anteriorly and more distinctly so posteriorly. 



GENETYLLIS LTITEA. 95 

The dorsal cirri are large, more or less vertical, partly covering the dorsum and the feet, 
though to a less extent than in the previous genus. The inferior cirrus, which is behind 
the bristles, is vertical. Both are less developed than in Notophyllum. Bristles compound, 
more slender than in the latter, but conforming to the same type. The dorsal spine and 
scanty bristles of the former genus have disappeared. 

This is placed by Grube (1879) under the Phyllodocidas proper, and not with the 
Eulalia-group. Segment 2, like segment 3, smaller than the peristomial segment and 
with two tentacular cirri. Levinsen observes that the eyes and the nuchal appendages 
are large whilst the dorsal cirrus covers the foot. 

In Genetyllis ocidata of the ' Challenger ' the oblique muscles not only pass down by 
the sides of the nerve-cords, but certain fibres, probably from the vertical, pass above 
them to the opposite side. The great ventral spine in this form is black. 



1. Genetyllis lutea, Malmgren, 1865. Plate LXVIII, figs. 18 and 19— feet; Plate LXX VII, 

fig. 13 — bristle. 

Specific Characters. — Head rounded — ovate, with two nearly circular black eyes of 
variable size. Tentacles subulate and tapered. The tentacular cirri also have finely 
pointed tips. Ventral surface of the prostomium bipapillose with a smaller posterior lip 
than in Notophyllum. Body linear with transverse lines in each segment. The superior 
division of the foot carries a somewhat reniform cirrus, or rather a lamella almost semi- 
circular anteriorly with the inner edge reduced, broadly reniform, but with a dorsal 
peak, and with projecting inner lappets in the middle of the body. The spinigerous 
region is bluntly conical, bears a yellow spine, and has more slender bristles than in 
Notophyllum, the distal end of ,the shaft being dilated and obliquely striated, with a series 
of spikes on the ridge. The terminal blade is of moderate length, obliquely striated, curved, 
finely tapered, and the edge is minutely serrated. The ventral cirrus is obovate with a 
slight peak superiorly, and is soldered to the posterior surface of the foot by its inner 
edge. The colour in life is stated to be fine yellow. In spirit it is a dull yellow. 

Synonyms. 

1865. Genetyllis lutea, Malmgren. Nord. Hafs-Annul., p. 93,, Tab. xiv, f. 32. 

1867. „ „ Idem. Annul. Polych., p. 20. 

1869. „ „ Mcintosh. Kept. Brit, Assoc. (1868), p. 337. 

1874. „ „ Idem. Ann. Nat. Hist., ser. 4, vol. xiv, p. 196. 
„ „ „ Malm. Op. cit. Groteb., p. 79. 

1875. „ „ Mcintosh. Invert, and Fishes St. Andrews, p. 119. 
1879. „ „ Tauber. Annul. Danic, p. 86. 

1883. „ „ Levinsen. Yidensk. Meddel. Foren. Kjobenh., p. 205. 

1893. „ ,, Idem. Vidensk. ud. "Hauchs," p. 326. 

1901. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. viii, p. 223. 

Habitat — From the deep sea fishing boats, St. Andrews (E. M.) ; in 100 fathoms, 
St. Magnus Bay, Shetland, and off the Hebrides (Dr. Grwyn Jeffreys). 



96 GENETYLLIS CITRINA. 

Norway (Canon Norman). 

Head rounded, ovate, narrower in front, with two nearly circular black eyes, often 
with a lens-like thickening ; the size of the eyes, how T ever, differs in a series of specimens, 
showing that too much weight need not be placed on this character. It is probably 
connected with sexual changes. The tentacles spring from the anterior part of the 
prostomium, and are subulate organs with finely tapered tips. The two shorter and 
two longer tentacular cirri have also finely pointed tips. The ventral surface of the 
prostomium is bipapillose, with a groove leading backwards to the mouth. The latter 
has a smaller posterior lip than in Notophyllum. Occasionally white grains occur behind 
the eyes, forming a kind of transverse bar. 

Body linear, rounded dorsally and flattened ventrally. Dorsally the segments have 
transverse lines, while ventrally a double line of slight elevations maps out a central area 
which is but little depressed. No specimen retained a caudal process. Of a uniform dull 
yellowish colour, paler beneath; the dorsal cirri, especially in front, are darker, from 
brownish pigment-grains. 

The superior division of the foot (Plate LXVIII, fig. 18 [tenth] and 19 [fiftieth]) is 
considerably shorter than in Notophyllum, and has no prolongation upward and inward, 
or trace of spine or bristles. It carries a somewhat reniform cirrus, which varies a little 
in outline according to the region of the body, being almost semicircular anteriorly, with 
the inner edges reduced, broadly reniform, but with a dorsal peak, and with projecting 
inner lappets in the middle of the body. 

The inferior division is represented by a bluntly conical, bifid, free process bearing 
the bristle-tuft and a yellow spine. 

The bristles (Plate LXXVII, fig. 13) have rather long translucent shafts, more slender 
than in Notophyllum poly noides, but also with a backward curve towards the tip. The distal 
end of the shaft is dilated and marked with fine striae obliquely directed downward and 
backward, whilst the free edge has a series of spikes. The terminal process is of moderate 
length, curved, and finely tapered, the edge being minutely serrated. Fine striae also run 
obliquely downward and backward to the dorsal rib at the broader or inferior part of the 
process. 

The inferior cirrus is obovate or oval, with a slight peak superiorly, is soldered 
to the posterior base of the division by its inner edge, and is thus kept vertical. 

This species seems to be an inhabitant of water of some depth. No food of moment 
seemed to be in the alimentary canal. 

Specimens from Shetland in July had many well-developed ova. 



2. Genetyllis citrina, Mcintosh. Plate LXVIII, figs. 20 and 21— feet ; Plate LXXVII, 

fig. 14 — bristle. 

Specific Characters. — Head rounded ovate, with two rather large black eyes ; tentacles 
probably similar to those of G. Intea, but they were absent; tentacular cirri similar. 
Body about three inches long, much tapered anteriorly. Posteriorly two short caudal cirri. 
Dorsal surface convex, ventral depressed in centre with an elevated ridge on each side. Of 



GENETYLLIS HIBERNICA. 97 

a brilliant chrome-yellow throughout. Dorsal cirrus unequally cordate, broadly lanceolate, 
longer posteriorly than anteriorly, and marked by a series of lines and reticulations from 
a central rib. The ceratophore is a massive short and bluntly-conical process. Imbricate 
cirri borne more or less horizontally, leaving the centre of the dorsum bare. Setigerous 
region short, bifid, supported by a black spine, and carrying shorter bristles than in 
G. lutea, with shorter terminal processes. 

Habitat. — Procured on a stone coated with corallines, a yellow sponge, and several 
Ascidians brought up by a fisherman's hook in the Minch, August, 1865. 

Head rounded ovate, with two black eyes of considerable size. The four tentacles 
had been lost before minute examination, but they probably conform to those of the 
previous species. The tentacular cirri are similarly arranged — two shorter anterior and 
two longer posterior. 

Body about three inches in length, much tapered anteriorly. Posteriorly it terminates 
in two caudal cirri. The dorsal surface is convex, the ventral marked by the two raised 
lines on each side of a slightly depressed central area. The proboscis is enclosed. 

The entire animal is of a most brilliant chrome-yellow colour, deepest on the middle 
third, which here and there shows blackish-brown patches on the lamellae. It tinged the 
water with a yellowish mucus, and also dyed the spirit in which it was immersed of the 
same hue. 

The dorsal region of the foot (Plate LXVIII, figs. 20 and 21) has a massive short 
bluntly-conical process, devoid of spine or bristles, and bearing the unequally cordate 
lamella, which is marked by a series of lines and reticulations from a central rib. The 
imbricate lamellae are borne more or less horizontally along the sides of the dorsum, 
leaving the central part bare. The semicircular gap at the base of the lamella fits the 
rounded extremity of the division to which it is attached. The short setigerous region is 
bifid at the tip, and supported by a black spine and a group of bristles, shorter than in 
G. lutea (Plate LXXVII, fig. 14), and with translucent shafts, slightly bent, and with a 
dilated distal end, which has a few spikes on each side. The terminal process is finely 
tapered, and shorter than in G. lutea. The edge shows no distinct serrations, though the 
adherence of particles would indicate them. Attached to the ventral and posterior parts 
of the region is an irregularly reniform lamella (ventral cirrus), vertical in position. The 
inferior border is rounded, but the superior is truncated and with a tendency to a point 
at the upper and outer angle. The cirri, both superior and inferior, vary a little in shape 
throughout the body. 

The stone on which it was brought up had a yellowish sponge on part of its surface, 
but the connection between the Annelid and it is unknown. It crawled actively amongst 
the Ascidians and other growths to escape capture. 

3. GrENETYLLis hibernica. Plate LVIII, fig. 8— head ; Plate LXIX, figs. 1 and 2— feet; 

Plate LXXVII, fig. 15— bristles. 

Specific Characters. — Head rounded ovate; eyes two, black and of considerable size; ten- 
tacles and tentacular cirri as in allied forms. Body linear, about one inch long, terminating 

70 



98 ETEONE. 

in an anal papilla—probably flanked by two cirri. Dorsal surface bluish in spirit, ventral 
surface pale cinnamon, both being iridescent. Dorsal and ventral cirri of a cinnamon 
colour. Dorsal region of the foot less than in G. lutea, the dorsal cirrus ovato -lanceolate 
in front or ovate posteriorly, veined and carried horizontally, the tip more acute than in 
the former, but less acute than in G. citrina. Setigerous region has a single yellow spine. 
Bristles have a well-marked dilatation at tbe end of the shaft and with minute spines 
guarding the base of the terminal process which is finely tapered. Ventral cirrus borne 
vertically, reniform. A mamilla (papilla) occurs at the inner border of the foot. 

Habitat— Coast of Galway, Dr. E. P. Wright, 1868. 

Head (Plate LVIII, fig. 8) rounded ovoid, with two black eyes of considerable size. 
Tentacles and tentacular cirri as in allied forms. 

Body linear, about an inch long, slightly tapered in front, more so posteriorly, 
and terminating in the anal papilla, which was probably flanked by two cirri. The 
dorsum in spirit has a bluish tint, and the ventral surface a pale, cinnamon hue — both 
being iridescent. The lamellae were of a cinnamon colour. 

The dorsal region (pedicle) of the foot (Plate LXIX, figs. 1 [tenth] and 2 
[fortieth]) is less than in G. lutea — forming a short truncate process supporting the 
cirrus, which is comparatively large and cordate, the "bite" at the base being less than 
in the two foregoing species. The lamella has the same leaf -like series of veins, but on 
the whole has a more acute tip. It is carried somewhat horizontally, so that in series an 
imbricate arrangement is produced. The lamellae are rather more acute posteriorly. 

The bifid setigerous process is of average length, and bears a pale spine and a series 
of translucent bristles, the shafts of which are slightly curved and have a well-marked 
dilatation at the tip, with minute spines guarding the base of the terminal process, which 
is finely tapered (Plate LXXVII, fig. 15) and by no means long. The ventral cirrus is 
borne vertically, and has a somewhat reniform outline. A distinct mamilla occurs at the 
inner border of each foot. 

So far as can be observed this does not appear to be the young stage of either of the 
foregoing. 



Genus XXXVI. — Eteone, Savigny, 1820. 

Head elongated antero-posteriorly or somewhat trapezoidal in spirit, with or without 
eyes ; two pairs of short tentacles ; two pairs of tentacular cirri, the first on the first 
segment, the second on the following. Body devoid of metallic lustre ; segments similar. 
Proboscis with cordate or with pointed papillae ; an obtuse rigid papilla on each side of the 
aperture in extrusion. Anal cirri leaf -like. Feet simple ; dorsal cirrus sessile, simple, 
and leaf -like, much larger than the ventral cirrus. Dilated ends of the shafts of the 
bristles with large hook-like spines and short terminal blades. 

Small Annelids — chiefly characteristic of the northern seas, and sparingly distributed. 

The body-w^all of JEteone sjpetsbergensis (Fig. 43) is distinguished by the massive 
longitudinal muscles — both dorsal and ventral. The dorsal form a broad arch in section 



ETEONE. 



99 



with a rounded thick inferior border on each side ; whilst the ventral have a tendency 
to a crescentic arrangement of each mass, for the outer border (in section) curves 
inward dorsally. 

The alimentary canal has a thick coat of longitudinal muscular fibres exteriorly — 
apparently only a basement-membrane with a few circular fibres intervening between 
it and. the nucleated outer cellular layer. 

The oblique muscles are of great strength, and thus in section the ventral median 
line is deeply grooved. They meet in the mid-ventral line in Eteone spetsbergensis, but 
they show intervals in E. picta. The nerve-area is thus generally pushed further inward 
than in Phyllodoce, and both trunks and ganglia have a firm sheath with longitudinal 
fibres. The interganglionic cords are separate. At intervals strong transverse bands 
of muscle pass above the cords from side to side. The circular muscular fibres are 
largely developed in the dorso-lateral regions. 

Ehlers (1864) characterises the genus thus: Body elongated; head with four 

dv 




Fig. 43. — Section of Eteone spetzbergensis, to indicate the muscles. The same letters are used as formerly. 

tentacles ; two pairs of tentacular cirri ; segments similar with simple feet ; dorsal cirrus 
small, leaf-like, with the base resting on the dorsum, much larger than the ovate ventral 
cirrus; two leaf -like anal cirri. Claparede (1868) includes Mysta in this genus, as also 
does Levinsen (1883). 

Grube (1879) mentions that the tentacular cirri are short, as also the tentacles, 
which are only a third the length of the short trapezoidal head. Some genera lose 
the dorsal cirri in the second segment. In others segment 1 has one tentacular cirrus, 
segment 2 also one. Eyes small, often mere points, and frequently disappear in spirit. 
Proboscis with two cordate papillge ; upper surface smooth, or covered with pointed 
papilla, or ringed, or with sparsely distributed papillae having minute processes (hook- 
like). Body does not show metallic lustre, and when coloured has specks. Small as 
a rule; characteristic of northern and arctic seas and sparingly distributed. 

Grube groups the species of Eteone as B. — tentacular cirri on first segment; B s . 

only one pair of eyes. 

(a) Head as broad or broader than long. E. arctica, E. picta, E. flava. 

(b) Head more or less longer than broad. E. longa, E. pusilla, E. spetsbergensis. 
Gravier 1 mentions a strong band of transverse muscular fibres from side to side 

1 ' Bull. Sc. Fr. Belg./ t. xxix, p. 368, pi. xvii, f. 13. 



100 ETEONE PICTA. 

below the gut in Eteone foliosa, De Quatrefages, and apparently in the line between the 
feet (as it is not figured). 

In May (1903) Leschke found larvse of Eteone in the Bay of Kiel. 



1. Eteone picta, De Quatrefages, 1865. Plate XLV, fig. 3; Plate LXIX, figs. 3 and 4— 

feet; Plate LXXVII, fig. 22— bristle. 

Specific Characters. — Head pale, flattened, shovel-shaped, with an evident constriction 
opposite the eyes, in front of the swellings from which the tentacular cirri arise, and 
another a little behind the tip. Eyes black, situated rather near each other at the 
posterior third of the Tiead. Four short tentacles are attached anteriorly. Two subulate 
and short tentacular cirri, which are less than the transverse diameter of the head (in pre- 
servation). Body from one to three inches long, firm, somewhat flattened, gently dilating 
behind the head, and again diminishing a little towards the tail, where it ends in two 
ovoid yellowish cirri. Ground- colour pale yellowish, and each segment laterally has a 
reddish-brown or rose-red spot opposite the foot, and two others on each side of the 
middle line. Thus there are four rows in addition to the spots on the dorsal cirri. A 
patch of yellowish pigment occurs between the foregoing spots in the anterior third of the 
body. The posterior third is in some pinkish — apparently from the ova. Dorsal cirrus 
broadly lanceolate or somewhat conical, and firmly fixed to the peduncle by almost the 
entire base. Setigerous process is bifid and comparatively short. The shaft of the stout 
bristle is dilated, and, antero-posteriorly, has two hook-like spines of large size (laterally a 
median curved spine and a smaller in front and behind). The terminal serrated blade is 
comparatively short, broad at the base, and rapidly tapered to a fine point. Ventral 
cirrus bluntly lanceolate, reaching almost to the tip of the setigerous process, but 
shrinking within it at the fiftieth foot. It is borne on a distinct basal process (cerato- 
phore) which projects from the setigerous lobe of the foot. 

Synonyms. 
1865. Eteone picta, De Quatrefages. Annel., h, p. 147, pi. 7 bis, f. 18 — 23. 

1874. „ „ Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 197. 

1875. „ „ idem. Invert, and Fishes St. Andrews, p. 120. 

„ „ „ Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., t. ii, p. 62. 

1879. „ „ Tauber. Annul. Danic., p. 91. 

1890. „ „ Giard. Bull. Sc. Fr. Belg., t. xxii, p. 77. 

1897. „ ,, Gravier. Ibid., t. xxix, p. 308, pi. xvi, f. 14; pi. xvii, f. 5—7 • pi. xviii, f. 9—11 ; 

pi. xix, f. 20; pi. xx, f. 10, 11; pi. xxiii, f. 2, 
1904. „ „ Allen. Journ. Mar. Biol. Assoc, n. s. vol. vii, p. 224. 

Habitat. — Not uncommon under stones between tide-marks and in fissures of rocks, 
Guernsey and Herm ; Whitby (Dr. Carrington) ; deep water off St. Andrews Bay, and 
in the stomach of the cod and haddock, St. Andrews (E. M.). 

Head pale, flattened, shovel-shaped, with an evident constriction opposite the 
eyes — in front of the bosses from which the tentacular cirri arise, and another occurs a 
little behind the tip. The eyes are black and situated rather near each other at the 



ETEONE PICTA. 101 

posterior third of the head. Four short tentacles are attached anteriorly, and two 
tentacular cirri spring from the peristomial segment — one laterally and one dorsally. 
They are subulate and short, less than the transverse diameter of the head (in preserva- 
tion). 

Body from 1 to 3 inches in length, firm, flattened, though more convex dorsally, 
gently dilating behind the head, and again diminishing, but not much, towards the tail, 
where it terminates in two ovoid yellowish cirri. The ventral surface has a median 
furrow. The colour is pale yellowish, especially towards the posterior area of the 
intestinal tract. Each segment has a reddish or madder-brown spot laterally opposite the 
foot, and two others on each side of the middle line. Thus there are four rows, besides 
those made by the spots on the dorsal lamellae. A patch of yellowish pigment occurs 
between the foregoing transverse groups of spots in the anterior third of the body. In 
some specimens the spots are of a deep rose-red, which, with the general iridescence of the 
surface, gives a striking aspect. The posterior third occasionally is pinkish, apparently 
from the ova. The under surface is pale. 

The foot (Plate LXIX, figs. 3 [twelfth] and 4 [thirtieth]) has dorsally a broadly 
lanceolate or somewhat conical cirrus, which stands stiffly outward with a slight slope 
backward. It is firmly fixed to a prominent papilla by almost the whole breadth of its 
base and is not easily detached. Both papillae and lamellae increase in size from before 
backward, the difference between the tenth and the fiftieth foot, for instance, being 
marked. The hypodermic streaks form a fan-like arrangement from the central basal 
region. The setigerous process is bifid, comparatively short, and bears a series of stout, 
slightly curved bristles (Plate LXXVII, fig. 22), that is, the bristles are curved, when 
viewed laterally, straight when viewed antero-posteriorly. The distal end of the shaft is 
dilated, and antero-posteriorly presents two curved hook-like spines of large size, while 
laterally it shows a median and a smaller curved spine in front and behind. The terminal 
blade is comparatively short, broad at the base, and rapidly tapers to a fine point. It 
is minutely serrated on the anterior edge, the points being directed distally. 

The inferior cirrus is bluntly lanceolate, extending at the tenth foot almost to the 
tip of the setigerous process, but at the fiftieth shrinking within it in lateral view. It is 
borne on a distinct basal process which projects from the setigerous lobe of the foot. 

Reproduction. — Some of the examples obtained between tide-marks at Guernsey 
and Herm in July and August contained masses of minute eggs, but no ripe form was 
seen. 

Habits. — They are very restless Annelids, darting through the water by spasmodic 
coiling of the body as in other forms, and partial to boring in sand. They often coil 
themselves into a ball, the posterior end being rolled concentrically. Consequently, most 
of the examples killed in spirit are coiled or twisted in a characteristic manner. 

The Meone fucata of M. Sars 1 approaches this form very closely, though he mentions 
only three purplish spots on the dorsum — a lateral and two median. The shortness of 
the upper tentacular cirrus, however, may be diagnostic of the Norwegian form. 

(Ersted's ffleone maculata? is another closely allied form. 

1 c Bidrag. Christ. Fauna/ iii, p. 26, 1873. 

2 'Annul. Dan. Consp./ p. 29. 



102 ETEONE ARCTICA. 

It is difficult to know what the Nereis lineata of Montagu 1 is. It may refer to this 
species. 

2. Eteone arctica, Malmgren, 1867. Plate LVIII, fig. 9 — head; Plate LXIX, figs. 8 and 

9— feet ; Plate LXXVII, fig. 18— bristle. 

Specific Characters. — Head (in spirit) conical, with the diameters nearly equal, the 
antero-posterior, however, being the longer. No eyes. Tentacles small and tentacular 
cirri short. Body elongated, with elevated lateral areas at the bases of the feet in each 
segment. Two somewhat ovate anal cirri. Proboscis, in extrusion, is papillose at the tip 
and has minute papillae apparently in rows on its surface, but the specimens are imper- 
fectly preserved. Dorsal cirrus ovate and separated by an interval from the setigerous 
lobe. The latter is somewhat conical and bears bristles with two strong though not 
long spines, besides smaller spikes on the ridges of the dilated ends of the shafts. 
Terminal sabre wide at the base, but rapidly tapers to a fine point. The ventral cirrus 
is lanceolate, and extends outward about as far as the tip of the setigerous lobe. 

Synonyms. 

Malmgren. Annul. Polych., p. 27, Tab. ii, f. 12. 

Ehlers. Sitz. Phys. Med. Soc. Erlangen, iii, p. 79. 

(?) Mcintosh. Ann. Nat. Hist., ser 4, vol. xiv, p. 197. 

idem. Invert, and Fishes St. Andrews, p. 120. 

idem. Trans. Linn. Soc. Zool., i, p. 502. 

Wiren. Ohsetop. c Vega' Exped., p. 399. 

Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 208. 

Habitat. — Southport sands (Dr. Carrington, 1866). 

Greenland (' Valorous ') ; Spitzbergen (Ehlers) ; Siberia and Behring's Strait (Wiren). 

Head (Plate LVIII, fig. 9) conical with the diameters nearly equal, though in one 
the antero-posterior is the longer. No eyes are visible. The tentacles are small, and the 
tentacular cirri short. 

Body elongated, with the lateral areas in each segment forming fillets at the base of 
the foot. Posteriorly it terminates in two somewhat ovate cirri. 

The exserted proboscis shows papillae at the tip, and minute papillas in lateral rows 
on its surface, but the organ is imperfectly preserved and only partially extruded. 

The foot carries dorsally an ovate cirrus, separated by a considerable interval from 
the setigerous process. It thus agrees with the lamella of E. arctica. The setigerous 
lobe is somewhat conical and has a series of bristles with a distinct curvature below the 
enlarged tip, which has two strong though not long spines besides smaller serrations on 
the ridges. The terminal process is wide at the base but rapidly tapers to a fine point. 
The edge is apparently minutely serrated, but the points are seen with difficulty. The 
ventral cirrus is lanceolate and extends outward about as far as the tip of the setigerous 
lobe. 

Though the eyes are absent, this form would seem to come nearest E. arctica, but 

1 l Trans. Linn. Soc.,' vol. vii, p. 83, 1804. 



1867. 


Eteone arch 


1871. 


)> > 


1874. 


3) ) 


1875. 


}) J 


1878. 


)1 ) 


1883. 


>> y. 


>) 


)) >, 



ETEONE ARCTICA. 103 

there are certain differences, such as the shape and length of the ventral cirrus, which 
Malmgren states is sub-obtuse and distinctly longer than the setigerous lobe. It differs 
from E. longa l in the shape of the dorsal cirrus and other points. The condition of the 
proboscis prevented the determination of the number of the terminal papillas. 



A form which has been described as a separate species, though apparently only a 
variety of Meone arctica, merits a special description from its characteristic structure. 



Eteone arctica var. Robertiana. Plate LXIX, figs. 8 and 9 — feet. 

Head flattened, bluntly conical, with a distinct furrow on each side. Two subulate 
tentacles. ~No visible eyes in the preparation. The fused cephalic and peristomial 
region is separated by a constriction from the succeeding segment. The tentacular cirri 
are little tapered and considerably shorter than the transverse diameter of the region. 
Body of the typical form, terminating in two linear fusiform or almost cylindrical cirri. 
Pale, slightly iridescent. Dorsal cirrus ovoid at the tenth foot and shorter than the 
setigerous process, ovato-lanceolate at the thirtieth and projects further outward than 
the setigerous process. Still further backward it becomes more acutely lanceolate and 
the peduncle is longer. Setigerous lobe largest anteriorly, bifid, and carries bristles, the 
shafts of which have dilated tips with a larger and a smaller curved hook and minute 
spines passing downward to the shoulder. The terminal blade is of average length, is 
broad at the base, tapers gradually to a fine point and has a serrated edge. The ventral 
cirrus is lanceolate, about as long as the setigerous process at the tenth foot, and some- 
what longer and more acuminate posteriorly. 

Synonyms. 

1874. Eteonella Robertianse, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 197. 

1875. „ „ Idem. Invert, and Fishes St. Andrews, p. 120. 

Habitat. — Found when digging for littoral Annelids near low-water mark at the 
West Rocks, St. Andrews (R. M.). Other pale specimens were sent alive (E. M.). 

Head differs from that of the ordinary form by the more evident separation of the 
fused cephalic and peristomial segments — a constriction dividing them from the succeeding 
region. The head has a flattened bluntly-conical form with a distinct furrow on each 
side, and with two short subulate tentacles on each side anteriorly. Two minute but 
distinct papillse occur dorsally in the centre of the peristomial segment immediately behind 
the head. Posteriorly the peristomial segment seems to form a boss at each side, from 
the posterior part of which two slender and little tapered tentacular cirri proceed. In 
the preparation they are considerably shorter than the transverse diameter of the region. 

1 Some examples of E. longa from Greenland have many small white ova, showing a distinct 
micropyle, attached to the bristles and other parts. 



104 ETEONE SPETZBERGENSIS. 

The body is most dilated anteriorly, only narrowing a little immediately behind the 
head. It then tapers gradually towards the tail, which is terminated by two linear, 
fusiform, or almost cylindrical cirri. It is rounded dorsally and flattened ventrally, and 
is pale with a slight iridescence throughout. 

The foot (Plate LXIX, figs. 8 [10th] and 9 [70th] ) has dorsally a more or less ovoid 
cirrus which at the tenth foot is comparatively small, and truncated at the base, where it 
abuts on the peduncle. The setigerous process is greater in diameter and projects 
further outward. At the thirtieth foot it has become ovato-lanceolate, and extends 
considerably further than the setigerous lobe. Behind the foregoing it is more acutely 
lanceolate, and its peduncle is longer. The streaks in the hypoderm of the organ have a 
fan-like arrangement. The setigerous lobe is largest anteriorly, is bifid, and has a series 
of pale slightly curved bristles (Plate LXXVII, fig. 18), the shafts of which have slightly 
dilated tips with a larger and smaller curved hook and minute spines passing downward 
to the shoulder. The terminal process is broad at the base, tapers gradually to a fine 
point, and has a serrated edge. It is of average length, The ventral cirrus is lanceolate, 
about as long as the setigerous lobe at the tenth foot, and somewhat longer and more 
acuminate posteriorly. 

This species seems to approach most closely to Eteone arctica, Mgm., from which it 
differs in the absence of eyes (which, however, occasionally disappear in spirit in other 
examples), in the ovato-lanceolate superior cirrus, and in the somewhat narrow lanceolate 
shape of the ventral cirrus, instead of the subobtuse lamella of E. arctica. Malmgren's 
figure (Plate II, fig. 12, c'), however, shows a closer approach than his description. The 
greater size and leaf-like condition of the dorsal cirri in E. Robertiana in contrast 
with E. arctica is also a distinguishing feature. 

On the whole it would appear that Eteonella Robertiana, is only a well-marked variety 
of Eteone arctica. 



3. Eteone spetzbergensis, Malmgren, 1865. Plate LXIX, figs. 5 and 6 — feet; PI. 

LXXVII, figs. 16-16 6— bristles. 

Specific Characters. — Head somewhat elongated, separated dorsally from the peri- 
stomial segment by converging lines ; bluntly rounded in front. Tentacles short, the upper 
in front of the ventral. Two pale eyes occur posteriorly, but they disappear in the spirit 
preparations, in which the head is also contracted into a short cone. Tentacular cirri 
short, the ventral being the longer. Body three and a half inches long, about equally 
tapered anteriorly and posteriorly, and terminating in two linear fusiform cirri (Malmgren). 
Several preserved examples show a brownish pigment-band on each side dorsally, within 
the bases of the feet. Proboscis in extrusion is clavate, with two median and two lateral 
rows of papillae, which begin as prominent organs and end in low transverse elevations. 
On the ventral aspect four longitudinal bands are also present, but they show only low 
transverse folds or wrinkles. The aperture is dorsal, somewhat triangular, and papillose. 
Dorsal cirrus at the tenth foot is sub-circular, but posteriorly the peduncle lengthens, and 
the organ becomes renif orm with the larger end external. The setigerous process increases 



ETEONE SPETZBERGENSIS. 105 

in size from before backward, and is nearly equally bifid, with a dorsal elevation 
posteriorly. The dilated ends of the shafts of the bristles have two unequal spines, and a 
spinigerous ridge on each side. Terminal blade broad inferiorly, tapers to a fine point, 
and has a serrated edge. The ventral cirrus is oblong-ovate at the tenth foot, and 
gradually increases in transverse diameter from this region backward. Its outer margin 
projects a little beyond the tip of the setigerous process. 

Synonyms. 
1865. Eteone spetsbergensis, Malmgren. Nord. Hafs-Annul., p. 102, Tab. xv, f. 38. 
1867. „ „ idem. Annel. Polych., p. 26. 

1874. „ andreapolis, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 197. 

1875. „ „ idem. Invert, and Fishes St. Andrews, p. 120. 

1879. „ spetsbergensis, Theel. K. Svensk. Vet. Handl., Bd. xvi, No. 3, p. 31, pi. ii, f. 21 and 22. 

„ „ ,, Tauber. Annul. Danic, p. 90. 

1901. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. viii, p. 223. 

1903. „ „ idem. Mar. Invert. S. Africa, vol. iii, p. 35. 

Habitat. — Tossed on shore on the West Sands, St. Andrews, after a storm in 
February. Under a stone on shell-gravel above low- water mark, Lochmaddy. 

Extends to Norway (Canon Norman) ; Canada, Dr. "Whiteaves (W. C. M.) ; and to 
Nova Zembla (Theel) ; South Africa (W. C. M.) ; Behrings Sea (Marenzeller) . 




Fig. 44. — Head of Eteone spetsbergensis, from life. 

Head, (Fig. 44) somewhat elongated, separated dorsally from the peristomial 
segment by converging lines, and bluntly rounded in front between the two short 
tentacles, the upper of which is the shorter, and placed anterior to the other. The 
tentacular cirri are also short, the inferior being the longer. In the preparation they are 
about half the transverse diameter of the peristomial segment. Two pale eyes (woodcut) 
occur at the posterior part of the head, but they disappear in spirit-preparations, in 
which the head is also contracted into a short cone. 

Body three and a quarter inches long, about equally tapered anteriorly and posteriorly 
(Malmgren), somewhat flattened, and with the segments marked in the typical manner. 
The tail is incomplete, but Malmgren states that it bears two linear fusiform cirri. The 
colour inclines to pale yellow anteriorly, especially in the lamellae, which are uniform in 
tint throughout. After the first third the intestine gives the body a pinkish or skin colour. 
The surface has a pearly iridescence, especially dorsally. The proboscis in extrusion is 
clavate, with two median and two lateral rows of papillae, which begin as prominent organs, 
and end distally in low transverse elevations. On the ventral aspect four longitudinal 
bands are also present, but they have only low transverse folds or wrinkles. The aperture 
of the organ is dorsal, and forms a papillose triangle. Gregarines w^ere common in the 
intestine of that from St. Andrews. Several examples from Canada show in spirit a well- 
marked brownish pigment-band within the bases of the feet on each side. 

71 



106 ETEONE PUSILLA. 

The foot (PL LXIX, figs 5 [tenth] and 6 [hundredth]) agrees with that described 
and figured by Malmgren as characteristic of E. spetsbergensis, though the close approach 
of several to the same type gives rise to some uncertainty. The dorsal cirrus is described 
as obliquely obovate, and such would fairly apply to it behind the anterior region, but, for 
instance, at the tenth foot (Fig. 5) it is nearly subcircular, and the peduncle is shorter 
than in the posterior feet, so that the lamella closely approaches or touches the setigerous 
lobe beneath. Its outer border extends considerably beyond the tip of the setigerous 
process. Though somewhat thick the fan-shaped arrangement of its hypodermic strands 
are visible. From the fiftieth to the hundredth foot (Fig. 6) and posteriorly, the peduncle 
lengthens, and the lamella becomes almost reniform — only the larger end of the organ 
is external. Malmgren's description, therefore, of obliquely ovate will apply best to 
those between the fiftieth and seventieth feet. The setigerous process likewise increases 
in size from before backward, is nearly equally bifid at the tip (the spine projecting in 
the split), and presents a dorsal elevation in the posterior feet. The pale bristles (Plate 
LXXVII, figs. 16, 16 a, 16 b) have slightly curved shafts, the tips of which have a 
shoulder, are dilated, and terminate in a hook-like spine on each side, one being larger 
than the other. A row of minute spines runs from the larger hook obliquely downward 
and backward. The terminal blade is broad inferiorly, and tapers to a delicate tip. The 
edge is serrated. The details of the tip of the shaft vary a little from front to tail. The 
variations in the bristles are shown in Fig. 16 from St. Andrews, Fig. 16 a from Gaspe 
Bay, Canada, and Fig. 16 b from Lochmaddy, North TJist. The inferior cirrus is ovate- 
oblong, more rounded (shorter) and deeper at the tenth foot, and gradually increases in 
transverse diameter from this region backward. Its outer margin projects a little beyond 
the tip of the setigerous process. 

Reproduction. — A fragment of what appears to be the same species, procured at 
Lochmaddy in August, contained the male elements well advanced. A Canadian example 
had large ova in the coelomic space and extending into the bases of the feet. It was 
procured in July. 

Habits. — A long, agile, restless annelid, constantly gliding to and fro, and pushing its 
flattened snout with promptness and facility through mud or sand. When on a suitable 
site an undulating motion takes place from before backward, apparently to assist in 
respiration. When irritated it coils itself into a ball and displays many contortions. 



4. Eteone pusilla, (Ersted, 1843. Plate LXV, fig. 6— head; Plate LXIX, fig. 7— foot; 

Plate LXXVII, fig. 17— bristle. 

Specific Characters. — Head spathulate — a blunt, elongated cone, with the short 
subulate tentacles at each side anteriorly, and separated antero -posteriorly from each 
other. Eyes indistinct, only a faint trace of one remaining. Tentacular cirri short and 
subulate, about half the diameter of the peristomial segment. Body two inches or more 
in length, normal in shape ; anal cirri (fide Malmgren) elongate linear-fusiform. Dorsal 
cirrus obliquely ovate, touching the setigerous process anteriorly (in the preparation), but 
posteriorly the longer pedicle carries it outward. Posteriorly the lamella is somewhat 



1843. 


Eteone pus r i 


1851. 


■>■> ) 


1865. 


■>•> ) 


■>•> 


Eulalia , 


}) 


Eteone „ 


1867. 


■>•) ) 


1874. 


y> j 


1879. 


j) i 


1890. 


)) } 


1896. 


>; ) 



ETEONE PUSILLA. 107 

rhomboidal. Setigerous process slightly bifid. The terminal dilatations of the shafts of 
the bristles have a long curved spine and several smaller ones on each side. The distal blade 
is of moderate length, broad and slightly bulged inferiorly, and tapering somewhat 
abruptly to a fine point. The edge is minutely serrated. Ventral cirrus irregularly 
ovate, and more or less pointed posteriorly. 

Synonyms. 

CErsted. Consp. Annul., p. 30, f. 84. 

Grube. Fam. Annel., pp. 57, 130. 

Malmgren. Nord. Hafs-Annul., p. 102, Tab. xv, f. 37. 

De Quatrefages. Annel., ii, p. 122 (?). 

idem. Ibid., ii, p. 151. 

Malmgren. Annul. Polych., p. 26. 

Malm. Op. cit., G-oteb., p. 81. 

Verrill. Check List, U. S. Comm. F. and F., (sep. copy) p. 8. 

Giard. Bull. Sc. Fr. Belg., t. xxii, p. 77. 

Michaelsen. Polych. deutsch. meere, p. 37. 

Habitat. — Dredged in 100 fathoms, St. Magnus Bay, Shetland, 1867 (Dr. Gwyn 
Jeffreys) . 

Canadian and American coasts. 

Head (Plate LXV, fig. 6) spathulate, forming a somewhat elongated blunt cone, with 
the short subulate tentacles at each side anteriorly and separated antero-posteriorly from 
each other. The eyes in the preparation are invisible, or only a faint trace of one remains. 
The tentacular cirri are also short and subulate, being little more than half the diameter 
of the post-cephalic segment. 

Body about two inches or more in length, nearly equally narrowed anteriorly and 
posteriorly, though the tip of the tail is the more attenuate. It is rounded dorsally, 
flattened ventrally, the latter surface having a median groove anteriorly. The anal cirri 
are absent, but, according to Malmgren, they are elongate linear-fusiform. 

The foot (Plate LXIX, fig. 7) bears an obliquely ovate cirrus, which at the tenth foot 
is more or less sessile, that is, its pedicle or ceratophore is shorter. The lamella, which 
has its greatest diameter external and the lower edge dependent, thus touches the 
setigerous process, whereas the longer pedicle, e. g. from the thirtieth to the seventieth 
foot, carries it further outward, a feature still more marked at the hundredth foot. The 
lamella in the latter has also become somewhat rhomboidal, the long dorsal and ventral 
edges being straight, the outer and inner rounded. From the thirtieth to the seventieth 
foot the process projects considerably beyond the tip of the setigerous region, but this 
condition is less marked at the hundredth foot, probably from the elongation of the 
latter. The setigerous process is somewhat bifid, and it has a series of slightly curved 
bristles (Plate LXXVII, fig. 17), the ends of the shafts being enlarged and furnished 
with a long curved spine and several smaller ones on each side. The distal blade is of 
moderate length, broad and somewhat bulged inferiorly, and tapered rather abruptly 
to a fine extremity. The edge is minutely serrated. The inferior cirrus is also irregularly 
ovate at the tenth foot, with a tendency to a point at the tip, which, with that of the 



108 MYSTA BAKBATA. 

superior lamella, projects beyond the setigerous region. The inferior lamella becomes 
more pointed externally from the thirtieth foot backward, and at the hundredth the tip 
has also a tendency to a point ventrally. 

A curious two-jointed parasite occurs on the dorsal lamella of the fiftieth foot. 

This species presents certain resemblances to the Eteone cxca of Ehlers, 1 from the 
6 Porcupine ' Expedition of 1869, but the latter has no eyes, and the tentacular cirri seem 
to be longer. The presence or absence of eyes, however, in spirit-preparation is an 
uncertain character. 

Eteone flava, Fabr., a form not uncommon in more northern waters, has not yet been 
found in British seas. What relationship the Eteone fucata of M. Sars 2 has to this or 
allied forms it is difficult to say. In all probability it is identical with one or other of the 
forms mentioned by Malmgren. 



Genus XXXVII.— Mysta, Malmgren, 1865. 

Head bluntly conical. Eyes two, conspicuous, situated at the posterior border. 
Tentacles short and subulate, as in Eteone. Tentacular cirri, two on each side, unequal, 
subulate, fixed to the first segment. Body linear, slightly flattened, tapered towards 
each extremity. Two anal cirri. Proboscis long, sub-cylindrical, thickened anteriorly, 
papillaB ovate-fusiform in linear rows. Foot with the dorsal cirrus sub-orbicular, seti- 
gerous lobe bifid ; ventral cirrus ovato-lanceolate. Bristles have the ends of the shafts 
dilated with a large curved spine, and a smaller at the tip with minute spines on the 
ridges. The terminal blade is wide beneath, tapers to a fine point, and has a serrated edge. 

This genus resembles Eteone, and by some is made a sub-genus. 

1. Mysta barbata, Malmgren, 1865. Plate LVIIL fig. 10— head ; PL LXIX, fig. 10- 
foot; PL LXXVII, fig. 19— bristle. 

Specific Characters. — Head (in spirit) bluntly conical, with two conspicuous eyes 
posteriorly. At the truncated apex are two short subulate tentacles on each side. 
Tentacular cirri two on each side, a dorsal and a ventral, the former more than twice the 
length of the latter, thick at the base, but tapering to a fine point. Body slightly 
diminished towards the head, and more distinctly so posteriorly. A brownish band passes 
from the head backward within the line of the lamellse on each side, and a median stripe 
of similar hue runs along the ridge of the dorsum. Dorsal cirri sub-orbicular, having a 
longer transverse diameter in the anterior feet, and a longer obliquely vertical diameter 
in the posterior feet. The setigerous lobe is somewhat small and bifid. The bristles 
show a large curved spine and a smaller one opposite it, and the ridges of the dilated 
ends of the shafts are spinous. The terminal blade is wide beneath, tapers to a nne 
point, and the edge is serrated. The ventral cirrus is ovato-lanceolate. 

1 ' Zeitsoh. f. w. Zool./ Bd. xxv, 1875, p. 42, Taf. ii, figs. 12—14. 
3 ' Vidensk.-Selsk. Forliandl./ 1871, (sep. copy) p. 4. 



1865. 
1867. 


Mysta barbi 


1871. 


>} >> 


1874. 


•>■> >) 


1879. 


•>•> }j 


1879. 


}) }} 


1883. 


'"> )> 


1901. 


•>•> }> 



MYSTIDES. 109 



Synonyms. 

Mysta barbata, Malmgren. Nord. Hafs- Annul., p. 101, Tab. xv, f. 34. 
idem. Annul. Polych., p. 26. 

Ehlers. Sitz. Phys. Med. Soc. Erlangen, III, p. 79. 
Malm. Op. cit. Groteb., p. 81. 
Tauber. Annul. Danic, p. 89. 
Theel. K. Vet. Akad. Handl., Bd. xvi, 3, p. 34. 
Wiren. Chgetop. f Vega' Bxped., p. 400. 
Mcintosh. Ann. Nat. Hist., ser. 7, vol. viii, p. 223. 

Habitat. — Procured in the trawl a little east of the island of May, January, 1891. 

Spitzbergen (Ehlers) ; Nova Zembla (Theel) ; Norway (Canon Norman) ; Siberia and 
Behrings Strait (Wiren). 

Head (Plate LVIII, fig. 10) in the preparation bluntly conical, with two conspicuous 
eyes posteriorly. At the truncated apex are two short subulate tentacles on each side. 
The tentacular cirri are two in number on each side, viz. a dorsal and a ventral, the 
former more than twice the length of the latter, somewhat thick at the base, but 
tapering to a fine point. 

Body slightly diminished towards the head and more distinctly tapered posteriorly, 
somewhat rounded on the dorsum, flattened on the ventral surface, and probably 
terminated by two cirri posteriorly. A brownish band passes from the head backward 
on each side within the line of the lamellaa, and a median stripe of similar hue occupies 
the ridge of the body. 

The foot (Plate LXIX, fig. 10) seems to correspond with Malmgren's description and 
figure, though it must be mentioned that there is considerable difficulty in following the 
distinctions of the various species given by the author. The somewhat imbricate dorsal 
cirrus may be described as sub-orbicular, having a longer transverse diameter in the 
anterior feet and a longer obliquely vertical diameter in the posterior feet, with a 
tendency to a point externally, where it goes much beyond the setigerous process. The 
latter is somewhat small, bifid, and bears the bristles (Plate LXXVII, fig. 19), which 
present a shoulder, from which a denticulated ridge passes to a large curved spine, with 
a smaller spine opposite it, the blade of the terminal process being between them. It 
(terminal process) is wide beneath and tapers to a fine point, and the edge is serrated. 

The ventral cirrus is proportionately broad anteriorly, narrower and more pointed 
posteriorly, and it projects considerably beyond the setigerous lobe. Malmgren describes 
its form as ovate, with a rounded apex. In this form it was rather ovato-lanceolate. 

Reproduction. — The specimen is a female laden with minute ova — apparently free in 
the coelomic space, so that the breeding season would appear to be in spring (May). 



Genus XXXVIII.— Mystides, Theel, 1879 (char, emend.). 

Tentacles, four long, or short, lateral rather than terminal ; tentacular cirri, three 
on each side, viz., on the first, second, and third segments ; eyes, two, large. The British 



110 MYSTIDES LIZZLE. 

species has comparatively short tentacles, but they may have been injured. Body linear, 
elongate, segments numerous (forty-eight, Theel). Dorsal cirrus renif orm ; seti- 
gerous lobe strong, bifid ; ventral cirrus ovate or ovato-rectangular. Distal end of shaft 
of bristle has a hook-like spine with a smaller opposite it. 

Theel 1 founded this genus on the great length of the three pairs of tentacular cirri. 
Eteone and Mysta have but two tentacular cirri ; Phyllodoce and Arteritis have four. His 
example had no anal cirri, the body terminating in an obtuse cone. 

Levinsen (1883) gave Mystides two to three pairs of tentacles, small dorsal lamellge 
(cirri), and three pairs of tentacular cirri. 



1. Mystides Lizzie, n.s. Plate L VIII, fig. 11 — head; Plate LXIX, figs. 11 and 12 — feet; 

Plate LXXVII, fig. 20— bristle. 

Specific Characters. — Head bluntly conical, with two large eyes widely separated and 
situated towards the posterior border. Anterior tentacles slender and minute, situated 
laterally. Three tentacular cirri — gradually increasing in length — at each side. They 
are longer than the homologous organs in Eteone and appear to approach in length those 
of Mysta, which, however, has only two. Body about three inches long; segments distinctly 
marked by a deep furrow in front and by a lozenge-shaped depression after the anterior 
third. Dorsal cirrus reniform, with a hilus at the base. The posterior differ from the 
anterior chiefly in their tendency to form a point at the free dorsal edge, and in their 
smaller size. The setigerous lobe is strong, bifid, and bevelled inferiorly. Distally the 
dilated ends of the bristles have a curved, hook-like spine directed towards the serrated 
edge of the terminal blade and with a spinous margin below it. A smaller spine occurs 
on the opposite side. The terminal blade is broad at the base and gradually tapers to a 
long and fine point. Its edge is minutely serrated. The ventral cirrus is ovate or ovato- 
rectangular, barely reaching the tip of the setigerous process at the tenth foot, but more 
nearly reaching it posteriorly, where the tip is less obtuse. Three madder-brown or 
purplish bands on the dorsum, viz. a band at each side, and a median, which, however, 
posteriorly is nearly severed into two by a pale median line. The pale portions between 
the coloured belts also increase in distinctness posteriorly, so that the animal has a boldly- 
marked series of bars. The dorsal cirri, with the exception of a few anteriorly, which 
have a dark speck or two, are pale. 

Habitat. — Tossed on the West Sands, St. Andrews, after a storm in February, 1870, 
when it was picked up by my sister. 

Off Cape Sagres, < Porcupine,' 1870. 

Head (Plate LVIII, fig. 11) bluntly conical, with two large eyes, widely separated 
and situated towards the posterior border. The anterior tentacles are slender and minute 
(whether from injury or otherwise is unknown), and they appear to be placed laterally 
rather than at the tip of the snout. Three tentacular cirri occur on each side (on the 
first, second, and third segments), and they increase in length from the first to the third. 
They are longer than the homologous organs in Eteone, and appear to approach in length 

1 'Ann. Polych. Nouv.-Zemb./ p. 35, 1879. 



MYSTIDES LIZZLE. Ill 

those of Mysta, which, however, has only two, and they are also of considerable thickness, 
though tapered distally. They are much longer than those of Langerhans' Mystides 
bidentata, which was also found by De St. Joseph at Dinard. 

Body about three inches in length, flattened, and with the segments distinctly marked 
by a deep furrow anteriorly, and by a lozenge-shaped depression after the anterior third. 
It is very slightly tapered anteriorly, but the diminution posteriorly is more pronounced. 
The ventral surface has a median line anteriorly, and the segments are marked by 
alternate elevated and depressed transverse bars, the former having an elevated and 
projecting cushion or boss at each side. The specimens were in spirit before they were 
observed, but three madder-brown or purplish bands could be distinguished. They 
were incomplete posteriorly, and Theel's example also was apparently imperfect. 

The foot (Plate LXIX, figs. 11 [tenth] and 12 [fiftieth]) has superiorly a large reniform 
cirrus of considerable thickness, having an umbricate arrangement, and being somewhat 
cordate at the base. Its hypoderm is coarsely areolar and scarcely streaked. The 
papilla supporting it is distinct. The anterior (e.g. tenth) differ from the posterior feet 
chiefly in the tendency to form a point on the free dorsal edge, and in their smaller size. 
The setigerous region is a strong bifid process, bevelled inferiorly, and bearing a series of 
stout, translucent bristles, with a slight curvature distally. The dilatation at the tip 
of the shaft is peculiar, presenting distally on each side a curved, hook-like spine directed 
towards the serrated edge of the terminal blade, and with a denticulated margin beneath. 
The other side of the distal end has a smaller spinous process (Plate LXXVII, fig. 20), 
also with a slightly denticulated edge below it. The shoulder is not clearly seen in every 
bristle, but it is distinct in those near the dorsal edge. The transparent terminal blade is 
broad at the base, and gradually tapers to a long and fine point. The edge is minutely 
serrated a considerable distance beyond the base. The minute structure of these bristles, 
if they agree with the foregoing, is not clearly defined by Theel, but his species was 
minute. 

The ventral cirrus is ovate or ovato-rectangular, barely reaching the tip of the 
setigerous lobe at the tenth foot, but more nearly reaching it posteriorly where the tip is 
less obtuse. 

The lamellas of the ' Porcupine ' example have a tendency to greater definition and 
angularity. 

Reproduction. — The specimen from St. Andrews is a female, with the eggs fairly 
developed, though not ripe, in February. De St. Joseph found a female of Mystides 
bidentata, Langerhans, with natatory bristles from the sixty-sixth segment backward, 
whilst the eggs were greenish. 

This form appears to fall under the genus Mystides of Theel, 1 a small example of 
which came from Nova Zembla. The peculiar conformation of the first segment which 
dorsally overlaps the head, the short lateral tentacles, the long tentacular cirri, the longer 
dorsal cirri, and the structure of the bristles sufficiently distinguish it. Its comparatively 
large size, in contrast with its northern relative, and the distinctive coloration are also 
noteworthy. It is named after my sister, who procured it on the West Sands of 
St. Andrews. 

1 ( K. Sc. Akacl. Handl./ Bd. xvi, 3, p. 35. 



112 ALCIOPIML 



No member of the Alciopidse has yet been met with in British waters, but there is 
no reason why such may not be found — e. g. off the southern shores of England or off the 
west coast of Ireland, as at Valencia. The remarkable fauna — both littoral and pelagic 
— of the latter region would lend countenance to this expectation ; as also would that 
occasionally found in the Moray Frith. 



9 



1% 



/ 






C,^.^ 



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v ^5 






v-4£ 



ff 53 



^ 



V 



Fig. 45.— Peculiar pelagic type near Syllidse and Alciopidse. The caudal cirri are reduced. 

In this connection, a minute form (fig. 45), about an eighth of an inch in length, 
found in the tow-net in St. Andrews Bay, in July or August, 1900, may be mentioned, 
especially as it does not conform, so far as can be ascertained, to any known species. At 
first sight it may be mistaken for Ioida, but a close inspection shows its marked distinction 
from that form and from the Syllidas, as well as its separation from the Phyllodocidge and 
Hesionidse. Unfortunately it had been stained and mounted in balsam before it came 
under observation, so that the precise structure of the feet and the more delicate processes 
has yet to be worked out. 

The head somewhat resembles that of Ioida in carrying two large black eyes on the 
dorsum, and apparently another pair on the ventral surface. The head-region is small 
and deeply cleft in the middle line, the two lateral flaps (one on each side) on which the 
eyes are situated being slightly angular in front, and trending from the median fissure 
outward and backward, the central part being coarsely fibrillated, as if from ruptured 



ALCIOPIDJE. 113 

muscular fibres, the ends of which occur on each side of the first segment- junction. A 
thin membranous flap dotted with cells and nuclei occurs in the front of each eye, and a 
projecting narrower process of the same tissue apparently below it, whilst a subulate 
tentacle, also thickly studded with deeply-stained nucleated cells passes forward a little 
below the frontal edge. All the structures just described are delicate, and are 
probably diaphanous during life. Moreover, they differ wholly from the appendages of 
the male and female buds of Autolytus, and do not approach those of the Alciopidse. 
A funnel-shaped depression with a median notch ventrally appears to indicate the oral 
region, which thus is wide, and this segment, which is not clearly differentiated from 
the head, bears on each side a foot directed obliquely forward and outward, and supported 
by a strong translucent spine and a series of translucent bristles. The direction of these 
feet shows that they have special functions — probably both protective and tactile. The 
bristles are comparatively stout, and have a distinct curvature of the shaft below the 
terminal dilatation, which is marked by a few oblique central strise. The distal end 
is bevelled, and the terminal piece is of moderate length and simply hooked. These 
bristles are borne by an elongated setigerous region slightly tapered from base to apex. 
The dorsal cirrus somewhat resembles that of a Syllis, being long, slightly tapered and 
articulated, and the same deeply-stained minute nuclei occur in these as in the cephalic 
processes, but they are proportionally more numerous. They arise from the base of 
the foot, and seem to have a short ceratophore, though the state of the preparation 
rendered this indistinct. A short, simple, subulate ventral cirrus, again, is attached 
inferiorly a little exterior to the base of the foot. The first foot is somewhat spindle- 
shaped, and the rest present an ovate or elliptical dorsal process, apparently with a 
special spine, but as only dorso-ventral views are obtainable further investigation is 
necessary. Every foot, from the second to the last, is furnished with a very long, 
translucent tuft of swimming-bristles. 

The body has fifteeen segments — diminishing gradually from the head backward, 
the segment-junctions being boldly marked. The pygidium is slightly bipapillose, and 
has two long caudal cirri which are longer and larger than the cirri immediately in front. 
A pale moniliform streak passes from the mouth backward in the central line, and 
probably represents the alimentary canal. 

The foregoing has the general outline of the Syllidae, and the appendages of the head 
resemble those of the male of Autolytus (Polybostrichus), and also of Krohn's Syllis 
prolifera, 1 the jointed cirri and the structure of the bristles especially recalling the latter. 
The great size of the spines, and the presence of a considerable dorsal spine, however, 
show divergence, and the minute structure of the head and mouth likewise differs. The 
whole aspect, however, suggests relationship with a budding form, and in the present 
uncertainty it is perhaps well to leave it in a separate position. 

The precise position of this form must, for the present, remain in abeyance until 
more material is obtainable, but it conforms to no British type hitherto observed. 

1 ' Archiv. f. Naturges./ Bd. xviii, p. 66, Taf. iii, fig. 1 (1852). 



72 



114 HESIONKLE 



Family VIII. — Hesionid2e (Sars). (Hesionea, Grube, 1851.) 

Head with two pairs of eyes, and two or three tentacles, and generally two 
biarticnlate palpi. Nuchal organs primitive ; ciliated surfaces in direct contact with the 
exterior (Racovitza). Body of moderate length. Pygidium with 2 anal cirri. Proboscis 
protrusible, armed or unarmed. Anterior segments (1 — 4) distinct or more or less fused, 
each carrying two pairs of tentacular cirri. The other segments uni- or bi-ramous ; 
spines black ; superior division generally reduced, furnished with simple bristles ; inferior 
division with compound bristles. Dorsal and ventral cirri filiform, faintly articulated. 

Segmental organ. — Hesione (Fallacia) has a very large genital funnel, not known to open 
to the exterior, but connected at one point with the open segmental trumpet. Exit of 
genital products has not been observed. In others, as in Ojohiodromus, at the moment of 
reproduction the organ develops a large genital funnel for the transmission of the sexual 
products (Goodrich). Fage, again, describes a cilio- phagocytic organ in connection with 



nc 
Fig. 46.— Section showing muscles of Ophiodromus flexuosus, D. Ch. 

the segmental apparatus in Hesione pantheria, a form, however, in which it does not 
transmit the sexual elements. 

It is interesting that Dr. Thomas Williams T long ago observed that the segmental 
organs occurred in the hollow of the foot as a beautifully looped, extremely delicate, 
slender, membranous ciliated tube. Further, that in the female the glandular yellowish 
mass which spreads into the hollow base of the cirri and feet can be traced into organic 
connection with the ciliated tube, and with that limb of the loop in which the vessels are 
most developed. He considered that the ova passed into one of the limbs of the ciliated 
tube. His description thus nearly approached the general structure as recently shown. 

In transverse section of the body-wall (Fig. 46) the dorsal longitudinal muscles are 
thick externally and tapered to the dorsal median line. The ventral longitudinal muscles 
are slightly curved and end abruptly internally against the somewhat large nerve-area 
which has the oblique muscles inserted dorsally, whilst externally are the hypoderm and 
cuticle. This family, therefore, agrees with Nephthys in the relations of the nerve-area. 

1 ' Philos. Trans./ 1858, p. 127, pi. viii, fig. 21. 






HESIONIDJE. 115 

The few forms of the Hesionidee known were grouped by 0. Fabricius, Linnseus, and 
others, amongst the Nereids. They formed the ninth genus of Savigny's Nereids. 
Subsequent authors generally placed them near the Phyllodocidse and Syllidae, though 
Benham separated them from the former by the Aphroditidse, with which they have little 
in common. 

Grube first linked the HesionidaB to the Phyllodocidas — under the title Phyllo- 
docea Hesionea. 1 In his last publication 3 on the Hesionea he defines the segments as 
often 22, though they may reach 81. The head is rounded, rhomboidal, narrowed in 
front, heart-shaped, with two or three simple, short, two- or three- jointed tentacles ; two 
or three segmented sub-tentacles (CErsted's palpi). Two pairs of eyes. On each side 
three or four (rarely one or two) tentacular cirri, slightly tapered, and sometimes with 
short segments. Dorsal cirri thread-like, on a ceratophore ; ventral and anal cirri 
shorter and devoid of a ceratophore. Foot simple or bifid; ventral bristles compound. 
Proboscis long and cylindrical, or shorter and almost ovoid, smooth or sometimes with 
a crown of papillae around the opening. In some, a tooth, as in the Syllidse, or a pair 
of jaws, or papillae ; occasionally tooth and jaws combined. 

His main sub-divisions are : (1) Head with two very short tentacles, without sub- 
tentacles (or with two). Fallacia, Hesione. (2) Head with two simple tentacles, and 
two or three jointed sub-tentacles (palpi). Magalia, Kefersteinia, Gastalia, Periboea, 
Syllidia, Psamathe fusca, Johnst. (3) Head with three simple tentacles and two sub- 
tentacles. The ventral bristles with three teeth. Irma, Orseis, Oxydromus, Podarhe, 
Gyptis, Ophiodromus, Tyrrhena, Leocrates, etc. In the foregoing synopsis he makes a 
special genus (Kefersteinia) for Psamathe, Johnston, yet likewise includes under a separate 
sub-section Psamathe fusca, Johnston, apparently having overlooked their identity. He 
noted that in Stephania flexuosa, Claparede, the sexes were distinguished by their 
colouration ; and that in the family generally the spines were black. In the Philippine 
annelids he paid special attention to the armature of the proboscis as an aid in classifica- 
tion. None of the British forms, however, show this condition. 

Ehlers (1864), who gives similar characters to Grube, characterizes the Hesionea as 
having short flattened bodies with comparatively few segments, often brightly coloured 
and transversely striped. The head is distinct — with tentacles only, or with tentacles, 
palpi, and four eyes. The following segment has large tentacular cirri. Foot large, 
single, or with a minute dorsal division ; jointed or simple capillary bristles ; long dorsal 
and ventral cirri. Anal segment with lateral processes and anal cirri or papillae. The 
extruded proboscis is short and smooth, and is followed by the stomach, the aperture to 
which has or has not papillae. He grouped the genera as follows : (a) Head only with 
tentacles. (1) Four tentacles. Hesione, Savigny. (2) Head with five tentacles. Podarhe, 
Ehlers. (b) Head with both tentacles and palpi; palpi two-jointed, (a) Foot with 
two well-marked divisions ; three tentacles. Ophiodromus, Sars. (b) Foot with a single 
division, or with two, the upper tubercular; two tentacles. Gastalia (Sav.), Sars. 
Ehlers (3) Palpi three-jointed, longer than the two tentacles. Periboea, Ehlers. 

i < Fam. Annel./ p. 58 (1851). 

2 Seniles, ' G-esellsch./ 1879. See also his work on the ' Philippine Annelids/ 1878. 



116 OPHIODROMUS. 

De Quatrefages (1865) again made a family, the Hesionea, for their reception, 
holding that they and the Syllids were analogous groups, allied to each other, yet pre- 
senting special modifications of the digestive system and other parts. He grouped them 
according to the condition of the feet, the length of the body, and the number of tentacles 
and tentacular cirri. His first genus Myriana contains a doubtful form, possibly from 
its length, Myrianida. He included several members of the family amongst his Syllids, 
instituting, for instance, a new genus for Gastalia fusea — of which he made two species, 
viz. Kefersteinia cirrata and K. Glaparedii. 

Claparede (1868) criticised De Quatrefages for lamenting the absence of correct 
structural details of the group, for he had overlooked Keferstein's description of Psamathe, 
and yet removed the genus Kefersteinia to the Syllidse without adequate reason. 

The Hesionidse formed one of the families of the Syllidiformia vera of Levinsen l 
along with the Nereidas, Syllidse, Nerillida3, and Sphserodoridse, and in Benham's 2 more 
recent classification, which has certain features in common with the foregoing, it is 
further linked to the Aphroditiclae (a relationship difficult to appreciate) under the sub- 
order JSTereidiformia. 

Gravier (1900) arranged this family like his predecessors, viz., according to the 
uniramous or biramous condition of the feet, the number of tentacular cirri, the condition 
of the proboscis, the tentacles, and the structure of the feet. 



Genus XXXIX. — Ophiodbomus, Sars, 1861. 
Oxydromus, Grube, 1853. 

Head trapezoidal with two pairs of eyes; tentacles filiform; palpi thick, Particu- 
late; six long, non-articulate tentacular cirri on each side. Body of moderate length. 
Proboscis short, sub-globose or shortly cylindrical; unarmed. Feet bifid, sub-equal, 
cylindrical, each lingule elongate, almost cirriform; dorsal cirrus long, non- articulate ; 
ventral cirrus shorter, arising from tip; dorsal bristles capillary, inferior compound- 
falcate. Body-wall as in Fig. 46. 

Goodrich (1901) observed that the genital funnel in Ophiodromus is smaller than in 
Hesione, but is more intimately connected with the segmental organ, so that the internal 
opening " can scarcely be said to exist as such." 3 

In the genera Leocrates (Tyrrhena), Ojohiodromus, and Irma, a gradually closer and 
closer union is established between the funnel and the segmental trumpet. In Irma, 
where the genital funnel is well developed, it is so completely fused with the segmental 
trumpet that the two organs can be distinguished only by their histological structure. 
The segmental organ thus acquires a large trumpet-shaped opening into the ccelom and 
acts both as a genital and an excretory duct (Goodrich). 

1 c Yidenskab. Meddel. Nat. Foren. Kjobenh./ 1883, p. 180. 
3 ' Camb. Nat. Hist./ ii, pp. 258 and 308. 
3 f Quart. J. Micr. Sc./ n. s., vol. xliii, p. 714. 



OPHIODROMUS FLEXUOSUS. 117 



1. Ophiodeomus flexuosus, Delle Chiaje, 1825. Plate LVII1, figs. 12 and 13 — head; 
Plate LXIX, fig. 13— feet; Plate LXXVIII, figs. 1 and 1 b— bristles. 

Specific characters. — Head small, distinct; eyes four, the anterior pair being the 
larger and wider apart. Tentacles three — two lateral and a median — besides a pair of 
palpi, which have a large basal joint, and are external and lateral in position. Body 
2 J inches long, dilating behind the head, attaining its maximum about the anterior third, 
and then tapering to the tail, which has two long cirri. The first four segments bear 
long tentacular cirri. Colour, various shades of lustrous brown ; a whitish streak, often 
with a bluish lustre, runs backward from the median tentacle, and divides the head into 
halves. The dorsum is transversely banded at intervals with belts of fine iridescent blue. 
Ventral surface a deep madder brown. Proboscis large, cylindrical, devoid of papillse or 
jaws. Foot elongated, biramous ; dorsal division has a long cirrus, a slender, tapering 
setigerous process, with a spine and a tuft of elongated, slender bristles, the shafts of 
which are longitudinally striated and have faint transverse touches, after the manner of 
the ventral series, but less distinct. Ventral division has a long setigerous region 
enclosing the spine, with, posteriorly, a stiff fan of jointed bristles, the terminal sabres of 
which have a claw at the tip blocked by a process beneath, and a serrated edge. 



Synonyms. 

1825. Nereis flexuosa, Delle Chiaje. Mem., II, pp. 368, 401, and 425, tav. xix, fig. 8. 

1841. „ flexuosa, Delle Chiaje. Descriz., Ill, p. 97, V, p. 103, tav. 129, fig. 8. 

1855. Oxydromus fasciatus, Grube. Arch. f. Naturges., p. 98, Taf. iv, f. 1 and 2. 

1861. Ophiodromus vittatus, M. Sars. Forh. Vid.-Selsk. Christ, and Nyt. Mag., XI B, p. 87. 

1864. Oxydromus fasciatus, Grube. Insel Lussin, p. 82. 

„ Podarlce albocincta, Ehlers. Borstenw., p. 190, taf. 8, f. 2 — 5 ?. 

1865. Nereis punctata, Johnston. Cat. Brit. Mus., p. 280. 

„ Ophiodromus vittatus, De Quatrefages. Annel., II, p. 657. 
1867. „ ., Malmgren. Annul. Polych., p. 31. 

1869. „ „ Mcintosh. Rep. Brit. Assoc. (1868), p. 339. 

,, „ ,, Mcintosh. Trans. R. S. Eclin., vol. xxv, p. 410, pi. xv, figs. 7, 8. 

1870. 8 tephania flexuosa, Claparede. Annel. Nap., Suppl., p. 118, pi. xii. fig. 1. 

1873. Ophiodromus vittatus, G. O. Sars. Bidrag. Christ. Fauna, p. 29. 

1874. „ „ Malm. Op. cit. Goteb., p. 81. 

„ „ flexuosus, Marenzeller. Beitr. z. Kennt. Adriat. Annel. in Sitz. Wien Akad., p. 23. 

1875. „ vittatus, Ehlers. Zeitsch. f. w. Zool., Bd. xxv, p. 44. 
1879. „ „ Tauber. Annul. Danic, p. 92. 

1883. „ „ Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 238. 

1885. ,, flexuosus, Cams. Fauna Medit., p. 238. 

1893. „ vittatus, Levinsen. Vidensk. Ud. " Hauchs," p. 328. 

1898. „ flexuosus, De St. Joseph. Ann. Sc. Nat., 8 e ser., t. v, p. 339. 

1905. ,, „ Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 324. 

1906. „ „ Bohn. Ann. Sc. Nat., 9 e ser., t. iii, p. 72. 



118 OPHIODROMUS FLEXUOSUS. 

Habitat. — Dredged somewhat frequently on a bottom of tenacious grey clay and mud 
in 4 — 8 fathoms in Lochmaddy, and especially in Ardmaddy Bay, and one or two were 
found at the verge of extreme low water in the same bay under an immersed stone and 
in sandy mud whilst digging for littoral annelids; on old oyster-beds and amongst 
tangle-roots in 5—7 fathoms in West Voe, Scalloway, Shetland; 100 fathoms in St. 
Magnus Bay, Shetland (Dr. Gwyn Jeffreys) ; 80—110 fathoms off the coast of Ireland, 
and in 15—20 fathoms in the Bay of Gralway, in the 'Porcupine' Expedition of 1869, 
on a bottom of muddy sand with pebbles; 9—125 fathoms 50 miles west of Valencia, 
Ireland (Dr. Gwyn Jeffreys); 81 fathoms off Cape Finisterre, 'Porcupine,' 1870; 
Station IV, bottom, Moray Frith, July, 1894 (Dr. H. C. Williamson). 

Extends to Norway (Sars), shores of France, Mediterranean. Burrows of Synapta on 
the shores of France (Perez). 1 

Head (Plate LVIII, fig. 12) small, distinct, with four eyes, the anterior being 
somewhat larger and wider apart, and sometimes elongated transversely. Tentacles 
three, a lateral on each side and a median somewhat shorter. The palpi have a thick 
basal region, the distal being slender and tapering so that the structure resembles a 
church spire. 

Body 2J inches long, dilates behind the head, attains its maximum about the anterior 
third, and then tapers to the tail. It is thus fusiform. The tail terminates in two long 
slender cirri which are shorter, however, than those of the fourth foot from behind. The 
first four segments bear modified limbs in the form of long cirri. The dorsum is of 
various shades of lustrous brown, whilst a whitish streak running backward from the 
central tentacle divides the head into halves. The dorsum is transversely banded at 
intervals with belts of fine iridescent blue, the bands extending to the tips of the feet. 
In a specimen 2J inches long there were a dozen conspicuous belts, besides less evident 
minor streaks posteriorly. The body-line at the base of each foot has a patch of the 
same colour, continued across the rugse of the segment in fine blue lines, but they vary in 
different specimens. The head in some has the same fine bluish colour. The ventral 
surface of the body is dark madder-brown. When placed in spirit OpModromus loses 
its tints in a moment, and generally breaks into several pieces. The same result happens 
on immersion in impure sea- water. 

The proboscis is large (Plate LVIII, fig. 13), devoid of jaws or papilla, and in 
extension it becomes cylindrical, or has a swollen base and a distal rim. The buccal 
orifice is capable of great dilatation. 

The body is garnished at each side with long resplendent tufts of bristles which 
glance with all the varied hues of the rainbow, while from each foot two long hair-like 
cirri project. The elongated foot is distinctly biramous (Plate LXIX, fig. 13), thus differ- 
ing from all the Hesionidso except Schmarda's Girrosyllis (Pseudosyllis, De Quatrefages) 
and (Ersted's Cast alia. The dorsal lobe (a) consists of the long cirrus (b), which springs 
from a basal segment (ceratophore), and tapers slightly towards the tip, and of the 
setigerous division, which tapers, as a slender process, outward to a point and envelops 
the spine. From the upper and posterior part of this division a tuft of slender, elongated 

1 'Proc. Verb. Soc. Sci./ Bordeaux, 1904—5. 



OPHIODROMUS FLEXUOSUS. 119 

and tapering simple bristles arises. When fully developed the shaft (Plate LXXIX, 
figs. 1, la, and 1 b) is longitudinally striated and has faint transverse touches, after the 
manner of the ventral series, though much less distinct. The ventral division of the foot 
presents a setigerous region, likewise carried outwards as a long tapering process (d) 
enclosing the spine, and from the posterior surface of the base of which issues the 
somewhat stiff fan of compound bristles (Plate LXXVIII, figs, la and 16). As usual 
in such forms the anterior segments possess bristles with somewhat shorter terminal 
processes, while the posterior (fig. 1 a) have their tips more elongated. Besides, in each 
foot the tips vary in length, the shorter being met with superiorly and inferiorly. The 
claw at the tip of the terminal division is somewhat indistinct, from the blocking of its 
outline by a process beneath. The serratures of the sabre are very fine and scarcely 
distinguishable towards the tip. The shaft of the bristle is obliquely striated towards 
the articulation, longitudinally throughout the rest of its extent, except at the pale base, 
and marked by a close series of transverse strise. 

In structure these bristles differ from allied British forms, e. g. Gastalia fusca and 
Gastalia punctata, each of which has compound bristles, and the peculiarly clawed serrated 
terminal process. Thus in the first-mentioned the larger size, the structure of the shaft, 
and the coarser serratures on the terminal process, distinguish it from Gastalia punctata, 
while the latter again has a much broader and proportionally larger terminal blade with 
coarser serratures and a better defined claw than in Ophiodromus, and though the shaft 
of the bristle more closely resembles it than that of Gastalia fusca yet it characteristically 
differs therefrom. The distinctions between Ophiodromus and the Pariboea and Podarke 
of Ehlers are not so clear ; indeed, the latter appears to be allied in the closest manner, if 
it is not identical. Unfortunately Ehlers does not figure the bristles on a large scale, so 
that their minute structure is uncertain. From the inferior border of the foot arises the 
inferior cirrus (e) which tapers gently towards the tip, and is shorter than the dorsal. 

Reproduction. — A large number of orange-coloured ova were emitted from a ruptured 
female in August, and those procured by Canon Norman off Bergen, Norway, in July 
showed the same feature. The ova are comparatively large. 

Claparede (1870) states that the ova of the Mediterranean examples were of a 
greenish-brown and Oil mm. in diameter. 

Habits.— Few annelids display so much activity and irritability. If the head is 
touched it darts vigorously backward, and if the tail is irritated it rushes as rapidly 
forward — with a wriggling sinuous motion. It is very sensitive to admixture of the sea- 
water with fresh. Thus, when conveying living specimens from North Uist in 1865 for 
a coloured drawing, the jar was replenished with sea- water at Dunvegan. Unfortunately 
the mists and rains obscured the surroundings of the creek — which had several streams of 
fresh water debouching into it, and it was mortifying to see the fine examples become 
pale and die in a few seconds after the dilute sea- water touched them. In other Hesio- 
nidas the changes in colour have been mentioned by several observers. Thus Professor 
Haswell l noticed that specimens of Halla parthenopeia changed from bright yellow to 
deep purple when placed in spirit. Mr. Crossland tells me that in Halla (Aglaura) fulgida, 
Sav., of the Red Sea : " The worm becomes dark when exposed to the air, whilst fresh 

1 ' Proc. Linn. Soc. N.S.W./ x, p. 752. (1886.) 



120 CAST ALIA. 

water dissolves out the same pink colour as is taken up by 70 per cent, alcohol. The 
addition of a little alcohol to the sea- water containing the living animal causes the mucus 
so abundantly secreted by the large dorsal cirri to become purplish. The alcoholic 
solution is deep reddish -brown by transmitted light, but shows a blue fluorescence when 
seen from above. These tints remained for several weeks, but gradually faded into a 
yellowish-brown colour." Yet in pure sea-water it is by no means difficult to keep 
Ophiodromus alive for a long time, and clayey mud is not deleterious. It swims 
through the water with more grace and even greater agility than Nephthys, but breaks 
into fragments if much interfered with. Even the separated posterior end keeps up a 
constant and violent motion — darting about in various directions for some time. While 
rupture of the body thus frequently occurs, yet reproduction of the posterior end readily 
takes place. 

Delle Chiaje's description 1 (1841) is: Head with four tentacles, furnished with a 
proboscis ; body yellowish-brown, banded transversely with white ; tail with two cirri ; 
tentacular cirri on six segments. His figures bear a close resemblance to the present 
species, though the identification is not free from doubt. On the whole it is probable 
that the northern form is only a variety of the southern, and it may be that it is a com- 
mensalistic species on Astropecten and Luiclia, though this has not actually been observed. 

The Hesione Steenstrupii of De Quatrefages (1865) from Guettary seems to approach 
this form very closely. 

The Oxydromus fuscescens of Marenzeller 2 (1875) from the Adriatic appears to be 
very closely allied. 

Levinsen (1883) states that Tauber considers this the Stephania fiexuosa of Delle 
Chiaje, but he differs from him. 



Genus XL. — Castalia, Savigny, 1820. 

Head somewhat square, with four eyes obliquely arranged. Tentacles two ; palpi 
biarticulate. Tentacular cirri two to three pairs, filiform and long. Body scolopendri- 
form; proboscis firm, barrel-shaped, aperture papillose, with or without two thickened 
ventral ridges (so-called jaws). Foot biramous ; superior division minute with or without 
simple dorsal bristles; inferior division large with compound (falcate) bristles, the 
terminal pieces being bifid. Dorsal cirri longer than the ventral. 

This genus was established by Savigny (1820) to include such forms as the Nereis 
rosea of 0. Fabricius. All he states is that this species, while agreeing generally with 
the heterogeneous assemblage of forms under his Nereides, differs in that the tentacular, 
dorsal, and anal cirri are long and smooth, and that there are two divisions in the foot. 

(Ersted 3 describes the palpi as like the tentacles and non-articulate ; four tentacular 
cirri ; four eyes ; minute, slender, pellucid, and edentulous maxillge ; pinnse two, superior 
minute, inferior large trilobed, one spine in superior, three in inferior division ; dorsal 
bristles capillary, ventral falcate. 

1 < Descriz./ v, p. 103, Taf. 109, fig. 8. 

2 ' Sitzb. d. K. Akad. d. Wiss./ i Jahrg. 1875, p. 15, Taf. ii, fig. 1 (sep. copy). 

3 ( Annul. Dan. Consp./ p. 23. (1842.) 



CASTALIA PUNCTATA. 



121 



The term Castalia was indicated by Savigny 1 in 1820 as suitable for the Nereis 
rosea of 0. Fabricius, and therefore has priority over Dr. Johnston's title Psamathe 
(1836), 2 and though the species described by the authors are different, there is no need to 
separate them generically. In the original description of Dr. Johnston the proboscis was 
described as edentulous, and the feet as uniramous, but OErsted 3 pointed out the presence 
of teeth, and showed that the feet were really biramous. 

Grube 4 (1879) observed that the genus was readily divided into two groups, in the 
first of which the dorsal division of the foot has from three to five bristles, whereas in 
the second the dorsal division has few and very small bristles — sometimes only one or 
two spines. 



1. Castalia punctata, 0. F. Miiller, 1788. Plate XL VI, fig. 2; Plate LXIX, fig. 14- 
foot ; Plate LXXVIII, figs. 2-2 b— bristles. 

Specific Characters. — Head somewhat square, pale yellowish, with two closely 
approximated red eyes on each side, the anterior pair obliquely elongated and wider apart 
as well as showing a cuticular lens anteriorly. Two slender tentacles on the anterior 
border ; external to each is a palpus with an enlarged basal segment, and a more slender 
distal region separated by a shoulder. Three pairs of long slender tentacular cirri, each 
arising from a prominent ceratophore on first three segments. Body about an inch long, 




Fig. 47. — Muscles of Castalia punctata, O. F. M., as seen in section of the body-wall. 

most massive anteriorly, gradually diminishing posteriorly, where it terminates in two 
long cirri. Colour of dorsum dull yellowish, crossed transversely by dark belts which 
become only touches posteriorly. The posterior two thirds is tinted olive from the 
intestine. Ventral surface dull yellow or salmon- colour, with a brown stripe in the 
median line, which increases in density posteriorly. The attenuated cirri are pale. 

The proboscis is firm, barrel-shaped, with a broad median notch ventrally, flanked 
by an elevated ridge with a papillose edge (ten papillse — Sars), and with four prominent 
papillas along the dorsal arch. Foot biramous, the dorsal spine piercing the base of the 
dorsal cirrus, which has a ceratophore, and is of proportionally great length, diminishing, 
however, from before backward. Dorsal bristles slender, pale, tapering, and simple. 

1 ( Syst. Annel./ p. 45-46. 

2 ' Loud. Mag. Nat. Hist./ ix, p. 14. 

3 'Annul. Dan. Consp./ p. 23. 

4 e Schles. Gesell./ loc. cit. 

70 
o 



122 CASTALIA PUNCTATA. 

Inferior division large, with a median, a dorsal, and a ventral papilla. The spine projects 
at the tip of the median. Bristles jointed, the shaft having finer markings than in 
Castalia fusca, its tip dilated, with oblique striae, and a distinct shoulder from which 
the terminal process projects. The latter is longest and most slender in the middle 
of the foot, and terminates in a curved claw with a secondary process beneath. 
The ventral cirrus subulate, extending in the preparations a little beyond the ventral 
papilla of the setigerous process. The body-wall (fig. 47) is powerfully muscular, the 
dorsal longitudinal muscles being separated in the middle line, whilst the ventral form 
two strong curved bands in transverse section. The oblique muscles are attached to the 
upper edge of the nerve-area. 

Synonyms. 
1776. Nereis punctata, 0. F. Miiller. Zool. Dan. Prodr., No. 2633. 
1780. „ rosea, 0. Fabricius. Fauna Gronl., No. 284, p. 301. 
1788. „ punctata, 0. F. Miiller. Zool. Dan., II, p. 28, Tab. lxii, f. 4 and 5. 
1791. „ „ Gbnelin. Linn. Syst. Nat., i, pt. 6, p. 3118. 

1799. „ rosea, 0. Fabricius. Skr. Naturh., s. 5, p. 175, Tab. iv, f. 14—16. 
1806. „ punctata, Turton. Linn. Syst. Nat., iv, p. 89. 
1825. „ rosea, De Blainville. Diet. Sc. Nat., xxxiv, p. 446. 
1828. Nereimyra rosea, idem. Ibid., lvii, p. 468. 
1843. Castalia punctata, (Ersted. Annul. Dan. Consp., p. 24, f. 15, 63 — 65, and 69 (excl. Fabr. 

and Sav. fide Malmgren) ; Reg. Mar., p. 78. 
1844-5. „ „ idem. Kroyer's Nat. Tids. (Drobak), p. 408. 

,, „ „ Sars. Nyt Mag., vi, p. 209. 

1848. Halimede venusta, H. Rathke. Beitr. Fauna Norw., p. 168, Taf. vii, f. 1 — 4. 
1851. Castalia punctata, Grrube. Fain. Annel., pp. 58 and 131. 

Sars. Nyt Mag., vii, p. 382. 

idem. Christ, vid. Selsk. Fork, 1861, p. 89 (sep. copy, p. 19). 
Danielssen. Reise, 1858, p. 117. 
De Quatrefages. Annel., II, pp. 102 and 664. 
idem. Ibid., p. 106. 
„ ,, rosea, idem. Ibid., p. 107. 

1867. „ punctata, Malmgren. Annul. Polych., p. 31. 

„ Psamathe punctata, Parfitt. Trans. Devon. Assoc, ii, pt. i, p. 233. 
„ „ pustulata, idem. Ibid., p. 234. 

1869. Castalia punctata, Mcintosh. Rep. Brit. Assoc. (1868), p. 339. 

idem. Trans. R. S. Edin., xxv, pp. 411 — 412, pi. xvi, f. 1. 

idem. Ann. Nat. Hist., ser. 4, vol. xiv, p. 197. 

Malm. Op. cit. Groteb., p. 82. 

Mcintosh. Invert, and Fishes St. Andrews, p. 121. 

Mobius. Jahresb. Comm. deutsch., p. 170. 

Tauber. Annul. Danic, p. 92 (partirn). 

Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 239. 

Hornel. Trans. Liverp. Biol. Soc, vol. v, p. 245. 

Levinsen. Vidensk. Ud. " Hauchs," p. 328. 



1853. 


)> 


1860 < 


3rl. „ 


>> 


>> 


1865. 


Psamathe 


jj 


Castalia 



1874. 

>) 
1875. 

}) 
1879. 
1883. 
1891. 
1893. 



Habitat. — Dredged in deep water near the Bell Rock, off St. Andrews Bay, and on 
Serpulce attached to Fusus antiquus on the same ground ; procured by the fishermen's 



CASTALIA PUNCTATA. 123 

lines (E. M.) ; between layers of shale near low water-mark at the East Rocks, St. 
Andrews (R. M.) ; interstices of laminarian roots in Bressay Sound, Shetland ; Cumbrae 
(Dr. Robertson). 

Extends to various parts of Norway, where it was first found by 0. F. Muller on 
oyster shells. 

Head somewhat square, pale yellowish throughout with two closely approximated 
dark red eyes on each side, the anterior pair obliquely elongated and wider apart, and 
showing a cuticular lens anteriorly. From the anterior border two slender tentacles 
arise, and external to each is a palpus having an enlarged basal segment with a more 
slender conical distal region separated by a marked shoulder. Three pairs of tentacular 
cirri follow, the dorsal cirrus of the third pair being the longest process in the body. 
These cirri are very long, slender, slightly tapered organs, and arise from a prominent 
ceratophore. 

Body about an inch in length, most massive anteriorly, and gradually diminishing 
posteriorly where it terminates in two long cirri. The feet are least prominent anteriorly, 
and gradually increase in length as the body diminishes, though posteriorly they again 
become small. The ground-colour of the dorsum is dull yellowish crossed transversely 
by dark belts (two in each anterior segment), which do not make a solid bar except in 
the centre of the dorsum, the sides forming an elongated space in which the pale ground- 
colour of the dorsum appears. Even the dense central portion of the bar is for the most 
part crossed by fine pale lines. The dark pigment-belts become less and less complete 
towards the posterior end, where they form only transverse touches. The posterior two- 
thirds of the dorsum (behind the proboscis) has an olive-hue from the alimentary canal. 
A dark speck occurs at the base of each foot, one at the base of the cirrus, and one 
between the feet. These touches become more distinct posteriorly as the central pigment 
wanes. The dorsal blood-vessel, which alters its course at the posterior end of the 
proboscis, also enlivens the region. The grouping of the dark transverse belts on the 
massive anterior region renders it one of the darkest parts of the body. The ventral 
surface has a dull yellow or salmon-hue with a brown stripe, which increases in density 
posteriorly, in the median line. The longitudinal pigment-striae are interrupted at the 
segment-junctions so that the band is elaborate. The cirri are all pale and of extreme 
tenuity. The proboscis in extrusion is a short barrel-shaped structure — with a broad 
median notch ven tr ally — flanked by an elevated ridge showing prominences or papillae, 
and having four prominent papillae along the dorsal arch. The whole organ is firm and 
resilient, and the aperture is spacious. 

The intestinal canal is ciliated. 

The foot (Plate LXIX, fig. 14) is distinctly biramous, the dorsal spine piercing the 
base of the dorsal cirrus. The latter springs from a peduncle (ceratophore) and is of 
proportionally great length, diminishing, however, from before backward. The dorsal 
bristles are slender, pale, tapering, and simple, ending in a smooth point (Plate LXXVIII, 
fig. 2). The inferior lobe is largely developed, and has a median, a dorsal, and a ventral 
papilla, so that the region appears to be trifid. The spine pierces the median papilla, and 
the bristles form a broad fan, the ventral, however, being shorter than those above them. 
The bristles are compound (Plate LXXVIII, fig. 2 a)., the shaft differing from that in 



124 CASTALIA PUNCTATA. 

Gastalia fusca in having finer transverse markings. The tip is slightly dilated and has 
obliqne stria3 with the transverse markings, and a very distinct shoulder, from which the 
terminal process goes. In anteroposterior view the tip is somewhat fusiform with two 
blunt points on each side of the origin of the terminal piece. The latter (Plate LXXVIII, 
fig. 2 b) is largest and most slender in the middle of the foot, broadest in the ventral 
series. The edge is finely serrated and the tip somewhat diminishes and terminates in a 
curved claw with a secondary process beneath. Though the serrations diminish toward 
the tip, yet there is no marked difference between the upper and the lower bristles as in 
Gastalia fusca, Johnst. Some of the longer tips show a curve. The median bristles of the 
example from St. Magnus Bay have unusually long terminal processes. 

The ventral cirrus arises nearer the body than the ventral papilla, and has a similar 
structure to the dorsal, only it is shorter. All the cirri are pale. 

In life the ceratophore of the dorsal cirrus, which has the dark pigment-patch, is 
richly ciliated, and the cilia are continued on the skin in the immediate neighbourhood 
towards the intermediate dark patch. The blood-vessel internally passed up one side of 
the ceratophore and down the other, but did not seem to enter the cirrus proper. 

Habits. — This species appears for the most part to frequent the deeper water, and 
the finest examples are got there. It has much of the lively habits of Ophiodromus, and is 
irritable and impetuous. When irritated, it gathers its cirri about it and swims rapidly 
through the water. Yet with all its beauty, it, like the gaudy Amblyosyllis, clings to the 
under surfaces of stones, and hides in crevices, so that, unless these are only its resting- 
places, it has little field for the display of its attractions. 

Reproduction. — The fine pinkish colour of the feet is due to the presence of large ova 
(September), which are grouped in masses at the base and extend for some distance into 
the foot. 

It is probably this species which Garstang 1 found breeding at Plymouth in August. 

Two very distinct species have been included by Dr. Gr. Johnston in Britain on the 
one hand, and by several Continental authors on the other, under the head of Psamathe 
punctata, viz. the present form and P. fusca, Johnst. Neither Malmgren nor De Quatref ages 
recognized the distinctions. The latter indeed groups them as two species under the 
genus Psamathe, but, like CErsted and Johnston, he has not appreciated the differences. 
The synonym Gastalia punctata, (Erst., is placed under each without allusion to the fact 
that CErsted shows two bristle-tufts (dorsal and ventral) in each foot, whereas De 
Quatrefages states as a generic character: " Pedes unirames." The bristles of the two 
species, besides, are quite different. 

Amongst the figures of "Marine Insects" given by Baster is one 2 which probably 
represents this species. 

This annelid appears to be both the Nereis punctata and N. pnstidata* of Montagu, 
the latter being a greenish variety dredged off Torcross, Devonshire (MS. Drawings, 
Linn. Soc, Plate LII, fig. 3). The N. fascicular ia of the same observer (MS. Drawings, 
Plate XXXVIII, fig. 2) is closely allied, if not identical. 

1 ' Journ. M. B. A./ vol. iii, p. 225. 

2 ' Opuscula Subseciva/ Tab. iv, fig. 4. (1762.) 

3 This specific name is also given for an Eulalia in MS. Tab. 51, fig. 4. 



CASTALIA ABCTICA. 125 

H. Rathke (1843) formed a new genus and species for this form which he correctly 
recognized as 0. F. Midler's Nereis punctata. 

Tauber (1870) merges the Psamathe punctata of Johnston in this species, but they 
are distinct alike in minute structure, colouration, and habitat. 



2. Castalia arctica, Malmgren, 1867. Plate LVIII, fig. 14 — head; Plate LXIX, fig. 
15— foot; Plate LXXVIII, figs. 3 and 3 a— bristles. 

Specific Characters. — Head somewhat square with two tentacles anteriorly and four 
rather small eyes (reddish in spirit), the anterior pair wider apart. Tentacular cirri as 
in Castalia fusca. The palpi are two-jointed. Body about half an inch long, some- 
what fusiform, and terminating in two long anal cirri. Proboscis with a papillose rim 
(ten papillse — Theel, Levinsen, etc.), the single example from the ' Porcupine ' not being 
definite in this point. Two prominent brownish hooked maxillae, ventro -lateral in 
position. Foot biramous, two slender spines passing into the ceratophore of the dorsal 
cirrus, which is more or less segmented. No bristles are present in the division. 
Setigerous region of the inferior division diminishes towards the tip, and has a conical 
papilla in front of the tip of the spine. Bristles curved towards the tip of the shaft, 
which is dilated and with a shoulder from which the terminal process proceeds. The 
latter is slightly curved and diminished distally, has a serrated edge, and ends in a minute 
claw with a secondary process beneath. Ventral cirrus comparatively short and 
subulate. 

Synonyms. 

1790. Nereis aphroditoides, Fabricius. Fauna Grcenl., No. 278, p. 296. 

1867. Castalia Fabricii, Malmgren. Annul. Polychget., p. 32 (?). 

„ „ arctica, idem. Ibid., p. 32. 

1883. „ aphroditoides, Wiren. Chast. 'Vega' Exped., p. 401. 

„ „ Fabricii, Levinsen. Vidensk. Meddel. Foren. Kjobenh., p. 239. 

1889. „ aphroditoides, Marenzeller. Arch. f. Naturges., 55^ p. 128. 

Habitat. — Dredged in 90 — 125 fathoms fifty miles west of Valencia on the west coast 
of Ireland by Dr. Gwyn Jeffreys. 

Sengiarak, Greenland (O. Fabricius). Siberia and Behrings Strait (Wiren). 

Head (Plate LVIII, fig. 14) somewhat square with the usual tentacles anteriorly 
and four rather small eyes (reddish in the preparation), the anterior pair wider apart. 
The tentacular cirri resemble those of Castalia fusca. The palpi are two-jointed. 

Body appears to be about half an inch in length, is somewhat fusiform in outline, 
and terminates posteriorly in two long cirri. The proboscis, unfortunately, was injured 
in drawing, so that its subsequent examination was unsatisfactory. It had a papillose 
rim, but whether so many papillae occurred as in the figure is an open question. Two 
brownish hooked maxillae occur ventro-laterally and project during the extrusion of 
the cylindrical or barrel-shaped organ. 

The foot (Plate LXIX, fig. 15) is biramous, two slender spines passing into the base 



126 CASTALIA ARCTICA. 

of the ceratophore of the dorsal cirrus. The ceratophore is proportionately shorter than 
in Psamathe fusca. The cirrus shows crenations towards the base and is more distinctly 
segmented towards the tip. The setigerous region diminishes distally, and has a conical 
papilla in front of the tip of the spine. The bristles form a vertical fan and have a 
distinct downward curve towards the tip of the shaft, which is somewhat dilated and 
furnished with a shoulder from which the terminal piece proceeds, the margin sloping 
from this upward to the pointed end of the shaft (Plate LXXVIII, fig. 3). The shaft 
is marked similarly to that in G. fusca, but the lines are much more delicate and less 
easily observed. The terminal blade is shortest ventrally and gradually increases in 
length superiorly, the blades of those on the dorsal edge, however, being shortened, 
though they are longer than those of the ventral edge. All are slightly curved and 
diminished distally, and end in a minute claw with a secondary process beneath. The 
serrations on the edge are finer than in G. fusca, yet there is a tendency to a more 
distinct series inferiorly, and then they gradually become finer and fade away distally. 
The bristle is thus intermediate in character between G. punctata and 0. fusca. 

The ventral cirrus is comparatively short and subulate. 

In all probability a flap existed at the tip of the setigerous process, but the state of 
the preparation was unfavourable for determination. It may also be smaller than in 
allied forms. 

Reproduction. — The specimen is a male with the reproductive elements well developed 
in June. 

There are certain discrepancies between this form and the Nereis aphroditoicles of 
0. Fabricius, especially in the structure of the foot, for, in addition to the single terminal 
papilla, Fabricius adds that there are two smaller papillae inferiorly. The general 
greenish hue with red lines make the species described by Fabricius conspicuous, and 
it may be that further investigation will show that the British form differs from it. 
In the uncertainty as to the minute characters of the older form it has been deemed 
best to include the present species under Gastalia arctica. 

In the structure of the foot this form differs both from Gastalia Fabricii, Malmgren, 
and G. midtipapillata, Theel, for, instead of the three papillse which characterize the tip of 
the setigerous region in these, the British species has only one conical papilla in front of 
the tip of the spine. 

Malmgren observes that in G. arctica the dorsal division has only a single spine, and 
in the British species the spine is so slender in contrast with those of G. Fabricii that 
Malmgren may have overlooked the fact that in some there are two. No details of the 
structure of the feet are given by Malmgren, and none of the bristles or of the proboscis, 
so that considerable doubt must remain as to the nature of these. 

Wiren amalgamates the G. Fabricii and G. arctica of Malmgren as the G. aphroditoides 
of Fabricius, the former, G. Fabricii, having two acicules dorsally and three or four 
ventrally, whereas G. arctica has only one dorsally and two ventrally. He considers the 
latter a variety of the former, but he gives no details of the structure of the foot and no 
figures. 

Marenzeller (1898) alludes to this form from Spitsbergen as having three spines, as 
in 0. Fabricii, Malmgren. 



CASTALIA FUSCA. 



127 



3. Castalia fusoa, Johnston, 1836. Plate XL VI, fig. 3 ; Plate LVIII, figs. 15 and 16— 
head; Plate LXIX, fig. 16— foot; Plate LXXVIII, fig. 4— bristle. 

Specific Characters. — Head small, square, with two black eyes close together on each 
side, the anterior elongated, placed obliquely, and with a cuticular lens. Tentacles, two, 
slender ; palpi with a massive basal segment from which the short distal process passes off 
at an angle. Tentacular cirri, two pairs, filiform, slightly segmented and of great length, 
the dorsal being the longer. Body 1-3 inches long, scolopendriform, slightly narrowed 
in front, and tapered to an attenuated tail with two long, slender anal cirri. Colour 
pinkish or fawn, yellowish-brown, and purplish-pink, darker in the centre, with the dorsal 
blood-vessel which makes a double curve behind the oesophageal region. Cirri pale. 
Proboscis short, firm, cylindrical, the aperture surrounded by a dense fringe of filiform 
papillse. Two thickened ventral edges (the "jaws" of Dr. Johnston). The body-wall 
(Fig. 48) is as muscular as in G. punctata, though, perhaps, the ventral longitudinal 




Fig. 48. — Muscles of Castalia fusca, Johnst., as seen in section; n. c. nerve-cords cut in the 
line of the ganglia. Other letters as before. 

muscles project less inferiorly. The strong oblique bands are attached to the firm summit 
of the large nerve-area, which has externally hypoderm and cuticle. 

Feet prominent, longer than the diameter of the body ; biramous, for two small 
spines pass to the base of the dorsal cirrus, which is very long and slightly tapered, with 
a large and long ceratophore. Articulations are distinct towards the tip. The setigerous 
(ventral) division is massive, with a blunt papilla for the spine, another a little above it, 
and a third process is formed by the short ventral cirrus. Bristles leave the foot between 
the posterior flap and the papillge in front, are yellowish, and boldly curved towards the 
tip of the shaft, which is striated longitudinally and transversely. The end of the shaft 
is not much dilated and is bevelled to a shoulder for the terminal blade which ends in a 
curved claw with a secondary process beneath. The edge of the blade has minute 
serrations at the base, then a series of larger curved teeth, followed by minute serrations 
which diminish and disappear before the tip is reached. 

Synonyms. 
1836. Psamathe fusca, Johnston. Loud. Mag. Nat. Hist., ix, p. 14, woodcut f. 1. 
1840. „ „ idem. Ann. Nat. Hist., iv, p. 230, pi. vii, f. 4. 



128 CASTALIA FUSCA. 

1851. Psamathe fusca, Williams. Rep. Brit. Assoc, 1851, p. 213. 

,, „ „ Grube. Fam. Annel., pp. 58 and 130. 

1853. Nereis punctata (villosa), Daly ell. Pow. Creat., ii, p. 158. 

1862. Psamathe cirrhata, Keferstein. Zeitsch. f. w. Zool., Bd. XII, i, p. 107, Taf. ix, 

f. 32—36. 

1863. „ cirrata, Claparede. Beobacht., p. 55, pi. xiv, f. 1 — 7. 

1864. „ fusca, Grube. Insel Lussin, p. 82. 

„ ,, „ Kolliker. Kurz. Ber. Schott., p. 9. 

1865. Kefersteinia cirrata, De Quatrefages. Annel., ii, p. 41. 
„ ,, Claparedii, Idem. Ibid., p. 42. 

„ Psamathe punctata (partim), Johnston. Cat. Brit. Mus., pp. 182 and 342. 
,, „ fusca, De Quatrefages. Annel., II, p. 101. 

1869. „ „ Mcintosh. Trans. R. S. Edin., xxv, pp. 412—413, pi. xvi, f. 2. 

1874. „ „ idem. Ann. Nat. Hist., ser. 4, vol. xiv, p. 197. 

1875. „ „ idem. Invert, and Fishes St. Andrews, p. 121. 
1879. Castalia cirrata, Langerhans. Zeitsch. f. w. Zool., xxxiii, p. 306, f. 41. 

„ Psamathe „ Marion. Ann. Sc. Nat., 6 e ser., viii, art. 7, p. 21. 
1885. Hesione fusca, Cams. Fauna Medit., p. 239. 

1890. Psamathe cirrata, G-iard. Bull. Sc. Fr. Belg., t. xxii, p. 78. 

1891. „ fusca, Hornel. Trans. Liverp. Biol. Soc, vol. v, p. 245. 
1899. Kefersteinia cirrata, Mcintosh. Nat. Sc. (May), p. 376. 

1904. „ „ Allen. Journ. M.B.A., n.s., vol. vii, p. 220. 

1906. „ „ Bohn. Ann. Sc. Nat., 9 e ser., t. iii, p. 71. 

„ „ „ De St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 228. 

Habitat. — Procured under stones between tide-marks at various points around the 
British shores, and in Shetland in the roots of tangles in the laminarian region. It also 
occurs in fissures of rocks — as many as thirty or forty having thus been obtained on 
raising a large slab at Lochmaddy — where it is also found in the laminarian region. 
Plymouth (Allen). 

St. Vaast, Normandy (Keferstein) ; Naples (Claparede). Mediterranean (Cams). 
Marseilles (Marion). 

Head (Plate LVIII, fig. 15) small, square, entire in front, with two pairs of well- 
marked black eyes, the pairs on each side being close together, the anterior elongated and 
placed obliquely, and each furnished with a cuticular lens. Tentacles two ; palpi with 
a massive basal segment, from which the terminal division comes off at an angle. It is 
directed outward, and is comparatively short. The inner tentacles are slender, cylindrical, 
tapered a little at the tip, and arise from the upper and inner base of the foregoing pair. 
Tentacular cirri two pairs, filiform, slightly jointed and of great length, the dorsal being 
the longer. 

Body from one inch to three inches long, scolopendriform, slightly narrowed in front 
and tapered to a slender tail, which is terminated by two long slender cirri. 

Colour generally pinkish or fawn, yellowish-brown, or purplish-pink, darker in the 
centre, and enlivened by the red dorsal blood-vessel, which makes a double curve behind 
the oesophageal region. Cirri pale. Dr. Johnston considered that the purplish hue of some 
specimens spreads rapidly over the body when the animal is alarmed. As this occurred 



CASTALIA FUSCA. 129 

in April it is possible that it may have been connected with the development of the ova. 
Sir J. Daly ell, again, noted that the finest examples were scarlet, though some were 
yellow, but that the colour depended on the food. 

The proboscis (Plate LVIII, fig. 16) is short, firm, cylindrical, the circular aperture 
in extrusion being surrounded by a dense fringe of filiform papillae. It has two thickened 
ventral edges, which are simply the thickened parts of the organ in protrusion. The whole 
organ is firm, and well adapted for predatory habits. Dr. Gr. Johnston found that they 
devoured each other in confinement, and Sir J. Dalyell fed them on mussels. The latter 
author thought that the colour of the body was dependent on the food, but more probably 
it is due to the development of the reproductive elements. Intestine richly ciliated. The 
feet are prominent throughout, stretching, with the bristle-tufts, outwards more than 
the diameter of the body. Though apparently uniramous they are really biramous 
(Plate LXIX, fig. 16), for two small spines pass to the base of the dorsal cirrus, which is 
the only representative of the dorsal division. The ceratophore of the cirrus is both 
large and long, and the cirrus itself is very long, and is slightly tapered from the base 
towards the tip. Articulations are visible some distance above the base, and are very 
marked towards the tip. The setigerous lobe is massive, with a bluntly conical 
extremity, the tip being formed by a blunt papilla into which the spine goes, and another 
occurs a little above it ; while the ventral cirrus, which is comparatively short, unjointed, 
and little tapered, leaves the foot internal to the bristles, which project between a 
posterior flap and the papillae in front. The shafts of the bristles (Plate LXXVIII, 
fig. 4) are boldly curved towards the tip, are yellowish, striated longitudinally, except at 
the tip, where the striae are oblique, and marked by fine transverse striae. The end of 
the shaft is not much dilated and is bevelled to a shoulder, which is slightly bifid and 
gives attachment to the terminal blade. The upper end of the bevelled region is also 
bifid. The distal process is longest in the middle of the series, shorter superiorly and 
inferiorly, and ends in a curved claw with a secondary process beneath. The edge of the 
blade has minute serrations at the base, then a series of larger curved teeth, followed 
again by small serrations which diminish and disappear before the tip is reached. Filmy 
algae grow from the tip of the shaft and the terminal blade. 

Reproduction. — In May, when carrying ova, the body assumes a fine mauve or 
purplish-pink hue, which extends from the posterior region of the proboscis to the tail. 
In confinement the ova are discharged by rupture of the body, which then assumes the 
ordinary hue. The ova are pinkish and show a zona with markings externally. They 
were probably unripe. The scarlet specimens alluded to by Dalyell may have carried 
ova. Females with advanced ova are found at Plymouth in April (Allen). 

Habits. — It swims gracefully and actively through the water with an undulating 
motion, its cirri streaming from the sides, but if irritated it discards these organs, and 
even (as in spirit) breaks into fragments and throws off its feet. According to Bohn, 1 
the movements of this species are in accord with those of Eulalia, but its activity and 
irritability are quite distinctive. 

This species was first found by Dr. Johnston, who afterwards confounded it with the 
Gastalia punctata of CErsted, nor did Claparede and others notice the difference between 
1 Much of the literature of the subject has escaped this author. 

74 



130 LEOCRATES ATLANTICA. 

the latter and Dr. Johnston's species. The name (Psamathe) given by Dr. Johnston had 
already, in 1814, been nsed for a Crustacean by Rafinesque, and in any case lapses in 
favour of the earlier title of Savigny. There is no reason why the Psamathe cirrata of 
Keferstein should be separated. Fine examples under the latter title (Kefersteinia cirrata) 
were sent from Plymouth by Dr. Allen. 

Dr. Williams (1851) mentions that it feeds on algse, but this is doubtful. Dalyell 
found that it fed voraciously on mussels, " darting out its proboscis and absorbing a great 
quantity. The colour changes with the quality of its food." 



Genus XLI. — Leoceates, 1 Kinberg, 1865. 

Head with three tentacles, a pair of palpi, and a frontal tubercle ; eyes four. Body 
comparatively short, with few segments, ending in two anal cirri. Proboscis with two 
maxillas. Foot biramous; dorsal bristles capillary; inferior bristles compound (falcate). 

This genus was established by Kinbergin 1865 for a species from China — near Hong 
Kong. The author gives the proboscis only a single "maxilla" in the median dorsal 
aspect, but probably the second was overlooked. 

Claparede again constituted the genus Tyrrhena, as indicated, for the same type 
which Achilla Costa had discriminated under the name of Castalia Glajparedii in the 
manuscript of the fourth ' Annuaire du Musee Zoologique de Naples. 5 

Gi-rube 2 showed the identity of the genera established by the two preceding authors, 
and gave an amended description of the genus in his Philippine Annelids, with an account 
of a new species from Zamboanga. 

1. Leoceates atlantica, Roule, 1896. Plate LVIII, fig. 17— head; Plate LXIX, fig. 
17— foot; Plate LXXVIII, figs. 5 and 5 a— bristles. 

Specific Characters. — Head (in spirit) more or less quadrate, longer diameter antero- 
posterior, and generally with a median depression and a cordate posterior border. Eyes 
on an elevated ridge in middle of head ; anterior pair wide apart, large, rounded, and with 
a cuticular lens ; posterior pair also rounded or oblique. Palpi thick at the base with 
a short terminal segment. Lateral tentacles subulate and slender; internal to palpi. 
Median tentacle short and subulate. Tentacular cirri eight, with massive basal segment 
(ceratophore) and spine. Body about one and a half inches long, massive anteriorly, 
though tapering a little towards the head in life and more distinctly posteriorly. Rounded 
or convex dorsally, iridescent and minutely ringed, flattened ventrally, and with a deep 
median groove. First four segments fused, but with special nerve-twigs. Proboscis with 
a thick, frilled, anterior edge, a blunt papilla projecting from the central fold dorsally. 

1 ' Ofvers. af K. Vet.-Akad. Fork/ 1865, No. 4, p. 244. Claparede, in 1868, constituted the 
genus Tyrrhena for the same form, naming it after the Tyrrhene Sea opposite Tuscany. ' Annel. 
Neap./ p. 227. 

2 ' Jahresb. Schles. Gesellsch./ November 19th, 1879, s.a., p. 25; 'Anneliden fauna der 
Philippinen/ p. 105 (1878). 



LEOCPATES ATLANTICA. 131 

Chitinous teeth in mid-dorsal and mid-ventral lines. Feet sixteen in number, long and 
prominent. The dorsal lobe has — just below the ridge of the foot — the basal segment of 
the dorsal cirrus, the setigerous region in front being conical with a long black spine. 
Dorsal bristles simple, stiff, tapering to hair-like tips. Shaft marked by transverse strise 
almost to the tip, which has a regular series of minute spikes. Ventral division irregularly 
conical with terminal papilla for spines; upper slope downward and outward, inferior 
(more abrupt) downward and inward. The iridescent bristles form a dense tuft ; 
shafts with close series of longitudinal and transverse strias, slightly dilated distally and 
then bevelled — from the attachment of the terminal process to the point. Terminal blades 
longest in upper third of tuft, shorter dorsally, shortest ventrally. Each is flattened, 
slightly tapered and curved distally, where it is bifid — with a secondary process. The 
bevelled edge of the blade is attached by a web to the shaft. Ventral cirrus long, slender, 
subulate, and tapering — extending a little beyond the fleshy part of the foot. 

Synonyms. 

1896. Tyrrhena atlantica, Koule. Camp. 'Caudan/ Ann. Univ. Lyon, Aout-Septembre, 1895, p. 455, 

pis. xxi, xxiv, xxv, f . 9, 10, 24, 28 and 29. 
1901. „ „ Mcintosh. Ann. Nat. Hist., ser. 7, vol. viii, p. 227, pi. i, f. 5, 6 and 8. 

Habitat. — Several specimens were dredged in the 'Porcupine' Expedition of 1870 
at Station 9, on the Channel Slope, lat. 48° 6' N., long. 9° 18' W., in 539 fathoms, on a 
bottom of grey mud. Bottom-temperature 48 o, Fahr., surface-temperature 64°*0 Fahr. 
Off Ushant in 400 fathoms (J. E. Allen). 1 

In the ' Oaudan ' from a depth of 1410 metres in the Gulf of Gascony (Poule). 

Head (Plate LVIII, fig. 17) more or less quadrate, with the longer diameter antero- 
posterior, and generally with a median depression and a somewhat cordate posterior 
border, so that the eyes occur on an elevated ridge on each side. The anterior pair of 
eyes are wide apart, large, rounded, and with a cuticular lens in the centre, the darkest 
part of the pigment-ring, in spirit, being a crescent on the inner and posterior border. 
The posterior pair are rounded or oblique in the preparations, the latter, in all probability, 
being due to the presence of the pigment on the inner border. Both pairs lie in the 
middle of the head. From the outer and inferior angle of the head on each side the palpi 
project forward and downward, the segment at the tip being proportionally short, and 
Claparede observes that, in his species, it can be partly invaginated. The organ is small 
compared with that in the Syllids, is thickest at the base, and tapers towards the tip. The 
tentacle, arising from the head internal and superior to the former, is subulate, much more 
slender, and longer. In the median line between the posterior pair of eyes is a compara- 
tively short, subulate, median tentacle, which is easily overlooked, especially in an 
indifferent preparation. The edge of the proboscis projects in the preparations, and in 
the median line dorsally is a prominent papilla, which lies under the anterior border of 
the snout— fitting as it were between the palpi, and Claparede states that, in his form, it 
is surrounded in life by a circle of cilia. In these preparations the eyes are considerably 
1 I am indebted to Dr. Allen for this and other information on this species. 



132 LEOCRATES ATLANTICA. 

larger than in examples of L. Olaparedii from Naples, and the tentacular cirri are larger 
and longer. 

Body about one and a half inches long, massive anteriorly, though in life it probably 
tapers a little towards the head, as Delle Chiaje and Claparede show in L. Olaparedii, 
and more distinctly towards the tail. It is rounded or convex dorsally, iridescent and 
minutely ringed, somewhat flattened ventrally, and with a deep median groove. The tail 
terminates in a free median vent, with a long cirrus at each side. The tentacular 
cirri are eight in number on each side, and are long, tapering organs, springing from a 
massive basal segment (ceratophore), which is furnished internally with spines. The 
first four segments are fused, but, as Claparede showed in his species, they receive special 
nerve-twigs, and thus he thinks the view of De Quatrefages, that all the tentacular cirri 
arise from the buccal segment, untenable. The proboscis in situ presents a somewhat 
thick, frilled, anterior edge, the median papilla formerly alluded to projecting from the 
central fold of the dorsal arch. As Claparede pointed out in L. Olaparedii, chitinous 
jaws are situated in the present species at the anterior end of the organ in situ in the 
mid-dorsal and mid-ventral lines. The dorsal tooth at the free edge has the shape of a 
bifid fan, whilst the ventral resembles a conical process. The massive wall of the organ is 
chitinous and of a brownish hue, and extends backward quite a third the length of 
the body, terminating at a prominent aperture (the gut) in the centre of its posterior wall. 
Two flattened lateral cseca, also with dark pigment, occur in this region. The inner surface 
of the proboscis has two thick pads behind the teeth in front, a transverse bar on each 
side in the middle, and the posterior half is subdivided by deep grooves. It seems to be 
an efficient prehensile and suctorial organ, and its great proportional size probably is in 
relation to its functional importance. 

The bristled feet (Plate LXIX, fig. 17) are sixteen in number, long and prominent, 
and their resplendent tufts of bristles make them still more conspicuous. The dorsal 
lobe presents, just behind the ridge of the foot, the basal segment of the dorsal cirrus, 
while the setigerous region in front is acutely conical, the long black spine passing to the 
apex. Each bears a tuft of simple and somewhat stiff bristles, which taper gently to a 
hair-like tip. The whole bristle is marked by transverse strise, as if cameratecl, till 
within a short distance of the tip. Moreover, the tip is furnished with a regular series of 
minute spikes. The latter are much more conspicuously developed in L. Olaparedii from 
Naples, the spikes in some being continued far downward and forming a feature along 
each side of the bristle. It was the comparison between the dorsal bristles of the two 
forms that many years ago (long before the expedition of the ' Caudan ' was thought of) 
led to their separation. Claparede had overlooked the presence of these spikes in his 
species, as also has Eoule in the present form. The ventral division of the foot is 
irregularly conical, with a prominent papilla for the powerful black spine or two. The 
upper slope of the region is downward and outward, the inferior (more abrupt) is 
downward and inward. In shape the foot thus differs from that of L. Olaparedii, 
which is longer, and the inferior setigerous region has an abrupt truncated extremity, 
with, superiorly, a papilla for the spine. The ceratophore of the dorsal cirrus is 
carried erect in the specimens from the 'Porcupine, 5 whereas it is horizontal in 
L. Olaparedii, but this is not a point of much moment. The bristles (Plate 



LEOCEATES ATLANTICA. 133 

LXXVIII, figs. 5 and 5a) form a dense tuft. The shaft has a close series of 
longitudinal and transverse striaa, is slightly dilated at the distal end, and then bevelled, 
from the attachment of the terminal process to the point. The distal blades are longest 
towards the upper third of the tuft, shorter dorsally, and shortest ventrally. Each is 
flattened, slightly tapered, and curved distally, where it is bifid, with a secondary process 
beneath. Oblique strias slope from the serrated edge of the blade downward and 
backward. The bevelled base of the blade is attached by a web to the shaft. These 
bristles, though pale, are finely iridescent, and in some are brownish in the posterior 
region of the body. 

Comparing these bristles with those of L. Glaparedii (Plate LXXVIII, fig. 6) the 
whole bristle is more slender, the tips are longer and more delicate, and the bifid tip is 
more minute. The boldly bifid nature of the tip in L. Glaparedii and the more distinct 
secondary process are in contrast with L. atlantica, the second or inferior spur being 
curved and the secondary process adpressed. The serrations on the edge of the blade 
are similar. The backward tilt of the bristle is seen in both, but L. atlantica has also a 
tendency to a dorsal curve (i. e. a convexity toward the dorsal edge throughout). It is the 
same type of bristle in both, but that of the abyssal species has been modified. It is the 
same as that of Dalhousia atlantica of the ' Challenger,' l which agrees with this species 
except in the absence of maxillae. Should these be found in fresh examples, then the 
specific name given in the Annelids of the ' Challenger ' would stand. 

The ventral cirrus is of considerable length, slender, subulate, and tapering. It 
extends a little beyond the fleshy part of the foot. 

Claparede found the male elements in L. Glaparedii developing in winter. None 
have been seen in the present species. Eisig 2 notes the occurrence of hermaphrodite 
glands in Hesione sictda, and cites the remarks of Claparede on Leocrates (Tyrrhena) to 
show that such a condition may also occur in that species. The subject should be 
re-investigated in living forms. In the same paper Eisig gives his views as to the caeca 
of the gut of Leocrates (Tyrrhena,) having the function of a swim-bladder, but it cannot be 
said that either structurally or physiologically these organs lend themselves to this 
theory. 

What relationship the foregoing form or that of Claparede has to the various species 
described or alluded to by De Quatrefages 3 is uncertain, for there is little that is definite 
or that can be relied on in his treatment of the Hesionidae, as, indeed, Claparede long ago 
pointed out. In the number of the tentacular cirri Leocrates (Tyrrhena) agrees, with the 
genus Fallacia of De Quatrefages, but this is all that can be said with safety. 

Though the species had long been discriminated in my collection, the publication of 
the Annelids of the ' Cauclan ' by Prof. Louis Roule gives his title priority. He correctly 
describes the distinctions from L. Glaparedii in regard to the tentacular cirri and teeth. 
His account of the dorsal bristles, however, requires further elucidation, for he states that 
they are of two kinds — viz. a rare form, which is simple and delicate, tapering to a curved 
point ; the other a camerated bristle, which at the tip has a cap of granular matter giving 

1 ' Annelids of the Challenger/ p. 186, pi. xva, figs. 5 — 7. 
3 l Mitth. Zool. Stat. Neap./ II, pp. 298—300. 
3 f Anneles/ pp. 95 — 111. 



134 DALHOUSIELLA CARPEXTERI. 

it the aspect of a racquet. In the examples from the ' Porcupine ' not a few of the dorsal 
bristles had a granular parasitic mass near the tip, as, indeed, is liable to happen in spiked 
bristles. That the type of bristle should be so altered (as shown in Professor Poule's right- 
hand figure in fig. 29, Plate XXV) is peculiar. He has likewise omitted to record the 
presence of the spikes. His artist has, further, taken considerable liberties in his drawing 
of the ventral bristles (Plate XXV, Q.g. 28), but perhaps the style of plate adopted is 
unsuitable for the illustration of structures so delicate and yet so characteristic. The distal 
segment of the palpus has also been overlooked in Plate XXI, fig. 10, and the sub-frontal 
papilla is tapered to a point, whereas that in the specimens from the 'Porcupine' is blunt. 

Genus XLII. — Dalhousiella, 1 Mcintosh, 1901. 
Head devoid of a median tentacle, with a deep median furrow separating the closely 
approximated eyes. Tentacles about the length of the palpi. Tentacular cirri— eight 
pairs — from the peristomial segment backward, and furnished with spines. Tips slightly 
moniliform. Body typical of the Hesionidge. Proboscis with firm muscular wall, no 
teeth. Foot — dorsal division represented only by two small black spines at the base of 
the long dorsal cirrus— ventral, with compound (falcate) bristles. Ventral cirrus long 
and subulate. 

1. Dalhousiella Oarpenteri, Mcintosh, 1901. Plate LVIII, fig. 18— head ; Plate 

LXXVIII, fig. 7— bristle. 

Specific Characters. —Read typical ; devoid of a median tentacle. Tentacles appear 
to be about the length of the palpi, which have a short terminal segment. A deep median 
furrow separates the rounded lobes on which the large and closely approximated eyes 
(four) are placed. Eight pairs of tentacular cirri, with spines, and slightly moniliform 
tips. The papilla beneath the snout is small (or little elevated). Body typical, about one 
and a quarter inches long (in spirit), and with seventeen bristled segments. Two short 
anal cirri occur beneath the vent, but the extremity seems to be injured. Proboscis with 
a firm wall and glistening internal surfaces. Xo teeth. Shape of the feet somewhat 
uncertain, but they are long, and appear to diminish more towards the outer edge than in 
Leocrates. Dorsal cirrus arises behind the ridge of the foot, has two small black 
spines, and is long, tapering, and slightly moniliform. Xo dorsal bristles visible. 
Setigerous region tapers towards the outer extremity, where a papilla occurs. Spine 
black and powerful. Bristles have comparatively short shafts which are minutely striated 
longitudinally and transversely. When seen on edge the tips of the shafts are somewhat 
fusiform from the dilatation below the bevelled region. In lateral view the dilatation is 
less pronounced, though the diameter is greater, and the striaB are oblique. The dorsal 
limb of the fork is the longer, but is frequently broken. Edge of the distal blade is 
minutely spinous, even up to the base of the fork. Ventral cirrus slender and subulate, 
stretching beyond the tip of the setigerous process. 

1 The genus Dalhousia was named in honour of the Earl of Dalhousie, late Secretary for 
Scotland, vide c Annelids of the Challenger/ p. 186. 



DALHOUSIELLA CARPENTEBI. 135 

Synonym. 

1901. Dalhousiella Carpenteri, Mcintosh. Ann. Nat. Hist., ser. 7, vol. viii, p. 231, pi. i, f. 9 

and 10. 

Habitat. — Dredged in the ' Porcupine ' Expedition of 1870, at Station 9, on the 
Channel Slope, lat. 48° 06' N., long. 9° 18' W., in 539 fathoms, on a bottom of grey mud. 
It accompanied Leocrates atlantica, which, like Ophiodromns and other Hesionidae, has a 
fondness for mud — often of a most tenacious description. 

Head (Plate LVIII, fig. 18) agrees with the type observed in Dalhousia of the 
6 Challenger ' Expedition, and also with that in Leocrates, though the median tentacle is 
absent. The tentacles appear to be about the length of the palpi, which have a short 
terminal segment. A deep median furrow separates the rounded lobes on which the large 
and closely approximated eyes are situated. The preparation shows less disproportion 
than usual between the anterior and posterior eyes, but it is not good. Eight pairs of 
tentacular cirri, with spines, occur on the buccal and following segments, and they appear 
to have long and slightly moniliform tips, as in allied forms. 

Body of the typical form, about one and a quarter inches long (in spirit) and with 
seventeen bristled segments. The posterior end appears to be injured, though two short 
cirri occur beneath the vent. The papilla under the snout is small or little elevated. 

The proboscis has a firm wall and a glistening interior surface, but, though the parts 
in the mid-dorsal line are dense, no distinct teeth are present. 

The shape of the feet is uncertain in the preparation, but they are long, and appear 
to diminish more towards the outer edge than in Leocrates. The dorsal cirrus arises 
somewhat behind the ridge of the foot, has two very small black spines at the base, and 
is long, tapering, and slightly moniliform. No dorsal bristles existed in the preparation. 

The setigerous region tapers towards the outer extremity, where a papilla occurs. 
The spine is black and powerful. The bristles (Plate LXXVIII, fig. 7) have comparatively 
short shafts which are minutely striated longitudinally and transversely (internally) as 
in allied forms. When seen on edge the tips of the shafts are somewhat fusiform, from 
the dilatation below the bevelled region. In lateral view, though the diameter is greater, 
the dilatation is less pronounced and the striae are now oblique. The terminal blade is 
comparatively short, has a web connecting its bevelled base with the shaft, is slightly 
curved, and tapers very little to the deeply forked (longitudinally split) terminal region. 
The dorsal limb of the fork, viz. that opposite the serrated edge, is the longer, but is 
frequently broken. It is proportionally longer in the upper bristles of the tuft. The 
edge of the blade is minutely spinous, even up to the base of the fork. The ventral 
cirrus is slender and subulate, stretching beyond the tip of the setigerous process. The 
structure of this bristle would seem to be diagnostic, for though that of Stepha.nia flexuosa 
of Delle Chiaje, 1 as figured by Claparede, 2 is a step in its direction, yet not even uniform 
and continuous friction could make it resemble that of the present form, and this difference 
would suffice without referring to other distinctive characters of the species. 

1 'Mem. degli Anim. s. Vert./ II, pp. 308, 401 and 424, tav. xix, fig. 8; ' Descriz. e Not./ Ill, 
p. 97 : V, p. 103, tav. cxxix, fig. 8. 

2 ' Suppl. Annel. Neap./ 1870, p. 118, pi. xii, fig. 1. 



136 MAGALIA PERARMATA. 



Genus XLIII. — Magalia, Marion and Bobretzky, 1875. 

Cephalic lobe bearing four eyes, two palpi (jointed), and two tentacles. Buccal region 
with three pairs of tentacular cirri on each side. Proboscis armed with a stylet and two 
maxillse. Feet uniramous. 

1. Magalia perarmata, Marion and Bobretzky, 1875. Plate LIX, fig. 1 — head, and 2 — tail; 
Plate LXV, fig. 10— proboscis ; Plate LXIX, fig. 18— foot; Plate LXXVIII, figs. 8 
and 8a — bristles. 

Specific Characters. — Head somewhat quadrangular, eyes rather large, close together 
on each side about the middle ; anterior pair with lenses. Tentacles comparatively long, 
smooth, fixed over the palpi, which are bi-articulate. Buccal region under the head ; six 
pairs of articulated tentacular cirri, the dorsal of the second pair longest. Each has a 
basal spine. Body 7 — 8 mm. in length, slightly tapered anteriorly and posteriorly; 
segments 31 — 32. Anal cirri long and articulated. Colour, yellowish with a brown 
touch at each dorsal cirrus, and similar touches on the head and buccal region. Dorsal 
surface minutely striated transversely. Proboscis with a crown of papillae with barbed 
palpocils, and the adjoining surface (in extrusion) has long hair-like papillas. A 
symmetrical pair of jaws, and a median tooth. Intestine sacculated. Foot uniramous, 
dorsal cirrus long, and with a spine in ceratophore ; conical setigerous region with two 
spines; bristles slender, translucent, with long terminal pieces, especially superiorly, 
the tip minutely hooked and with a secondary process beneath. Ventral cirrus subulate. 

Synonyms. 

1875. Magalia jperarmata, Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., t. ii, pi. vi, f. 16 a — 11 

and pi. vii, f. 16. 
1904. „ „ Allen. Jonrn. M. B. A., n.s., vol. vii, p. 220. 

Habitat. — Not uncommon in dredgings from Queen's Ground, Asia Shore, and 
Millbay Channel, Plymouth (Allen). 

Under stones and in prairies of Posidonia, and in the coralline region, Marseilles 
(Marion and Bobretzky). 

Head (Plate LIX, fig. 1) somewhat quadrangular, bearing four eyes of con- 
siderable size situated near each other towards the middle of the region. The anterior 
pair have lenses. The tentacles are comparatively long, smooth, and are attached over 
the palpi which are bi-articulate. The buccal region lies entirely under the head, and 
the six pairs of articulated tentacular cirri are usually directed forward, the dorsal of the 
second pair being the longest, though it does not exceed the first dorsal cirrus. Bach 
has a spine at its base. 

Body 7 — 8 mm. in length (Marion and Bobretzky), slightly tapered anteriorly and 
more so posteriorly. Setigerous segments 31 or 32. The anal segment carries two 
long articulated cirri (Plate LIX, fig. 2). 

Colour yellowish, with a brown touch at the base of each dorsal cirrus, and similar 



SYLLIDJE. 137 

touches on the head and buccal region. The dorsal surface of the segments shows under 
the microscope transverse striae, somewhat irregularly arranged. 

The proboscis in extrusion (Plate LXV, fig. 10, from Marion and Bobretzky) has a 
crown of prominent papillae, soft and contractile, and with barbed palpocils, and the 
adjoining surface has long flexible hair-like papillae. Moreover a symmetrical pair of 
jaws occurs a short distance within the former, and a slightly tapered tooth or stylet in 
the middle attached to a dark chitinous socket. The intestine is somewhat sacculated ; 
the glandular region (urinary, Marion and Bobretzky) commences about the twenty-fifth 
segment. 

The typical foot (Plate LXIX, fig. 18) is uniramous, having the long cirrus dorsally 
with a spine in the ceratophore, and a somewhat conical setigerous region with a terminal 
papilla, supported by two spines, and bearing a fan-like tuft of slender, translucent 
bristles. These have slightly curved shafts which are a little dilated, striated, and 
bevelled at the tip (Plate LXXVIII, figs. 8 and 8a), and with long, slender, terminal 
pieces ; indeed the superior are so long and slender as to resemble those of Nereids, and 
the minute structure of the tip is indistinguishable. In the shorter forms the tip is 
hooked and a secondary process appears to be present beneath, whilst the ventral 
cirrus is subulate, and the edge of the blade is minutely serrated. 

Marion and Bobretzky state that the cirri of the segments adjoining the tip of the 
tail have no spines in the ceratophores. 

Reproduction, — Marion and Bobretzky found ripe males and females at Marseilles in 
December and January. 

Habits. — It can swim rapidly after the manner of a Syllis, though the body is much 
shorter, and the long yellowish cirri make it very conspicuous (Marion and Bobretzky). 



Family IX. — Syllim: (Alitor.). 

The Syllidae are characterized by a thread-like body, definitely segmented, and often 
with articulated tentacles and cirri, the former and some of the latter being occasionally 
very long. The head is rounded or quadrangular, has three tentacles— a median and two 
lateral — and usually four eyes. Palpi, in various degrees of development and separation, 
are generally present. Buccal segment with two tentacular cirri, rarely with bristles. 
The alimentary apparatus consists of a mouth opening into a pharyngeal cavity, through 
which the protrusible proboscis (pharynx, Schlundrohre), which has a chitinous wall, can 
be exserted. This organ in situ often shows a sinuous outline, has anteriorly a series of 
soft papillae at its margin, and a little behind a hard prominent tooth, or two maxillae 
(Grube). It terminates posteriorly in the proventriculus, a region often more or less 
barrel-shaped and marked by rows of dots from glands. The proventriculus is followed 
by a short portion— frequently tapering posteriorly— which ends in a dilated region, 
often with two lateral caeca, 1 and which is marked from the intestine behind by a 

1 Said by Benham (after Meyer ?) to be used for storing water. 

75 



138 



SYLLKLE 



constriction. Body terminated posteriorly by two cirri, sometimes with an intermediate 
papilla. The minute feet have dorsal and ventral cirri, or the latter are absent, are 
uniramous, and bear falcate bristles with a terminal process, which is either simple or 
bifid. Swimming dorsal bristles also occur in the sexual forms. 

Grube describes the stolones as having few segments, two or three tentacles, two 
palpi, with tentacular appendages and tentacular cirri ; two eyes, generally large. Feet 
with two fascicles of bristles ; bristles in some compound, in others simple and long. 
Cirri present or absent. Proboscis and proventriculus absent. 

The chief features of the body-wall (Fig. 49) in such species as Syllis armillaris 
is the proportionally large size of the dorsal longitudinal muscles, which form in trans- 
verse section a massive arch resting, as it were, upon the broad base formed by the 
incurvation of the lower edge. A slight median raphe carries the mesentery from the 
gut. Externally are the circular fibres, the very thin hypoderm, and lastly the thick 
cuticle. The arrangement of the fibres of these and the ventral muscles in transverse 
section corresponds with that of other families, such as the Nephthydidge. The ventral 
muscles approach each other very closely in the median line, a condition, however, which 




^X 



nc 



vm 



Fig. 49. — Muscles of the body-wall of Syllis armillaris (Miiller) (Erst., as seen in section. 

does not appear to be general in the group, so that the nerve-cords are pushed upward, 
only a pedicle, apparently of basement-tissue, connecting them with the external 
hypoderm. The oblique muscles pass below the cord to be attached to the raphe in the 
intervals. 1 Another feature of interest is the presence of a strong muscular slip — from 
the fan-like muscles of the spines — which goes straight downward at intervals through 
the ventral longitudinal muscles to the basement-membrane. As slips from the bases of 
the spines pass dorsally along the inferior border of the dorsal longitudinal muscles, and 
on the other hand, ventrally along the outer border of the ventral muscles, the movements 
of the spines and of the foot with its bristles are well provided for. 

Though De Quatrefages did not observe blood-vessels in the Syllidas, Ehlers, 
Claparede, De St. Joseph, Viguier, and Albert found them, the latter describing dorsal 
and ventral vessels, a branch to the dissepiment, and a caecum in the middle of the 
genital gland. Malaquin showed that the dorsal and the ventral vessels communicated in 
the cephalic region by a double loop. Such is, briefly, the condition in the Syllides and 

!• Claparede (1868) mentions that the Syllids have a large tubular fibre on the dorsal surface of 
the nerve-chain. This has not yet been observed in the British forms. 



SYLLIDiE. 139 

Eusyllides, the dorsal trunk anteriorly in the ganglionic region splitting into two, which 
pass anteriorly downward to form the ventral. In each segment, posteriorly, a vascular 
trunk unites the dorsal with the ventral ; and, in the genital segments, a vascular csecum 
leaves the ventral vessel a little behind the dissepiment and goes to the genital gland. 

Malaquin describes lymphatic glands along the ventral vessel in certain forms, and 
amoebicyte-glands in Syllis hamata. The blood is colourless, and the movement is from 
behind forward in the dorsal vessel ; from before backward in the ventral. 

Some forms are epigamous, others schizogamous or viviparous, and the remarkable 
condition of several has long given examples of the alternation of generations of various 
authors. The sexual distinctions between male and female buds, and between these and 
the nurse-stocks are characteristic. Alternation of generations in Annelids was first 
suggested by De Quatrefages (1843) for the condition in Syllis monilaris, Savigny, and it 
was further exemplified by Krohn (1852) in Syllis prolifera. Alex. Agassiz, again, in 
his account of Autolytus comutus, accurately distinguished the several generations, though 
he considered the parent-stock to be constantly devoid of sexual elements. It has since 
been shown 1 that sexual products (chiefly female) appear in segments 11 — 13 of the 
stock from which the stolon separates, and the same occurs in Procersea ornata and 
Myrianida. Calvin Mensch, therefore, considers it more a case of sexual dimorphism, 
and not a sexual alternating with an asexual generation. Epitokous conditions occur in 
Autolytus longeferiens and in other Syllids. Moreover, some carry eggs in a brood- 
pouch, whilst others bear them in sacs or in pairs along the body. In no group of 
animals, indeed, are the complexities of reproduction more pronounced. 

Segmental organs. — E. Horst 2 describes a Syllis-bud with extrusible segmental 
organs. It has large eyes, like an Alciopid, articulated dorsal cirri, and a brown spot at the 
base of the foot dorsally, and the terminal pieces of the bristles are bifid. At the ventral 
base of each foot, except the first, a large dark-brown sac is extruded. JSTo sexual products 
occurred in the body- cavity. He alludes to the observation of Ehlers 3 that in several 
species of Glycera, bladder-like bodies (branchiae) capable of extrusion were found. 

Goodrich 4 observes that in young Syllidse the genital funnel can scarcelv be said to 
exist, being represented only by a few crowded cells in the coelomic epithelium on the 
anterior face of the septum just above the opening of the small segmental funnel. In 
budding forms, such as Autolytus, Myrianida, and Haplosyllis, the genital funnel advances 
little on the foregoing in the regions from which genital products are not extruded. In 
those shedding the sexual products a large trumpet-shaped genital funnel is formed, as in 
the Phyllodocidge, on the anterior end of the segmental organ, the dilated canal of which 
transmits the products. Fage 5 gives similar observations, the minute nephrostome being 
replaced at maturity by a large genital funnel ; and points out that in the males of certain 
forms a peculiar transformation of the segmental organ occurs at maturity, a change 
which he associates with the formation of spermatophores, a kind of sexual dimorphism 

1 E.g. Mensch, 'Americ. Nat./ yol. xxxiy, p. 165 (1900). 

2 ' Notes Leyden Mus./ yol. xi, p. ] 1, pi. i and pi. ii, fig. 1 (1889). 

3 f Die Borstenw./ pp. 659 and 676. 

4 Op. cit., p. 728. 

5 Op. cit, p. 312, fig. 25, and pi. vi, figs. 9—12. 



140 SYLLIDS. 

thus being present, as in certain Spionidse. As no solenocytes are present the nephridial 
functions are performed by the special cells of the organ. This author agrees with 
Pruvot in thinking that the segmental organs have a mechanical function (by resistance) 
during locomotion, but this appears to require further elucidation. 

Though a conspicuous feature in the family no case of branching or lateral budding 
occurs amongst the British Syllidge. Langerhans describes an example of Syllis variegata 
with two heads, whilst Syllis ramosa of the ' Challenger ' has a furor for budding at all 
points. Neither has the Syllis vivipara of Krohn 1 from Nice, as recently and carefully 
described and figured by Goodrich 2 at Naples, been hitherto captured in our waters. 
The Neapolitan form has the head of a Syllis, with moderately long tentacles and cirri, 
which are all articulated, the proboscis has apparently about ten papillae and a single 
tooth towards the front in extrusion, and simple tips to the terminal pieces of the 
compound bristles. Mesnil 3 is inclined to consider all such cases (which are not to be 
confounded with parasitic forms in the body-cavity of Syllids) as associated with 
parthenogenesis. 

Novel instances of collateral budding have been lately described by H. Parlin 
Johnson 4 in Trypano syllis ingens and in T. gemmipara, the former from Pacific Grove, 
California, between tide-marks, and the latter from Puget Sound. In the last-mentioned 
the buds, about fifty in number, arise in front of the last twenty-four segments of the 
diminished caudal region of the annelid. They form a tuft like a series of fronds of a 
minute fern, have no swimming-bristles, only a median strand for an alimentary canal, 
two large nerve-cords, two large eyes, dorsal and ventral cirri, spines, and bristles. Both 
species approach generally the Syllis gigantea of the ' Challenger,' 5 though no buds were 
present in the latter. Mr. Cyril Crossland, who has done such able work on the Annelids, 
tells me that he procured a Syllid with a rosette of buds at the posterior end, at Wasin, 
British East Africa, living in a red sponge common in ten fathoms in that locality. The 
annelids are bright orange, with four brown eyes. The dorsal cirri are annulated, very 
long, and of a still deeper hue. The tentacles and ventral cirri are shorter, but also 
annulated. The rosette of young is of the same colour as the parent, but each is shorter 
and broader, and no cirri are visible under a lens. Their tails keep waving in the water, 
the attachment being only by the head. Mr. Crossland's forms were collected before the 
publication of the American author was issued, but his species is still undescribed. 

Forms commensalistic in sponges like the foregoing and Syllis ramosa of the 
c Challenger ' lead the way to what appears to be a parasitic Autolytus, found by 
Mr. Crossland, also at Wasin, Bast Africa. " This annelid was attached by the extruded 
proboscis to a specimen of a yellow Nemertean, another to a Polynoid, seven examples to 
Nicidon gracilis, n.s., and a single specimen to another Niciclon of the same species. The 
Syllids cannot be pulled off their prey without rupture except in the case of the soft 
Nemertean. In the polychset hosts the point of attachment is always near the parapodia. 

1 < Arch. f. Naturges./ Bd. xxxv, p. 197 (1869). 

2 ' Journ. Linn. Soc./ vol. xxviii, p. 105, pi. xiii (1900). 

3 ' Compt. Rend. Soc. Biol./ Paris, vol. liii, p. 270 (1901). 

4 c Biol. Bullet./ vol. ii, No. 6 (1900), and ' American Naturalist/ vol. xxxvi, No. 424 (April, 1902). 

5 < Annel. < Challenger/ p. 193. 



SYLLIDJE. 141 

The proboscis seems incapable of retraction, and the parasites do not move after removal 
from the host. The Syllid is colourless, except the posterior part of the gut, which is 
grey, has four very minute eyes, very long tentacles, and uniramous feet." 

Syllids have, moreover, long been celebrated for their phosphorescence. Thus 
Viviani 1 (1805) mentions and figures no less than three, viz. Nereis cirrhigera (Odontosyllis 
fulgurans ?), Nereis mucronata, and Nereis radiata. The British forms are as conspicuous 
in this respect as the foreign, one of the commonest (Eusyllis tubifex, Grosse) causing 
striking manifestations of the phenomenon from its vast abundance on the blades of 
tangles covered by Obelia. 

In this country the pelagic types chiefly belong to the genus Autolytus, but in foreign 
waters certain of the Syllids proper likewise appear to have a pelagic stage. Thus when 
Mr. Crossland was in the Pearl Fisheries' steamer in Mersa Harbour, Durur, he observed 
numerous "small Heterosyllids circling round on the surface after the manner of 
Heteronereids. They were rapidly discharging genital products, the water in the jar in 
which they were placed becoming milky in a minute or two. They were readily killed by 
adding a drop of spirit to the water, and under this treatment showed a brilliant green 
phosphorescence. The dorsum was marked with alternate oblique and transverse bands 
composed of minute dots. The dorsal cirri were slender and the ventral cirrus absent. 
Long swimming-bristles occurred superiorly, and the terminal pieces of the compound 
bristles were simple." 

Andrews 2 also observed the adults of Autolytus prolifer swimming at the surface of 
the sea in the evening. 

The relationships of the remarkable pelagic types found by Greef 3 off the Canaries, 
such as Pontodora and Pelagobia, cannot be accurately determined, though he was 
inclined to link them to the Syllidao. In general form and in the structure of the 
alimentary canal, nervous system, and bristles, they certainly appear to approach that 
family and the Hesionidse. 

The Syllids abound between tide-marks, under stones, amongst zoophytes, algas, 
shells, and sponges. In the laminarian region they frequent oyster-banks, mussel-beds, 
or occur in great numbers amongst zoophytes on the blades of the tangles. 

They are especially common in the Mediterranean and at Madeira (Langerhans), yet 
the colder British area is not far behind the more genial southern waters in the variety of 
the species which frequent it. Thus Langerhans thought the waters of Madeira 
surpassingly rich with forty species, yet our own waters hold nearly the same number, 
and, whilst a few may prove to be varieties of one form, others not included may be 
discovered by future investigators. 

They range to a considerable depth (600 to 1380 fathoms). 

Little was done by the early naturalists to assign a definite place to the Syllidse. 
Savigny made them the twelfth genus of his Nereids, and his description is fairly 
accurate for the period. 

Grube, in his ' Familien der Anneliden' (1851), gave order to the group, but he 

1 f Phosphorescentia Maris/ G-enuas, 1805, p. 11, tab. iii, figs. 1 — 6. 

2 < Journ. Morph./ 1892, p. 186. 

3 ' Zeitsch. f. w. Zool./ Bd. xxxii, pp. 245 and 247, taf. xiv (1879). 



142 SYLLTDM. 

separated the sexual forms of Autolytus in a distinct family, the Amytidea. The Syllids 
followed the Grlyceridse as the eighth family of Annelids. 

Dr. Thomas Williams (1851) was not able to demonstrate the segmental organs in 
the genus Syllis "by any manoeuvre." He thought they resembled those of Psamathe 
(Gastalia). 

In his " Grlanures," Claparede (1861), after careful study at Port- Vendres, gave a more 
or less complete classification of the Syllidse up to date. He took the frontal lobes (palpi), 
the armature of the pharynx, the different kinds of cirri, and various external features — 
for instance, the presence or absence of ventral cirri — as a basis, and he produced a 
useful classification for the period. He made thirteen genera. 

Ehlers, 1 after an extended historical account of the family, gives a classification, the 
main divisions of which rest on the presence or absence of palpi, the subdivisions having 
as their basis the presence or absence of bristles on the first segment. Under those with 
palpi, the number of tentacular cirri, the armature of the proboscis, the nature of the 
palpi, and the presence or absence of ventral cirri, are used to differentiate the genera. 
In those devoid of palpi, the main groups rest on the nature of the cirri, which may 
occur on all the segments, on certain segments, or be absent. 

In an appendix 2 to this part he gives an elaborate series of references as well as 
descriptions of new and other genera arranged under the various heads. While this 
able author has greatly increased our knowledge of the group, he shows, perhaps, a 
tendency to increase unnecessarily the number of the genera. 

The terminology of Kinberg 3 (1865) was as follows : — Median tentacle = his 
tentaculum ; lateral tentacles = his antennse (antennse interiores and antennse exteriores) ; 
palpi ; posterior part of cephalic lobe (Grube) = his segmentum buccale ; buccal segment 
(Grrube) = his segmentum primum corporis. Kinberg relied for discrimination on what 
he termed " maxillse," the papillse of the proboscis, the palpi, the condition of the 
tentacles and cirri, the eyes, and the bristles. 

De Quatrefages (1865), who placed the Syllidse between the Nereids and Hesionidge, 
describes salivary glands at the posterior part of the proboscis (trompe) in the form of 
small granular masses, or occasionally simple caeca. He probably refers to the gastric 
caaca, as he mentions that the intestine follows. As a rule blood-vessels, he says, are 
absent. Only in Grubea did he find a dorsal vessel. This author classified them 
according to the mobility or immobility of the feet, the presence of cirri (dorsal and 
ventral), the armature of the "gizzard," the structure of the head and its appendages, 
and the number of " antennas " and tentacles. He, however, included members of the 
Staurocephalidse, such as Prionognathus, Hesionidae, like Kefersteinia (Gastalia), and gave 
generic distinction to sexual forms like Ioicla. 

Claparede 4 thought the posterior region of the intestine in this group had a urinary 
function. In Trypanosyllis coeliaca he found lateral diverticula of the intestine, and he 

iBorstenw./ I (1864). 

2 Ibid., pp. 255—256. 

3 < Ofvers. af k. Vet.-Akad. Fork., 5 No. 2; p. 248 (1865). 

4 'Annel. Neap./ 1868. 



SYLLILLE. 143 

notes that it would be interesting to see if T. Khronii has similar cseca. His fine figures 
and good descriptions did much for this group. 

Eisig 1 considered the T-shaped glands or cseca of the alimentary canal of this group 
and the casca of the Hesionida3 to be swim-bladders. 

Langerhans 2 follows Ehlers, Malmgren, and others, in grouping the Syllidge. He 
makes three great divisions according to the condition of the palpi. Thus the Syllidse 
proper have these separate ; the Exogonese have them fused, but prominent ; whilst the 
AutolyteaB have them fused and grooved ventrally. These groups are subdivided by the 
condition of the pharynx, the nature and number of the tentacles and tentacular cirri, 
the condition of the head, and the first segment. He makes twenty-six genera of those 
from Madeira, and points out that Syllides = Pionosyllis, and Pterosyllis = Amblyosyllis, 
and that of forty-eight different genera by various authors only eleven stand. This 
author did much to place the classification of the group on a more rational footing. 

Langerhans (1881) found a canal in the tooth of the proboscis in several genera, and 
in Syttis aurantiaca, Grube, he met with a pair of glandular sacs on the dorsum of the 
proboscis. He gives interesting notes on the budding of the Syllids. 

Haswell 3 (1885) found that Syllis corruscans had ova of a dull colour anteriorly, 
whilst posteriorly an orange-coloured region bore sperms (with testicles, Malaquin). 
The passage from the one to the other occurred abruptly at the 100th segment. At the 
same epoch he observed a pair of large eyes on the first segment of the male region, and 
the part was by-and-by detached as a pelagic male. 

De St. Joseph 4 found Labrostratus parasiticus and a Lumbriconereid parasitic in the 
ccelom of various Syllids, viz. Eusyllis moniliconis, Syllis prolifera, Pionosyllis lamelligera, 
and Grubea clavata. In this case the parasitism may be transitory, for others occur on 
sea- weed (Bytiphloea). 

Malaquin, 5 in one of the most complete treatises on the structure, development, and 
classification of the group, describes the Syllidae as a Family of Errant Annelids 
(Polychseta), his diagnosis being: Cephalic segment provided with five appendages— viz. 
two palpi, two lateral antennae and one median, and two pairs of eyes. The peristomial 
(post-cephalic) segment has in general two pairs of tentacular cirri; sometimes only a 
single pair. The succeeding segments have feet formed only of the setigerous lobe of the 
ventral division, accompanied by a dorsal and a ventral cirrus. The dorsal division of 
the foot appears at the epoch of sexual maturity. Pygidium with two cirri. Bristles 
variable — simple or compound. The most general is the compound heterogomph with 
bidentate tip. Proboscis protrusible — of two regions, the anterior (pharynx) chitinous, 
cylindrical ; the second, the muscular proventriculus and ventriculus, being a secondary 
development of the pharynx of the larva. Reproduction distinguished by the appearance 
of secondary sexual characters ; the individual is transformed (epigamy), or gives rise to 
a bud (schizogamy). 

1 < Mitth. Zool. Stat. Neap./ t. ii, p. 255. 

2 ' Zeitsch. f. w. Zool./ Bd. xxxii (1879). 

3 <Proc. Linn. Soc. N.S.W./ iv (1885), p. 733. 

4 ' Ann. Sc. Nat./ 7 e sei\, v, p. 221 (1888),and Gravier/Bull. Mus. Hist. Nat., Paris/ vi, p. 417 (1900). 

5 ' Kecherches sur les Syllidiens/ Lille, 1893, pp. 477, and 14 plates (4to). 



144 SYLLID-ZE. 

He makes four great tribes : 

« ir -j .,, Li C throughout . Exogones. 

Sylhd* with | pal t ) fused . . { at the bage onl Bus llid& . 

ventral cirrus ) 1 o. -.-..-,, 

(..separate Sylhdes. 

Palps absent . Autolytes. 

He gives an elaborate list of the genera under each tribe, with careful references, 
and a few remarks. 

This author 1 maintains that the cephalic lobe in the Syllidse and other forms is 
homologous with the ordinary segments of the annelid; thus the lateral tentacles 
correspond to the ventral division ; the median to the dorsal cirri ; the palps to the 
ventral cirri, and the tentacular cirri (anterieures, laterales, posterieures) to the dorsal 
division. This is contrary to the views of his countryman, Pruvot, who holds that the 
cephalic region consists of three segments with three corresponding nerve-centres : 
(1) The stomatogastric with its appendages the palps; (2) the tentacular segment 
(segment antennaire anterieur) with the lateral tentacles ; (3) the posterior tentacular, 
with the median tentacle (composed of two fused). Of the two views Malaquin's has 
most in its favour. 

Malaquin 2 (1893) found the pharyngeal wall to have in Syllis ; 

(1) A thick layer of chitine. 

(2) An epithelium secreting the foregoing. 

(3) A thin layer of circular muscles. 

(4) A thick longitudinal muscular coat. 

(5) A layer of peritoneal endothelium. 

The proboscis has protractor and retractor muscles. 

In the case of the proventriculus, he describes from without inwards : 

(1) The peritoneal endothelium. 

(2) Thin external circular muscular coat. 

(3) Complex radiating muscles separated by transverse diaphragms. 

(4) Internal circular muscular coat. 

(5) Columnar epithelium of the digestive tube. 

(6) The cuticle. 

The ventricule (stomach) follows, and often presents T-shaped lateral caeca, which 
have the same ciliated epithelium as the ventricule. Malaquin thinks they are filled 
with water pumped in by the proventriculus. In those without these organs, the author 
considers that the anterior part of the intestine performs their function. He divides the 
intestine into the glandular secreting anterior, and the rectal or urinary posterior. 

He uses the terms " schizogamy " and "epigamy " to distinguish the reproductive 
processes in the Syllidee. 

The former (" schizogamy ") is applied to cases in which a non-sexual nurse-stock 
gives off sexual buds. In the latter (" epigamy ") the individual is entirely sexual. 
Epigamy (Malaquin) is accompanied by the enlargement of the eyes and the elongation 

1 ( Recherches sur les Syllidiens/ Lille, 1893, p. 487. 

2 Op. cit, p. 194. 



SYLLIDS. 145 

of the antennae, the development of swimming-bristles, the development of the genital 
glands in the median and posterior regions of the body, and changes in the segmental 
organs. They are parallel to the changes seen in Nereis. 

In a subsequent paper 1 he demonstrates that epigamy occurs in the family just 
mentioned (Nereids), in the Syllids, particularly in Exogone and Eusyllis, and in the 
Hesionidse, e. g., Kefersteinia cirrata. Further, that schizogamy is present in the Syllids 
and occasionally in the Capitellidae, and in this a part of the individual only acquires 
sexual characters and is separated. In the Syllids schizogamy has various forms, viz. : 
(1) Acephalous, ex. Syllis (Haplo syllis) hamata, Clap. ; (2) Tetraglene, ex. Try jpano syllis ; 
(3) Ghsetosyllis, ex. Syllis prolifera, Krohn ; (4) Syllis arnica, De Quatrefages ; (5) Ioida, 
ex. Syllis (Typosyllis) hyalina, Gr. In Autolytus the form assumed is that of Sacconereis 
for the female, and Polybostrichus for the male. 

He points out that at the period of maturity the eyes of the Syllids having direct 
reproduction enlarge, and become lateral in position, or move to the ventral surface. In 
the pelagic sexual forms the great development of the eyes is a necessary accompaniment. 
An additional pair of pigment-specks is occasionally seen in front of these two pairs, but 
they are not true eyes. 

He shows that in the group of the Syllides (Malaquin) as contrasted with the Autolyti, 
the sexual forms resemble each other, even to the distribution of the male and female 
sexual glands. Again, they differ from the Autolyti in the variation in different types. That 
variation exists not only in distinct genera, but in closely allied species in the same genera. 

Malaquin describes the small segmental organs as having the ciliated funnel projecting 
anteriorly from the septum, and with a long, narrow canal, which opens on the ventral 
surface by an extremely minute pore. 

He gives an account of several larval stages of Syllis hyalina frequenting Polyzoa, 
and indicates how the development runs parallel with that of Eusyllis, only the formation 
of the proboscis is more rapid. His earliest stage had two setigerous segments, and the 
larval pharynx was comparatively long. The intestine opened by an anus. The larva 
with five bristled segments had the proventriculus behind the pharynx, and the rounded 
palps projecting from the sides of the head. When the larva has seven or eight setigerous 
segments the structure is much the same as in the adult. 

In Syllis, Grubea, and Exogone the ciliated apparatus of the larva disappears and the 
phenomena are abridged. 

Bisig's 2 opinion that the lateral cseca of the alimentary canal in the Syllids performs 
the function of swim-bladders does not meet with much support, especially as such a 
provision is not distinctive of highly pelagic types — types in which, for instance, swimming- 
bristles are developed on specially modified feet. The bodies of these animals are so 
plastic at the reproductive season that the modifications of an organ designed to aid in 
distributing the sexual elements could not fail to have attracted notice, and to have more 
or less differentiated pelagic from non-pelagic types in this alone. 

The SyllidaB form one of the families under the group Syllidiformia vera of Levinsen 
(1883), and are placed between the Hesionidse and ISTerillidse, a position to which no great 

1 'Zool. Anzeiger/ No. 514 (October 5th, 1896). 

2 'Mitth. Zool. Stat. Neap./ ii, p. 255 (1881). 

76 



146 SYLLIILE. 

objection can be taken. Benham, again, makes the family the first of thirteen under his 
sub-order Nereidiformia, and thus links them to such divergent forms as the Aphroditidge, 
TomopteridaB, Nephthydidaa, Amphinomidge, Grlyceridas, Sphasrodoridas, Arieiidas, and 
Typhloscolecidas. It is true that Levinsen, under the second group of his Syllidiforma, 
viz. the S. spionina, associates the Spionida3 3 Chastopteridaa, Cirratulidas, Ariciidse, 
Chlora3mida3 (?), and Ophelida3 (?), but this is all that can be said in its favour — after 
substituting Nereidiformia for Syllidiformia. 

Pruvot distinguished two forms only in the Syllides, viz. Pseudocephale (Chastosyllis, 
Tetraglena, S. arnica) and Uucejphale (Ioida). 

In the Pseudocephale the oculiferous lobes are separated from the cerebral ganglia. 
In the Uucephale the lobes are fused and contain the cerebral ganglia. 

In Benham's 1 classification this is the first family of the sub-order Nereidiformia of 
the Branch Phanerocephala, though upon what principal is uncertain. It is followed by 
such as the Hesionidas, Aphroditidaa, Phyllodocidas, Tomopteridaa, and Nephthydidse, a 
classification which appears to be at variance with natural affinities and with structure, as, 
indeed, Miss Florence Buchanan has ably pointed out. It has been considered better to 
follow the arrangement given by Malmgren and others than to adopt this classification, 
wdiich, however, in its primary branches, viz. Phanerocephala and Cryptocephala, certainly 
gives expression to actual facts. 

Calvin Mensch 2 finds that in certain parent-stocks of Autolytus ornatus, Verrill, after 
the separation of buds at the fourteenth segment, sexual elements appear in the eleventh, 
twelfth, and thirteenth segments, and he suggests the possibility of the parent-stock 
developing into a sexual form subsequently, though he has not observed the changes in 
the eyes, bristles, and other parts — probably because the examples were too young. This 
may be a case similar to Haswell's. 

Hacker 3 (1896) figures a metatrochophore of Syllis with two eyes, three separate 
tufts of cilia on the sides, and a paratroch. Gland-cells occur on the ventral aspect, and 
the alimentary canal is simple and cylindrical. He gives a later stage, in which only a 
protrotroch and a paratroch remain, whilst the long space between them has fifteen 
segments, the head is more elongated (conical), and the eyes are at the posterior border. 
A large black pigment-speck is present. Each segment has a pair of hook-like bristles. 

In a note, in 1902, on the British Syllidae, 4 the increase in the number of genera 
and species was alluded to. 

Perhaps the most natural grouping of the Syllidse is that of those authors who 
arrange them primarily into (1) those with palpi fused but prominent (Exogonea), 
(2) those with palpi separate (Syllidea proper, Langerhans), and (3) those with the palpi 
fused (Autolytea). These groups are readily subdivided by the condition of the pharynx, 
the nature and number of the tentacles and tentacular cirri, the condition of the head and 
first segment, and especially the structure of the feet and bristles. Moreover, a fourth 
group, the Eusyllidea (Malaquin), in which the palpi are fused at the base only, may be 

1 f Camb. Nat. Hist./ vol. ii, p. 306 (1896). 
3 ' Zool. Anzeiger/ June 15th, 1896. 

3 "Pelagische Poly chat enlarven/' ' Zeitsch. f. w. Zool./ Bd. Ixh, p. 82, Taf. iii, figs. 6 and 7. 

4 Mcintosh, < Ann. Nat. Hist./ ser. 7, vol. ix, p. 296. 



NEBILLA ANTENNATA. 147 

interpolated between the Exogonea and Syllidea. Nerilla may be left at present in an 
uncertain position. 

Syllids especially abound in the Mediterranean and at Madeira (Langerhans), and 
they range from tide-marks to 1380 fathoms (Ehlers, Syllis abyssicola). 



Genus XLIV. — Nerilla, 0. Schmidt, 1848 (provisionally). 

This form (figs. 50, 51, and 52), as Malaquin truly states, is at present in an 
uncertain position, though most authors place it in relation with the Syllidse. 
De Quatrefages gave it a new title, viz. Dujardinia, and inserted it between Microsyllis, 




Fig. 50. — Nerilla antennata, Scnm. (after Claparede), from the ventral aspect. (Enlarged.) 

Schmardia, and the Amytidea. A brief notice of it is, perhaps, necessary, as it has been 
found in the British area. Its structure and reproduction need careful re-examination. 
It has been thought unnecessary to constitute a family for it, as some have done, in the 
present condition of information on the subject. No example has been seen by the author. 

1. Nerilla antennata, 0. Schmidt, 1848. 

Synonyms. 

1848. Nerilla antennata, 0. Schmidt. Reise nach der Fiiror, Neue Beitrag., Nat. Wiirmer, p. 38, 

Taf . Ill, f . 8 and, 8 a, fide ant. 
1851. „ „ Grube. Fam. Annel., p. 62. 



148 NERILLA ANTENNATA. 

1854. Nerilla antennata, Leuckart. Archiv Naturges., 1854, p. 327. 
1863. „ „ Claparede. Beobacht., p. 48, Taf. xii, f. 16—20. 

1865. Dujardinia „ De Quatrefages. Annel., II, p. 69. 

1877. Nerilla „ Barrois. Compt. Bend. Ac. So., t. lxxxv, p. 288. 

1883. „ „ Levinsen. Vidensk. Meddel. Foren. Kjobenh, p. 250. 

1884. „ „ Webster and Benedict. Bep.U. S. Comm. Fish for 1881, p. 711, pi. ii and 

iii, f. 17—23. 
1893. ,; „ Flor. Buchanan. Kep. Brit. Assoc, 1892, p. 358. 

Habitat. — Plymouth (Miss F. Buchanan). 

Faroe Islands (0. Schmidt). St. Vaast-la-Hougue, and other shores of France 
(Claparede and Barrois). Amongst corallines at Chausey (De Quatrefages). 

Head (fig. 51) rounded, bearing three rather long, articulated tentacles, a median 
and two lateral. The palpi (?) are small, clavate, and lateral in position. Byes four, 
brown, the anterior pair wider apart than the posterior ; all are provided with lenses. 



, U 



\ 



J/ 






Fia. 51.— Head of Nerilla antennata. Enlarged. After Claparede. 

Body (fig. 50) about 1 mm. long, and of nine segments, the first bearing a pair of 
articulated cirri shorter than the tentacles. The mouth is large and somewhat triangular, 
and it is followed by a muscular pharynx and a stomach. No constrictions appear on 



._ _.;o 



Fia. 52. — Foot of Nerilla antennata. Enlarged. After Claparede. 

the intestine, which in the middle region is glandular. Numerous tufts of vibratile cilia 
occur on the body, and one on each side behind the foot. Segmental organs are present. 
The body terminates in two cirri posteriorly. 

Foot (fig. 52) with only a lateral cirrus and simple capillary bristles. A glandular 
organ occurs in most of the segments of the body after the first, with a ciliated opening 
between the feet. The gland has refracting corpuscles (Claparede). 

Reproduction. — Ova large, developed in the lateral region so as to affect the outline 
of the body. In the males the sperms float freely in the body-cavity. 

Claparede thought that this form lay between the errant and the sedentary annelids. 
The free condition of the alimentary canal in the body-cavity, and the absence of 
dissepiments and compound bristles are interesting. Barrois (1877) found both males 
and females and observed the development. 

Levinsen (1883) makes a new family for this genus, viz. Nerillidse. Until further 
investigation is made, however, it is, perhaps, well to leave the genus as an appendix to 
the Syllidas. 



EXOGONE. 149 



Group L— EXOGONEA. 

Small syllids, with ventral cirri; palpi fused throughout. Segments few and 
variable. Appendages short. Proboscis straight. Bristles with simple tips to terminal 
pieces. Reproduction generally direct; rarely by stolons (Malaquin). 



Genus XLV. — Exogone, Mag. (Ersted, 1845. 

Head composed of two distinct segments ; palpi prominent, fused ; three clavate 
tentacles fixed to the middle of the head; eyes four; one pair of tentacular cirri (short) ; 
ciliated pits at the posterior part of the cephalic segments. Body small, linear, of a fixed 
number of segments. Proboscis short, with a single tooth anteriorly ; lateral casca of the 
stomach small. Two clavate or filiform anal cirri. Feet small, papilliform ; dorsal and 
ventral cirri sub-equal, clavate, or conical. A single fascicle of falcate bristles in the 
female ; in the male, all the segments (the first eight excepted) with falcate and long 
capillary bristles. Reproduction direct. Ova borne on the dorsum or on the ventral 
surface, with embryos and larvae in the latter case attached near the segmental orifice. 

This group received a notable contribution from the late distinguished anatomist of 
Wiirzburg, viz. Professor Kolliker, 1 who, in 1845, appended to a paper, by his friend 
H. Koch, in the f Xouveaux Mem. de la Soc. helvet.,' viii, an account of three forms from 
Naples, viz. Exogone (Erstedi, Exogone cirrata, and Cystonereis Edwardsi. The first has 
two pairs of tentacles, eggs in sacs at the sides, and long swimming-bristles. The second 
has the palpi apparently fused at the base, two pairs of tentacles, and also eggs in sacs at 
the sides. The third has four pairs of tentacles, no visible palpi, and the eggs are borne 
ventro-laterally. None of these correspond with Professor Graham Kerr's form from the 
Clyde with eight ovigerous sacs. 

Considerable confusion has occurred in this group from the imperfect descriptions of 
authors. Thus the Exogone Kefersteinii of Claparede 3 appears to be Exogone gemmifera, 
whilst Ehlers, misled by the supposed absence of the ventral cirri, constituted a new 
genus, ExotoJcas, for the inclusion of both as separate species. Claparede's genus, 
Poedophylax 3 is in the same position as the Exotohas of Ehlers, both referring to Exogone. 
The examples of the genus have hitherto been so rarely procured in this country that 
much yet remains to be done both in synonymy and investigation of structure. 

The genus seems to be cosmopolitan in its distribution, ranging from the Antarctic 
seas northward, a form very similar to Grubea pusilla, and bearing ova on the dorsum, 
for instance, occurring at Kerguelen, and entangled amongst the bristles of Loetmonice 
producta, just as an Irish member of the group was found on Aphrodita aculeata. It is 
probable that considerable additions will yet be made to the group in British waters, 
especially on the southern and western coasts. 

1 ' Nachwort zu dem vorhergehenden aufsatze/ op. cit., p. 13. 

2 'Beobacli. ueb. Anat. u. Entwickl./ p. 42, Taf. xii, f. 3—6. 

3 ' Annel. Nap./ p. 210. 



150 EXOGONE NAIDINA. 

De Quatrefages (1865) instituted the genus Urania for the Exogone pusilla of 
Dujardin, 1 which he distinguished from (Ersted's species. 

1. Exogone naidina, Mag. (Ersted, 1845. 

Specific Characters. — Head bluntly conical, the fused palpi being only marked by a 
notch dorsally, but clearly seen inferiorly. Eyes four, connate, the anterior the larger, 
and furnished with cuticular lenses. Median and lateral tentacles short and clavate. 
Body linear, slightly tapered anteriorly, and more so posteriorly, 2 — 3 mm. long, and with 
19 — 35 segments; pellucid, yellowish from the intestine; terminating posteriorly in two 
short, clavate cirri. Proboscis with a median tooth, proventriculus whitish and granular. 
Foot with the setigerous region obliquely truncated, and bearing falcate bristles with 
short terminal pieces furnished with a simple hook; capillary swimming-bristles in 
male. Dorsal cirri slightly tapered ; longest in the anterior third ; ventral cirrus shorter. 
Both were stated by M. (Ersted to be subclavate. Ova borne on the ventral surface. 

Synonyms. 
1845. Exogone naidina, M. (Ersted. Arch. f. Nat., xi, p. 20, Taf. 2, f. 1—14. 
1851. „ ,, G-rube. Fam. Annel., p. 62. 

1855. Syllis longiseta, G-osse. Ann. Nat. Hist., 2nd ser., xvi, p. 32, pi. iv, f. 14 — 21. 
1865. Exogone naidina, De Quatrefages. Annel., ii, p. 33. 
1874. „ „ Malm. Op. cit. Goteb, p. 82. 

1879. „ „ Langerhans. Zeitsch. f. w. Zool., Bd. xxxii, p. 563. 

„ „ „ Tauber. Annul. Danic., 95. 

1883. „ „ Levinsen. Yidensk. Meddel. Foren. Kjobenh., p. 244. 

1885. „ „ Cams. Fauna Medit., p. 232. 

1890. „ „ Giard. Bull. Sc. Fr. Belg., t. xxii, p. 78. 

1893. „ „ Levinsen. Yidensk. Ud. ' Hanchs/ p. 330. 

Habitat.— Weymouth (P. H. Gosse (?) ). 

This species has been entered on the responsibility of various authors who have 
considered Gosse's Syllis longiseta as identical with it. So far as can be made out this 
conclusion seems to be reasonable both as regards description and figures, but no oppor- 
tunity should be lost of re-examining this interesting form. 

Magnus (Ersted first described this small Syllid from Lille Belt near Striib, and his 
account agrees generally with that of Gosse. He shows the eggs on the ventral surface, 
and gives an interesting description of the development of the young from the egg up to 
the fourteenth day, when it is furnished with a median and two lateral tentacles, four 
separate small eyqs, four bristled segments, and two anal cirri. 

The Syllis longiseta of Gosse 2 from Weymouth, for which De Quatrefages 3 constituted 
the genus Gossia, is a bud with long swimming-bristles and a head like Exogone if the draw- 
ing and description can be relied on. The head shows three short clavate tentacles, and the 
tips of the bristles are simple. De Quatrefages repeats his reference to Gosse's species by 
1 ' Ann.. Sc. Nat./ 3 e ser., t. xv, p. 298, pi. v, fig. 9 and 10. 
3 c Ann. Nat. Hist./ ser. 2, vol. xvi, p. 32, pi. iv, figs. 14—21 (1855). 
3 < Annel./ II, p. 49. 



EXOGONE GEMMIFERA. 151 

making a second new genus (Syllia) for it. 1 The example from Weymouth secreted a 
tube fixed to the glass, and was procured amongst sea- weeds. It may be a male bud 
of Exogone gemmifera. 

2. Exogone gemmifera, Pagenstecher, 1863. Plate LIX, figs. 5 and 6. 

Specific Characters. — Head with fused palpi forming an even rim. Tentacles clavate. 
Eyes four, with lenses, connate on each side, the anterior the larger. Body about 3 mm. 
long, with 30 — 32 segments, yellowish or pinkish, bearing embryos along the dorsal or 
the ventral surface. Tail terminating in two slender cirri. 

Synonyms. 

1863. Exogone gemmifera, Pagenstecher. Zeitsch. f . w. ZooL, Bd. xii, p. 267, Taf. xxv and Taf. xxvi, 

f. 1, 2, 6, 7, 8. 

1874. „ naidina, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 198. 

1875. „ „ idem. Invert, and Fishes St. Andrews, p. 121. 

1884. „ gemmifera, Yiguier. Arch. Zool. Exper., II, p. 69, pis. hi and iv, f. 21 — 37. 
1890. „ „ G-iard. Bull. So. Fr. Belg., t. xxii, p. 78. 

Habitat. — Under a stone near low water-mark, East Pocks, St. Andrews (E. M.), 

1864. Lochmaddy, North Uist, between tide-marks. 

Between tide-marks at Cette, Coast of France (Pagenstecher) ; Bay of Algiers 
(Viguier). 2 

The Head has anteriorly a rounded border of the fused palps with a slight median 
notch. On the ventral surface a median groove indicates the palps more clearly. The 
cephalic region proper bears two brownish-red eyes on each side, the anterior the larger, 
and all furnished with lenses. A median and lateral tentacles are situated in front of the 
eyes. They are short, enlarged at the base, and the lateral are rather blunt distally, 
whilst the median is the longer. The tips of these have palpocils. The cephalic segment 
is closely united with the buccal, but, as Viguier points out, they are distinguished by a 
line behind the eyes. 

Body translucent, rather less than one eighth of an inch in length, slender, very 
slightly tapered anteriorly, but more distinctly so posteriorly, where it terminates in two 
slender cirri on each side of the anus, and Pagenstecher found refractive corpuscles near 
the bases of these cirri. The first segment has only a small cirrus and is separated by an 
undulating line from the buccal (Claparede and Viguier). A minute ciliated groove (sense 
organ ?) occurs immediately in front of the cirrus (Viguier). The second bristled segment 
is peculiar in having no dorsal cirrus. 

The proboscis has a single tooth anteriorly (Plate LIX, fig. 6), and Viguier indicates 
a break in the pigment about its middle. The proventriculus is situated opposite the 

1 Aimel., II, p. 80. 

2 It is interesting to find that this observer obtained his specimens by following a very old plan 
at St. Andrews — viz. by removing portions of rocks, stones, or calcareous algas at low water and placing 
them in vessels of sea-water, when the minute annelids hy-and-by leave the shelter of zoophytes, 
sea-weeds, or crevices, and appear at the water-line of the vessel. 



152 EXOGONE GEMMIFERA. 

seventh segment (fourth bristled) and is whitish, with transverse rows of glands to the 
number of about a dozen (Viguier). It has a rhomboidal outline or resembles a square with 
the angles rounded. The stomach behind has two lateral cseca (the salivary glands of 
Pagenstecher). On each side of the commencement of the proboscis is a brownish organ 
placed opposite the first setigerous foot. 

The intestine is of a yellowish or brownish hue, and has a moniliform outline, and 
oily granular matter in its glandular walls. 

The typical foot has a short and somewhat conical dorsal cirrus, and a ventral cirrus, 
with the intermediate setigerous region. The latter, in the males and certain females, 
has a tuft of long swimming-bristles between the dorsal cirrus and the setigerous region. 
The superior ventral bristle is the strongest, with a curved shaft and a short terminal piece. 
The next has a longer tapering terminal piece, then follow two with bifid terminal pieces 
like those of Autolytus (though Viguier's figure is peculiar), and, lastly, a simple curved 
bristle. The curvature of these inferior bristles resembles that of a gun-stock. 

The nine pairs of larvaB in the example from St. Andrews are flask-shaped, and with 
a reddish-pink granular mass of yolk at the enlarged central portion, and have two 
processes (caudal cirri) posteriorly at the attached end, and three at the anterior or free 
end, viz. the median and lateral tentacles. They corresponded, therefore, with the stage 
shown in Viguier's Plate IV, fig. 29, the French author having, under favourable oppor- 
tunities, given an excellent original account of the escape of the ova, their attachment 
to the region of the segmental aperture a little in front of the ventral cirrus, and their 
subsequent development. 

In a female, procured between tide-marks at Lochmaddy, about a quarter of an 
inch in length, pale — almost translucent — marked down the centre by a moniliform yellow 
band from the intestine, twenty-one pairs of flask-shaped, rose-pink embryos with a red 
spot in the centre occurred on the ventral surface. The head and nine segments only 
were devoid of them in front, whilst posteriorly the embryos nearly reached the tip, 
which may, however, have been broken. 

Viguier found the male with short swimming-bristles, and the perivisceral cavity 
from the ninth or tenth normal segment to the posterior end— with the exception of the 
tip of the tail — filled with sperms, the sexual region thus occupying about twenty 
segments. The short swimming-bristles begin somewhat behind the first sexual 
segment. The sperm-sacs (his seminal vesicles) appear to be fusiform and to stretcli 
across each segment to open at the ventral pore on each side, a little behind the ventral 
cirrus, at which point also a group of four bacillary corpuscles is found. 

In the female, Viguier found some with and some without the swimming-bristles, 
which, as in the male, commenced at the first sexual segment, or a little behind it, and 
continued to the ante-penultimate segment. The ova are developed under the intestine in 
pairs in the ccelomic space of each segment, which they by-and-by fill. They have 
externally the vitelline membrane and are extremely mobile, in short, almost amoeboid in 
their nature, for when ripe they pass outwards, apparently by the segmental aperture. All 
the ova are about the same stage in development, and appear to be fertilised after their 
escape and attachment. Viguier figures the two-, four-, eight-celled, and the morula 
stages, and also what he regards as invagination at a later stage, the aperture (mouth) 



EXOGONE GEMMIFERA. 153 

being on the ventral surface, but it afterwards shifts forward to the anterior end. Then 
the median tentacle appears, afterwards the two lateral and the caudal cirri on each side 
of the point of attachment. This is the stage subsequent to that which the sole example 
from St. Andrews had reached (Plate LIX, fig. 5). In front, that is at the distal or 
free end of the larva, are the three tentacles, the fused palps forming a ventral projection, 
the pharyngeal region and the proventriculus follow, then a reddish-brown vitelline 
mass with oleaginous globules, behind which is the intestine and the anus. The whole 
is enveloped by the vitelline membrane which, he (Viguier) holds, becomes transformed 
into the cuticle of the larva, as Stossich found in Serpula. The chief features observable 
before the larva leaves its parent are the appearance of eyes with lenses, tentacular cirri, 
the more definite outlining of the pharynx (proboscis) with its tooth, and outlining of 
the proventriculus, as well as the intestine, together with the occurrence of four 
bristled feet with dorsal and ventral cirri, the dorsal, however, being absent on the 
second bristled segment, as in the adult. 

Habits. — This form dwells between tide-marks, and seems to differ considerably 
from the pelagic types. It is sluggish — rolling in the vessel, and appears to be unable 
to use bristles or feet actively when carrying the embryos. On immersion in spirit the 
embryos on its ventral surface were thrown off. 

Exogone gemmifera, from Cette, was first described by Alex. Pagenstecher, in 1863, 
the year before the example from St. Andrews was found. His specimen was 3 mm. 
long, had thirty -two segments, and bore what he thought were buds from the ninth 
to the twenty-second bristled segment on the dorsum. Moreover, the dorsal and 
ventral surfaces of the buds corresponded with those of the parent. They presented 
head, mouth, "upper lip," four brownish-red eyes, and four bristled feet, and were attached 
by a posterior pedicle. The csecal digestive canal was brownish-red, and the proboscis 
had a tooth anteriorly. He notes sac-like' segmental organs between the seventeenth 
and twenty-ninth segments, and also the early eggs with the tentacles indicated in the 
embryo. His supposition that the embryos were buds has been fully dealt with by 
Viguier. 

The larval forms found on this species are very similar to those on Syllides pidliger, 
Krohn. 

Claparede (" Glanures," 1864) points out that Pagenstecher was wrong, but that in 
all probability the larvas in this (which he calls a Sylline, Grube) arose from eggs, as in 
Exogone and Syllides — attached to dorsal cirri (Krohn) — alternately with blanks. 
Claparede notes that it is a fact that in this and other forms long and short cirri occur 
alternately. The young are detached from Syllides pulliger when they have six segments, 
the third segment, however, being devoid of the dorsal cirrus. 

Giard (1890) thinks that this species is identical with E. liefer 'steinii, Claparede, and 
probably also with the Pdedojphylax claviger of the same author. 

Malaquin mentions that the young on being set free have the larval pharynx and the 
proventriculus (the so-called dipharyngeal condition), and when they have four setigerous 
segments the characters are nearly those of the adult. 

Malaquin found in September several mature male buds of thirteen segments of 
Exogone gemmifera, with swimming-bristles and heads recently formed, with two large 

77 



154 EXOGONE GEMMIFERA (?) SCHARFFI. 

anterior eyes on the ventral surface, and a smaller dorsal pair. No proboscis existed, only 
the alimentary canal. He decided these to be buds after the type of Tetraglene. 
Langerhans previously had found a similar condition. 

So long a period has elapsed since the examples (which unfortunately have been lost) 
occurred at St. Andrews and Lochmaddy that considerable doubt exists on some points, 1 
but they appear to pertain to this species. 



EXOGONE GEMMIFERA ?, var. SCHAREFI. 

Head a smoothly rounded lobe divided by a constriction. Behind the latter on each 
side is a pair of connate eyes with lenses, the anterior pair being large, the posterior 
minute. Body 3 — 4 mm. in length, and composed of about thirty-six segments. It 
terminates posteriorly in a slightly enlarged blunt process with two cirri. Proboscis a 
short, narrow, straight tube. Proventriculus also short. The feet have lost the dorsal 
cirri. Setigerous region conical, bearing a series of translucent bristles with the end of 
the shaft dilated and notched like the cannon-bone of a sheep. The upper bristles have 
a long tapering terminal piece, like that of Phyllodoce, whilst the inferior have a short 
terminal piece with a claw at the tip. 

Habitat. — Amongst the bristles and hairs near the tail of Aphrodita aculeata procured 
at Long. 55, R. I. Academy's Expedition, 1886. 

Head forms a smoothly rounded lobe with a constriction marking off an anterior 
from a posterior region. Just behind this constriction at each side is a pair of connate 
eyes, each with a clear lens-like body, the anterior large, the posterior minute. The 
example had no tentacles or tentacular cirri, but the margin of the snout may have had 
sensitive papillae. 

Body about 3 or 4 mm. long, elongated, composed of about thirty-six segments, the 
first following the region on which the eyes are placed. Each segment consists of a 
setigerous lobe and a short dorsal cirrus. 

The proboscis appears to be short, as the proventriculus commences at the fourth 
bristled segment and ends behind the fifth. The intestine then passes backward to the 
anus. Stretching from the mouth, which lies a little in front of the eyes, is a clear, 
straight, narrow tube, which may represent the proboscis thus curiously reduced. 

The posterior end terminates in a slightly swollen blunt process with two cirri. 

The foot has no dorsal cirrus in the preparation, but it may have been lost. The 
setigerous region is conical, and bears a series of translucent bristles with the distal end 
of the shaft curved and slightly dilated, the end being notched like that of a sheep's cannon- 
bone. The upper bristles have a long tapering terminal piece like that of a minute Phyllodoce. 
The inferior bristles have a short terminal piece with a claw at the tip, but the whole 
structure is so minute that whether it is simple or bifid could not be determined. 
Apparently it is simple. In lateral view the bevelled region of the dilated end of the 
shaft is truly Syllidean, and when seen on end the blunt point formed in those devoid of 
terminal pieces is also characteristic. 

1 e. g., the dorsal or ventral attachment of the embryos. 



XENOSYLLIS (?) KESTBERGL 155 

Whether this minute species lives amongst the bristles of Aphrodita or was only 
entangled therein in the dredge is unknown. 

This species somewhat resembles Claparede's 1 Exogone Kefersteinii from St. Vaast-la- 
Hougue, and may be a young form of this or an allied species such as Exogone gemmifera, 
yet the structure of the bristles, the form of the cephalic segment, and other details differ. 

The specimen has been lost or mislaid since the description was drawn up, so that 
drawings could not be made. 



Genus XL VI. — Xenosyllis, Marion and Bobretzky, 1875. 2 

So far as can be observed in the single example the following characters appear to 
coincide with this genus, the type of which is the Syllis scabra of Ehlers. 3 Head with 
comparatively large palpi fused at the base. Tentacles three, thick, short, and monili- 
form, as also are the tentacular cirri. Body 4 somewhat short and broad, with rather 
long setigerous processes to the feet, short, moniliform dorsal cirri, and a broadly lobate 
ventral cirrus. Bristles strong, terminal piece simple. 



1. Xenosyllis (?) Kinbergi, 5 n.s. Plate LIX, fig. 7; Plate LXX, fig. 2— foot; Plate 

LXXVIII, fig. 10— bristle. 

Specific Characters. — Head with two comparatively large palpi, fused at the base, 
visible ventrally, and prominent laterally. Median and lateral tentacles short, moniliform. 
Tentacular cirri two on each side, moniliform. Body 10 mm. long, having about fifty 
bristled segments, about equally narrowed anteriorly and posteriorly. Posterior end 
imperfect. Foot : dorsal cirrus short, stout, slightly tapered towards the tip ; nine to 
ten segments in the anterior third. Setigerous region comparatively long. Bristles 
strong, terminal piece with a simple hook — longer in the posterior region of the body. 
Ventral cirrus bluntly rounded at the tip, and in shape broadly lobate. Tip of the 
spines bent. 

Habitat. — Berehaven, Ireland, Royal Irish Academy's Expedition, 1885, along with 
Syllis comnta, H. Pathke. 

Head with comparatively large palpi fused at the base. The median and lateral 
tentacles are short, but distinctly moniliform, the former being the longer. Two small 
eyes occur on each side posteriorly, arranged in the usual trapezoid, the anterior pair 

i < Beobach./ p. 42, Taf . xii, figs. 3—6. 

2 < Ann. Sc. Nat./ 6 e ser., ii, p. 26. 

3 < Borstenw./ i, p. 244, pi. xi, figs. 1—3 (1864). 

4 No indication of a scabrous condition was seen. 

5 Named after the distinguished Swedish zoologist, Professor Kinberg, who has done so much 
valuable work in the Annelids. 



156 SPH^JROSYLLIS HYSTRIX. 

being the wider apart. In addition, a small pigment-speck is situated internal to the 
base of the anterior tentacles. 

Body 10 mm. long, about equally narrowed anteriorly and posteriorly, and composed 
of about fifty bristled segments, but the condition of the posterior end is doubtful. It 
terminated in two rounded lobes, but as the last foot and bristle-bundle projected from 
the outer side of each lobe, the condition is unusual and probably indicates injury. 

In the foot (Plate LXX, fig. 2) the dorsal cirrus is short and stout, slightly tapered 
towards the tip, and composed of nine to twelve segments at the anterior third, but the 
number apparently diminishes posteriorly. The setigerous region is comparatively long 
throughout the greater part of the body, and viewed from above is nearly cylindrical, or 
with a forward curve, and in some slightly narrowed at the base. In lateral view a 
slight hollow occurs below the spine superiorly, and then the margin slopes inward. The 
region is thus bifid at the tip. Each bears a projecting tuft of somewhat strong bristles 
of the ordinary type (Plate LXXVIII, fig. 10), the bevelled end of the shaft being 
smooth, and the comparatively short terminal piece having a simple hook distally, the 
figure representing one of those with shorter tips from the middle of the body. 

The anterior feet have bristles with more slender shafts and longer terminal pieces, 
the posterior stouter shafts and shorter terminal pieces. The tip of the spines in some is 
bent and projects slightly beyond the surface. The ventral cirrus is bluntly rounded at 
the tip and in shape broadly lobate. 

This form is distinguished by the comparative shortness of the body, the short but 
distinctly moniliform dorsal cirri, by the lobate form of the ventral cirri, and the simply 
hooked terminal pieces of the bristles. It has certain resemblances in the shortness and 
marked moniliform condition of the cirri to the Syllis scabra of Bhlers. 1 



Genus XLVII. — Sph^rosyllis, Claparede, 1863. 

Cephalic and peristomial segments coalescent. Palpi small, continuous with the 
head, fused dorsally. Tentacles (three), tentacular cirri (two), and dorsal cirri short, 
enlarged at the base. Eyes usually four. Body comparatively short, beset with papilla?. 
First segment with a single tentacular cirrus on each side 2 , and bristles. Pharynx with a 
single tooth ; proventriculus short. Ventral cirri short and filiform. Xo alternation of 
generations (Claparede). Ova borne on the ventral surface. 



1. Sph^rosyllis hystrix, Claparede, 1863. Plate LIX, figs. 3, 4, 8; Plate LXX, fig. 1 

foot; Plate LXXVIII, figs. 11-13— bristles. 

Specific Characters. — Head with the palpi tapering in front. Eyes four, red, the 
anterior pair having lenses. A median and two lateral tentacles much enlarged at the 

1 ( Borstenw./ i, p. 244, pi. xi, figs. 1—3. 

3 Claparede considers the pair on the united cephalic and peristomial segment tentacular cirri, 
whereas Ehlers gives five tentacles to the head. 



SPH^POSYLLIS HYSTBIX. 157 

base and tapering gradually towards the tip. A tentacular cirrus on each side. Body 
rather less than a quarter of an inch in length, tapered anteriorly. Segments thirty-two. 
About ten segments in front free from eggs, then a double series of pale rose-pink eggs 
follows. Proboscis has lateral glands and a tooth in front. Proventriculus with twelve 
rows of points (De St. Joseph). Foot has a small superior cirrus much enlarged (almost 
globular) at the base and marked by minute and scattered tubercles. Setigerous region 
with a fevy long papillge, two of which near the apex appear to be regular — viz. one 
pointing forward and the other backward. The falciform bristles bear a somewhat 
elongated terminal piece ending in a simple claw. A single simple bristle slightly 
curved toward the tip occurs amongst them, and from the ninth segment to the tail a 
tuft of long swimming-bristles. From the fifth segment there is a capsule with 
rhabdites in each foot ventrally. 



Synonyms. 

1868. Sphzerosyllis hystrix, Claparede. Beobach, p. 45, Taf. xiii, f. 36 and 37. 

1864. „ }i idem. Gllanures, p. 86, pi. vi, f. 1. 

1865. „ }} De Quatrefages. Aimel., ii, pp. 52 and 646. 

1869. „ „ Mcintosh. Trans. R. S. Edin., t. xxv, p. 416, pi. xv, f. 10; pi. xvi, 

f. 9. 

1874. „ „ Marenzeller. Sitzb. Wien., p. 25 (Adriat. Annel.). 

1875. „ „ Marion and Bobretzky. Annel. Marseil., p. 44; Ann. Sc. Nat., 6 e ser., 

t. ii, p. 44. 
1879. „ ,, Langerhans. Zeitsch. f. w. Zool., t. xxxii, p. 567. 

„ „ „ Tauber. Annul. Danic, p. 95. 

1884. „ „ Yiguier. Arch. Zool. Exper., 2 e ser., t. ii, p. 98. 

1885. „ „ Cams. Fauna Medit., p. 233. 

1887. „ „ De St. Joseph. Ann. Sc. Nat., 7 e ser., t. i, p. 204, pi. x, f. 79, 80. 

Habitat. — Under a stone, near low water, Lochmaddy, August, 1865. In the tow- 
net, off County Down, Ireland, August, 1902 (Professor Gr. S. Brady). 

Dinard, France (De St. Joseph), Madeira (Langerhans), Mediterranean, Atlantic, 
and North Sea. 

Head (Plate LIX, figs. 4 a and 8) with the palpi fused, except at the tip ; two lateral 
tentacles, much enlarged at the base and tapering gradually towards the tip. Their 
surface is somewhat rough. A median tentacle, rather larger, but of similar shape, 
arises just in front of the eyes (fig. 4 a). Eyes four, red, the anterior pair furnished 
with lenses. In an example from the Minch there were six eyes — two larger posteriorly, 
quite separated from each other, and two small round ones in front. De St. Joseph 
found the head (peau) covered with papillaa and incrusted with muddy particles. 

Body pale, nearly translucent, marked by a central moniliform yellow band, rather 
less than a quarter of an inch long, and tapered anteriorly to the small snout. Segments 
19 — 32 (30 — 38, De St. Joseph). The tail has two swollen cirri. 



158 SPH^ROSYLLIS HYSTEIX. 

Proboscis with a tooth and with a series of papillae. The proventriculus is typical 
of the family. 

De St. Joseph observes that the proboscis has lateral glands and a tooth in front. 
The proventriculus has twelve rows of points. In the lateral pouches of the ventricle 
is active ciliary circulation. 

In the foot (Plate LXX, fig. 1) the superior cirrus is small, much enlarged at the 
base — especially anteriorly, and the posterior cirri longest. It is marked by minute 
and scattered tubercles. The setigerous region presents long fleshy papillas, two of 
which, near the apex, appeared to be regular in position — viz. one pointing forward and 
the other backward. Each bears a number of jointed bristles (Plate LXXVIII, figs. 11 a 
and 11 b), and the somewhat elongated terminal piece with a simple claw at the tip and 
serrated along the anterior edge. In addition is a single simple bristle (figs. 12 and 13) 
slightly curved towards the tip, and apparently dorsal in position. Then a tuft of 
long simple bristles (swimming), stretching far beyond the body from the ninth segment 
to the tail. In an example from the Minch no elongated swimming-bristles are present, 
and the terminal pieces of the jointed bristles are somewhat longer. The ventral cirrus 
is slender and short. 

De St. Joseph describes a rounded gland containing rods at the base of each foot 
from the fifth to the last segment but one. 

Reproduction. — Professor Gr. S. Brady, who has for many years sent interesting 
specimens, procured several pelagic examples in the tow-net off the coast of Co. Down. 
All the females (Plate LXVI, fig. 3) carried eggs, but no example out of eight or ten had 
swimming-bristles. On the other hand, the only two males bore swimming-bristles 
(Plate LIX, fig. 4), and were distinguished in other respects from the females by the 
smaller number of segments, which, moreover, were more deeply cut, so that the aspect 
was moniliform. The long bristles commenced on the eleventh bristled foot and 
continued to the posterior end, which, however, was incomplete. These pelagic forms 
had the eyes considerably larger than the littoral. 

The females of these pelagic Syllids have a series of ova projecting freely from the 
dorsum in some cases, and in others from the ventral surface, and apparently fixed to 
dorsal or ventral cirri or between the feet, or to the feet by a tough thread. 

In an example of medium length they formed four irregular rows, occupying more 
than the median third of the dorsum, about five bristled segments at each end being free. 
By transmitted light the ova had a reddish speck (oleaginous ?), and readily became 
detached during the manipulations. 

In a small female considerable elongation of the bristles was present, but no capillary 
swimming-bristles could be distinguished. 

Habits.— So far as observed, these Syllideans frequent the under-surfaces of stones 
near low-water mark, especially in their earlier stages, but when carrying ova in the one 
case, or ripe sperms in the other, they appear to become pelagic. The great proportional 
size of the ova would seem adverse to pelagic activity, but their specific gravity is 
probably such as to aid the annelids in this nomad life, a supposition the more likely 
as the majority of the females obtained had no swimming-bristles. 



MICROSYLLIS. 159 

De St. Joseph found a male, 4 mm. in length, of thirty-eight segments, with natatory 
bristles and sperms from the eleventh to the thirty-fourth segment. In a female, 2*40 mm. 
long, of thirty segments without natatory bristles, the eggs were attached from the eighth 
to the twenty-second under the ventral cirri by a membrane. Each of the fifteen 
ovigerous segments had two eggs containing an embryo with a head, three tentacles, and 
two eyes, three segments furnished with a dorsal and ventral cirrus, a group of bristles 
with terminal pieces having a simple tip, and a single long simple bristle. The body of 
the embryo is terminated by an anal segment with two long cirri which are not dilated 
at the base like the other appendages. Internally is a rudiment of a proventriculus 
without a proboscis, and a mass of viteHus. They were ready to hatch. 

In another incomplete female without natatory bristles there were twelve older 
embryos (placed two and two behind the twelfth segment) with seven segments. 

Claparede (1863) pointed out that the fused palpi with the median groove approached 
Sylline, whilst the enlarged basal region of the cirri and their papilla? reminded him of 
the condition in Sphmrodorum {Ephesia) and the Siphonostomum villosum of Pathke. He 
observed cells with rhabdites in the feet, as in Sphserodorum. 

Viguier 1 describes a bud (of fourteen segments) of this species having two large 
ventral eyes and two smaller dorsal, and with swimming-bristles from the first post- 
cephalic segment backward. Malaquin thinks that the sexual elements had probably 
been shed. 

Sphserosyllis ovigera, Langerhans 3 (from Madeira), is entered in the fauna of 
Plymouth, and Dr. Allen kindly forwarded a preparation (slide) of it. The eyes (Plate 
LVIII, fig. 19) in this example are more closely arranged on each side, and no ova are 
visible. So far as could be observed the structure of the foot, the presence of the single 
slightly curved and pointed bristle, and the structure of the compound bristles all agree 
with S. hystrix. The skin has numerous papilla?, to which and to the dorsum sand-grams 
and mud adhere. The perusal of the description and figures of Langerhans does not 
conduce to greater certainty, for almost all would apply to 8. hystrix, unless the figure of 
the compound bristle is held to be accurate— viz. with a spine on the convex border of the 
enlarged terminal region of the shaft. Dr. Allen's example does not show any process of 
this kind. 



Genus XL VIII. — Miorosyllis, Claparede, 1863. 

The pre-oral and peristomial lobes not separated; only two tentacles. Palpi small. 
Tentacular and dorsal cirri very small. First segment with only one tentacular cirrus on 
each side. Body typical in shape, with 17—30 segments. Proboscis as in Syllis, with a 
tooth in front and a proventriculus. Foot with a small dorsal cirrus; short setigerous 
region ; bristles with simple tips. No ventral cirrus. 

1 Fide Malaquin, ' Arch. Zool. Exper./ 2 e ser., iv, p. 733 (1885). 

2 'Zeitsch. f. w. Zool./ Bd. xxxii, p. 547, taf. xxxii, fig. 23 (1879). 



160 EITSYLLIDEA. 



1. Miorostllis Marenzelleri, n.s. Plate LIX, fig. 9; Plate LXX, fig. 3— foot. 

Specific Characters. — Head smoothly rounded, with a slight lateral notch, behind 
which is a short lanceolate tentacle and near it a lens — indicating at least one eye on each 
side. Body about 3 mm. long, slender, and tapered towards the posterior end. Segments 
about thirty. Proventriculus stretches over three segments. Foot has a small dorsal 
cirrus. Setigerous lobe bluntly conical. Upper bristles have a slender tapering terminal 
piece considerably shorter than in Exogone. The others have a short terminal piece with 
a simple hook at the tip. 

Habitat. — Procured by the dredge in four to five fathoms in the Sound of Harris, 
August, 1872. 

Head smoothly rounded and having a very slight lateral notch to indicate sub- 
division. Just behind the notch is a short lanceolate tentacle on each side. Near this is 
a lens, showing that in life at least two eyes are present. 

Body about 3 mm. long, slender, somewhat tapered towards the posterior end, and 
marked laterally by the feet. The tail was injured. The bristled segments appear to be 
upwards of twenty-eight. The proboscis seems to be indicated by a pale region com- 
mencing behind a line between the tentacles and extending to the proventriculus. The 
proventriculus is considerably larger than in the Exogone from Ireland, and stretches over 
three segments. From this the canal passes backward to the tail. A more opaque central 
region in front is probably the tooth in the proboscis. 

The foot (Plate LXX, fig. 3) has a small conical dorsal cirrus. The setigerous lobe 
is bluntly conical. The upper bristles have a slender tapering terminal piece, considerably 
shorter than in the Irish Exogone (p. 154). The others have a short terminal piece 
after the fashion of Syllis, with a simple hook at the tip. The end of the shaft of these 
bristles is dilated, shouldered, and bevelled. 

The condition of the single example is such that only an imperfect account of it 
is possible. 

This genus is closely allied to Claparede's Pdedophylax, 1 in which a small dorsal and 
a ventral cirrus are present. 



Group II.— EITSYLLIDEA, Malaquin. 

Syllidse provided with ventral cirri; palpi fused at the base only. Tentacles and 
cirri filiform or cylindrical, presenting superficial constrictions, but not distinctly 
articulated. Reproduction direct (epigamy). 

Malaquin separates the genera by the structure of the proboscis. Proventriculus 
normal, ventricule (stomach) little developed. The epithelial coat is very fine and 



i c 



Annel. Nap./ p. 210. 



PIONOSYLLIS PROLIFERA. 161 

surrounded by a layer of circular and radiating fibres with nuclei. The radiating muscles 

are less developed than in Syllis, but penetrate the epithelial coat. Externally are 
longitudinal fibres and the endothelial epithelium (Malaquin). 1 



Genus XLIX. — Pionosyllis, Malmgrem, 1867. 

Tentacles, and tentacular and dorsal cirri, articulated, as in Syllis. Proboscis armed 
with a single tooth anteriorly and with ten soft papillae. Compound bristles subfalcate, 
terminal piece long and bidentate. Median and posterior segments with long swimming- 
bristles besides the falcate (Malmgren). Pionosyllis is distinguished by the presence of 
the single tooth from Syllides, which has an unarmed proboscis ; from Eusyllis by the 
elongated nature of the terminal pieces of the bristles, though they are bifid in both ; and 
from Syllis by the bifid terminal pieces to the bristles throughout. 



1. Pionosyllis peolifera, Krohn, 1852. Plate XL VI, fig. 1; Plate LIX, fig. 10 — 
head, etc.; Plate LXX, figs. 4-6— feet; Plate LXXVIII, figs. 14-14 c, 15, and 16 
— bristles. 

Specific Characters. — Head hexangular or sub-rectangular with four distinct reddish 
eyes obliquely arranged on each side, the anterior pair wider apart and with or without 
lenses. Palpi large and long. Median tentacle has 25 — 30 articulations, lateral 16 — 18. 
Tentacular cirri long and moniliform. Body three-quarters of an inch or more in length, 
of a dull straw colour, regularly banded with brown in front. On each side of the 
segment-junctions a lateral brown bar with a speck in front, whilst in the middle line a 
shorter bar occurs at a greater distance from the segment-junction and nearly in a line 
with the speck. Tail with two moniliform caudal cirri. Foot with a long and tapering 
dorsal cirrus of about 27 or 28 articulations. Setigerous lobe bluntly conical with 
two terminal papillae. Spines with a peculiar and short point. Bristles of moderate 
size, translucent, with the convexity of the enlargement at the distal end of the shaft, 
spinous. Terminal piece rather long, with a nearly straight spinous edge and a minutely 
bidentate apex. Ventral cirrus fusiform, lanceolate, and does not reach the tip of the 
setigerous lobe. 

Synonyms. 

1852. Syllis prolifera, Krohn. Arch. f. Xaturg., Bd. xviii, p. 66, Taf. hi, f. 1. 
1860. „ variegata, Grube. Ibid., Bd. xxvi, p. 85, pi. iii, f. 6. 

1863. „ lussinensis, idem. Ibid., Bd. xxix, p. 46, Taf. iv, f. 9. 

1864. „ fiumensis, Ehlers. Borstenw., I, p. 224, Taf. ix, f. 1 — 9. 
„ „ Armandi, Claparede. GTlanures, p. 70, pi. v, f. 1. 

„ „ hexagonifera, idem. Ibid., p. 73, pi. v, f. 2. 

1869. Pionosyllis Malmgreni, Mcintosh. Trans. K. S. Edin., vol. xxv, p. 414, Tab. xvi, f. 10, var. 

1874. Syllis lussine?isis, Marenzeller. Sitzb. k. Akad Wien, Bd. lxix, sep. abd., p. 30, Taf. iii ; f. 1. 

1875. „ variegata, idem. Ibid., Taf. lxxii, p. 147, pi. ii, fig. 2. 

1 ' Recher. Syllid./ p. 239. 

78 



16 2 PIONOSYLLIS PROLIFERA. 

1879. Typosyllis (Syllis) prolifera, Laiigerhans. Zeitscli. f. w. Zool., Bd. xxxii, p. 530, Taf. xxxi, f. 3. 

1885. Syllis prolifera, Carus. Fauna Medit., II, p. 226. 

1886. „ {Typosyllis) prolifera, De St. Joseph. Ann. Sc. Nat., 7 e ser., torn, i, p. 147, pi. via, 

f. 9—13. 
1900. „ variegata, G-ravier. Xouv. Arch. Mus., 4 e ser., Taf. ii, p. 158, pi. ix, f. 8. 

1903. „ prolifera, Mcintosh. Mar. Invert. S. Africa, vol. iii, p. 35. 

1904. Typosyllis prolifera, Allen. Journ. M. B. A., n. s., vol. vii, p. 219. 

1905. Syllis „ (= lussinensis, fiumensis, and Armandi, Clap.), Grraeffe. Arbeit. Zool. 

Stat. Triest, xv, p. 323. 
f 1906. „ {Typosyllis) prolifera, De St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 179. 

Habitat. — Abundant in fissures and cracks of the rocks to the north of the harbour 
St. Peter Port, Guernsey ; between tide-marks at Luccomb Chine, Isle of Wight, and in 
the same region at Herm ; between tide-marks, Outer Hebrides ; seven to eight fathoms 
in the West Voe of Burra, Scalloway. 

Adriatic, Mediterranean, shores of France, Madeira, whilst Malmgren procured his 
species from Spitzbergen ; South Africa. The range thus is wide. 

Head (Plate LIX, fig. 10) of the typical shape, with long palpi. Median tentacle 
has twenty-five to thirty articulations ; lateral somewhat shorter. 

M 

Fig. 53. — Plan of dorsal markings of Pionosyllis prolifera. 

Body of a dull straw-colour, banded with brown in front. On each side of the 
segment-junction is a short, brown bar (fig. 53), in each lateral region with a speck or 
brown touch in front; whilst, in the median line, a shorter bar occurs at a greater 
distance from the segment-junction. Occasionally the general hue of the anterior region 
is greenish-olive. It is rather pale posteriorly. The markings thus approach those 
observed in other forms, such as the Pionosyllis lamelligera of De St. Joseph, a species, 
however, in which the tentacles and cirri are not articulated. In some from Shetland 
the body is roseate anteriorly, with a black speck (which remains for some time in spirit) 
at the base of each dorsal cirrus. 

The foot (Plate LXX, figs. 4—6) has a long and distinctly articulated cirrus which 
tapers clistally and has about twenty-seven articulations. The setigerous lobe is bluntly 
conical with a receding slope ventrally. The spines, which have a peculiar and short 
point, pierce the lobe at the upper angle. The bristles (Plate LXXVIII, figs. 14 and 15) 
are translucent and of moderate or small size, and have the convexity of the enlarged 
region at the end of the shaft denticulated. The terminal piece is rather long, has a 
nearly straight spinous edge, and is minutely bidentate at the tip. 

The ventral lobe is somewhat lanceolate, diminishing at base and tip. It does not 
extend so far as the end of the setigerous region. 

Reproduction. — Several examples had buds, and in one this was of larger diameter 
than the nurse-stock, and of a fine salmon-tint, apparently from the ova. It was about 
a third the length of the adult, and often contorted itself in opposition to it. It had four 



PIONOSYLLIS PROLIFEPA. 163 

reddish eyes, and its foot is shown in Plate LXX, fig. 5. Moreover, in a female bud 
with swimming-bristles, from the West Voe of Burra, Shetland, the falcate bristles had 
stouter shafts and shorter terminal pieces (Plate LXXVTII, fig. 14 6). Some of the 
female buds in Shetland were of a pale lilac hue. 

At first sight amongst the rocks this widely distributed species somewhat resembles 
S. armillaris. It is easily discriminated by the longer dorsal cirrus, the structure 
of the bristles, and the longer ventral cirrus. It is a characteristic species of the 
southern shores of Britain, and if it is right to make Pionosyllis Malmgreni a variety of 
the same form, then it is sparingly represented on the west so far north as the Outer 
Hebrides, and it may occur so far as Shetland. 

A form which was found between tide-marks in Herm differs from the ordinary 
examples in the shortness of the dorsal cirri, which have only ten articulations (Plate 
LXX, fig. 6) . Moreover, in certain varieties, as in one from Luccomb Chine, a calcareous 
region, the bifid tip of the bristle is less evident (Plate LXXVIII, fig. 16). 

Krohn (1852) first discriminated this species when describing its budding and that 
of Autolytus prolifer, and it is quite recognizable from his account. He also alludes in a 
footnote to an allied form which he termed Syllis fissipara. 

Grube (1863) described the same species from Xeresine in the Adriatic as Syllis 
lussinensis, giving characteristic figures of the anterior segments and bristles. 

The Syllis fiumensis of Ehlers (1864), though he describes and figures the terminal 
region of the bristles as simple, appears to be this species. He gives a detailed account 
of the alimentary canal with good figures, and also describes the segmental organs and 
the budding. 

The same year (1864) Claparede described and figured his Syllis Armandi as 
common at Port-Vendres, giving an account of the general structure and reproduction. 
His figure of the bristle is indifferent. 

Marenzeller (1874) found this species plentiful in the Bay of Muggia. It was of a 
brownish-violet colour, with bands anteriorly and a speck at the base of t\\e dorsal cirrus. 
The intestine shone through the tissues as a brownish-green streak, and the posterior end 
of the body was yellowish-green. Anteriorly the dorsal cirri are alternately long and 
short. His figures of the tips of the bristles are good. A bud arose from the forty-eighth 
segment, with capillary swimming-bristles. He recognized its relationship with Krohn's 
S. prolifera. 

Langerhans (1879) mentions and figures a simple bristle with a bifid tip which he 
found from the twentieth to the thirtieth segment. He also observes that the budding 
takes place in the following segments, viz. 31, 33, 35, 36, 37, 41. 44, and that the ova are 
greenish. Vorticellds are frequently parasitic on it. 

De St. Joseph (1888) observed that in the stoloniferous examples violet ova or 
sperms occurred in two or three segments in front of the bud according as that was male 
or female. He mentions a male bud as rose-orange, and points out that the Ghxtosyllis of 
Malmgren l is a closely allied bud. He also describes one which had a regenerated head. 

The variety described as Pionosyllis Malmgreni, from Lochmaddy, shows no bars in 

, l ' Annul. Polycli./ p. 162, Tab. ix, fig. 51. 



164 PIONOSYLLIS (?) DIVARICATA. 

front, and the terminal piece of the bristles is rather broad. The rapid widening below 
the tip is noteworthy (Plate LXXVIII, fig. 14 c). One example has a developing bud, 
about the fortieth segment, with two widely-separated dorsal eyes, and the foot has a tuft 
of swimming-bristles which do not yet extend beyond the jointed forms. 

This species approaches Malmgren's P. compacta, from which, however, it is easily 
differentiated by the shorter terminal piece of the compound bristles, the longer and more 
distinctly moniliform cirri (Malmgren's form having these organs indistinctly articu- 
lated), the absence of the elongated simple bristles in the non-budding animal, and the 
greater length of the palpi. 

The Syllis (Typosyllis) variegata of Grube seems to be this form. 

G-ravier (1900) describes the pharynx as reddish ochre, and as armed with a large 
tooth in front. The Syllis (Typosyllis) exilis, S. Bouveri, and S. compacta of this author 
seem to be closely allied forms, probably also falling under Malmgren's genus Pionosyllis. 

De St. Joseph (1906) describes examples from St. Raphael with longer appendages 
than the British. Thus he gives the tentacles thirty-six, and the tentacular cirri no less 
than ninety articulations. The dorsal cirri, again, show fifty segments. He also mentions 
the violet eggs in the stolon. 



2. Pionosyllis (?) divaricate, Keferstein, 1862. Plate LIX, fig. 12— head; Plate LXX, 
fig. 7— foot; Plate LXXVIII, figs. 17 a and 17 b— bristles. 

Specific Characters. — Head resembling that of Syllis, with two large, simple palpi ; 
eyes four, red, in pairs close together on each side, the anterior considerably the larger, 
besides an additional eye-speck at the base of each lateral tentacle (Keferstein) but not 
visible in the spirit preparations, a long median tentacle just in front of the eyes, and two 
on the anterior border of the snout. Body about one inch long, with a considerable number 
of segments (45 — 55), tapering a little anteriorly and still more so posteriorly. Dull 
brownish, paler in the first fourth, and from this a dull, yellow, median stripe proceeds to 
the tail, with brownish spurs at each side crossing the dorsum at the segment-junctions. 
Under surface pale brownish. Proboscis is a short cylinder with a smooth, distal edge 
and a brown interior with a prominent dorsal tooth. The foot has a long, filiform, dorsal 
cirrus, often coiled like a screw. The dorsal division has a single, stout spine which 
appears only before the swimming-bristles at maturity. The ventral division has a very 
long, setigerous region slightly bevelled and bifid at the tip, with several spines, and a 
series of slender bristles having a short dilatation at the tip of the shaft which has a few 
serrations on the convex edge. The terminal blade is flattened, narrow, diminishing little 
towards the tip, which is rather abruptly hooked, and has a slender secondary process 
beneath. Ventral cirrus somewhat lanceolate, curved, and though carried far outward 
does not reach the tip of the setigerous lobe. 

Synonyms. 

1844-5. Syllis {Syllides) longocirrata, CErsted. Kroyer's Nat. Tids., p. 408 (?). 

1862. „ divaricata, Keferstein. Zeitsch. f. w. Zool., Bd. xii, p. Ill, Taf. ix, f. 45 — 47. 



PIONOSYLLIS (?) DIVARIOATA. 165 

1863. Syllis normannica, Claparede. Beobach. u. Anat., p. 40, Taf. xiii, f. 24. 

1879. Pionosyllis divaricata, Langerhans. Zeitsch. f. w. ZooL, Bd. xxxiii, p. 545. 

1886. „ longocirrata, De St. Joseph. Ann. Sc. Nat., 7 e ser., t. 1, p. 160, pi. viii, f. 21 — 29. 

Habitat. — -Under a stone in a rock-pool at Herm, and nnder stones between tide- 
marks at St. Peter Port, Guernsey. 

St. Vaast-la-Hongue (Kef erstein) ; Madeira, at some depth (Langerhans) ; Bochardien, 
France (De St. Joseph). 

Head (Plate LIX, fig. 12) like that of Syllis, with two large palpi, which thus 
conspicuously differ from those of Gastalia fusca ; eyes red, and the pairs of each side 
are close together, besides a small pair at the base of the lateral tentacles. A median 
tentacle of considerable length occurs just in front of the eyes, and there are two on the 
anterior border of the snout. 

Body about an inch in length, and it has a considerable number of segments. It 
tapers a little anteriorly and still more so posteriorly. It is of a dull brownish or pale 
fawn hue, somewhat paler in the centre, which has a few dark brown specks. The first 
fourth of the body is paler than the succeeding (probably from the pharynx), and is of a 
pale brownish hue. From this a dull, brownish-yellow, median stripe proceeds to the tail, 
having on each side a brownish stripe which, by sending a spur outwards at every 
segment- junction, cuts the brown of the dorsum into segments. The bases of the feet 
are occasionally paler than the inner region. The under surface is pale brownish. 

The proboscis is extruded as a short, firm process with a smooth distal edge, and 
a brown interior with a prominent dorsal tooth, which, according to De St. Joseph, has a 
poison-canal at its base, the issue of the organ being preceded by a ring of ten papillae, 
each being continued posteriorly (behind the ring) by an elongated caecal process, the 
function of which is unknown. 

The foot (Plate LXX, fig. 7) has a long pale dorsal cirrus — longer than that of 
Gastalia, indeed it resembles a mobile hair which is now gently extended, and again 
drawn in numerous screw-coils close to the body of the worm. The dorsal division of the 
foot is represented in the mature female by a single stout spine which scarcely projects 
beyond the integument, such probably being the precursor of the swimming-bristles, yet 
its occurrence at first caused the specific separation of the example. The inferior lobe is 
of moderate length in the ripe form, longer in the others, and slightly bevelled at the tip. 
It has several spines and a series of slender bristles (Plate LXXVIII, figs. 17 a and 17 b), 
which have a short dilatation at the tip with a few serrations on the convex edge. The 
terminal blade is flattened, narrow, diminishing very little towards the tip, which is 
rather abruptly hooked and has a slender secondary process beneath. The terminal pieces 
of these bristles diminish in length from the upper to the lower border of the fan 
projecting from the foot. The ventral cirrus is somewhat lanceolate, curved, carried far 
outward, and does not reach the tip of the fleshy setigerous lobe. The foot in the immature 
form is apparently considerably longer than in the mature, the ova, in the example 
under examination, distending the base. 

Reproduction. — De St. Joseph observes that this species produces directly without 
alternation of generations. In the ripe forms the segmental organs from the eighteenth 



1M PIONOSYLLIS (SYLLIS) HYALIXA. 

to about the thirty- sixth segment show marked ciliation internally, with a violet colour in 
the wall of the internal trumpet (pavilion), and orange at the external aperture at the 
base of the feet. At the same time a clear spine and a group of swimming-bristles 
develop beneath the dorsal cirrus. 

Advanced ova occurred in a female from Herm in July, and indeed, the example 
appeared to be almost ripe. 

Habits. — The species is somewhat smaller than Gastalia fusca and is at once 
distinguished by its brown colour. It is active and lively, wriggles through the water 
with the usual vigour of a Hesione, and is irritable and fragile. Those obtained broke 
into fragments. 

So far as can be determined this species is that originally described by Keferstein 
from St. Vaast-la-Hougue, and subsequently by Claparede as Syllis normannica, and 
though it is true Langerhans states that it has a simple tip to the terminal piece of its 
bristles, it is more likely that this feature — often so indistinct— was misinterpreted than 
that two forms so nearly identical exist on the shores of the Channel. The additional 
distinction given by De St. Joseph, viz. the structure of the first segments, would not 
seem to invalidate the foregoing conclusion. 

The occurrence of a spine for the support of the swimming-bristles, and the structure 
of the distal piece of the compound bristles, give the form an intermediate character — 
approaching the Syliids most closely, but also leaning to the Hesionidse. It may be 
placed provisionally in this group. 

De St. Joseph (1886) found Trichodina Auerbachii attached to the tentacles and feet. 

j 

3. Pionosyllis (Syllis) htalina, Ghrube, 1863. Plate LI, fig. 2; Plate LXX, fig. 24- 
foot; Plate LXXVIII, figs. 9 and 9 a— bristles. 

Specific Characters. — Head subpentagonal ; eyes four, dark-reddish, arranged in a 
trapezoid, anterior pair larger and wider apart and in some with lenses, posterior close 
together, occasionally with a small additional pigment-speck in front on each side. Palpi 
large, somewhat triangular. Median tentacle rather longer than the lateral, and all 
distinctly articulated (about twenty segments). Buccal segment slightly narrower than 
the succeeding. Tentacular cirri about the length of the dorsal cirri, a little longer than 
the breadth of the segment. Body about 18 mm. long (Grube), 90—127 segments, 
hyaline, showing the gut. Cutis dense and iridescent, occasionally with a pinkish-yellow 
colour. Two long anal cirri, thicker than the dorsal cirri, and much longer, with a 
median papilla. Feet with moniliform dorsal cirri, the articulations ranging from twelve 
to sixteen, but some are alternately shorter ; about half the breadth of the body-segment. 
Setigerous lobe forming a short bilobed cone — the spines piercing its apex, which is 
superior— leaving the longer curve inferiorly. About ten falcigerous bristles in the 
setigerous region, the terminal piece elongate and slightly curved. The jointed bristles 
occur beneath the spines, and have shafts with a marked distal curve and a bevelled dilata- 
tion, the convex edge of which has traces of serrations. The terminal piece is somewhat 



PIONOSYLLIS (SYLLIS) HYALINA. 167 

long, and bifid at the tip, the edge below being boldly spinous. The ventral cirrus is 
somewhat lanceolate and scarcely reaches the tip of the setigerons process. 

In the posterior segments the bristles are more slender, with short terminal pieces 
(about as broad as long— Grube) . 

Synonyms. 

1843. Syllis tigrina, Rathke. Nova Acta Leop. -Carol., xx, p. 165 (?). 
1852. „ fissipara, Krohn. Arch. f. JSTaturg., Bd. xviii, p. 68. 

1863. „ hyalina, Grube. Ibid., xxix, p. 45, Taf. iv, f. 8. 

1864. „ pellucida, Elilers. Borstenw., I, p. 239, Taf. x, f. 6—11. 
„ „ simillima, Claparede. Grlanures, p. 77, pi. v, f. 4. 

1867. „ borealis, Malmgren. Annul. Polych., p. 42, pi. vi, f. 42 (?). 

1868. „ simillima, Claparede. Annel. Nap., p. 199, pi. xii, f. 5. 

1874. „ macrocola, Marenzeller. Sitzb. k. Akad. Wien, Bd. lxix, 1 Abth., sep. abd., p. 37, 

Taf. in, f. 3. 

1875. „ hyalina, idem. Ibid., Bd. lxxi, p. 22 (sep. abd.). 

1879. Typosyllis (Syllis) hyalina, Langerlians. Zeitsch. f. w. Zool., Bd. xxxii, p. 535. 

1905. Syllis hyalina ( = fissipara, Krohn, etc.), Graeffe. Arbeit. Zool. Stat. Triest, XV, p. 323. 

Habitat. — Millbay Channel, Plymouth Sound (Dr. Allen). 

Lussin Grande, Neresino, Crivizza, in the Adriatic (Grube) . Quarnero, Adriatic 
(Ehlers). Common in 20 — 30 fathoms off Madeira and also between tide -marks 
(Langerhans) . Adriatic (Marenzeller). 

Head with the structure of Syllis ; two long palpi ; eyes comparatively small ; anterior 
pair the wider apart. 

Body of about 100 segments, the anterior end being slightly tapered, and gradually 
increasing to a little behind the anterior third, this region being of a yellowish brown 
colour, and each segment being banded with brown, which anteriorly is in the form of 
a double band — a longer at the segment-junction, and a shorter immediately in front. 
The succeeding region is uniformly thicker, and has a pinkish or salmon colour, a darker 
stripe marking the segment-junctions. Pharynx (proboscis) long, with a large dorsal 
tooth and twelve soft papillas in front. The proventriculus has thirty or more rows of dots 
and is reddish. 

In this species the dorsal longitudinal muscles have a different character from those 
of Syllis annillarls, which form a continuous and massive band on each side. In 
S. hyalina they are considerably thinner, and are cut into a larger upper band and a 
smaller inferior division on each side (Fig. 54), the former stretching to a point a little 
above the origin of the dorsal cirrus, and the latter terminating at the base of the spine. 
The ventral longitudinal muscles are also somewhat thin, and show a marked differentiation 
on each side of the muscular slip from the base of the spine. 

The foot (Plate LXX, fig. 24) has a dorsal cirrus of 22 — 23 articulations ; 
setigerous region forms a short cone, with the edge below the apex split, the longer 
margin being inferior. Ventral cirrus lanceolate, and does not reach the tip of the 
setigerous region. Bristles (Plate LXXVIII, fig. 9) translucent, with curved shafts, 
dilated and bevelled ends, which show traces of serrations on the convex border. The 



168 



PIONOSYLLXS (SYLLIS) HYALINA. 



terminal pieces are rather long and boldly bifid at the tip, the edge below being spinous. 
As usual, the spinous edge with the hooks at the tip is directed upward. 

The observation of Langerhans that in the caudal segments a simple bristle occurs 
dorsally and ventrally, in addition to the ordinary jointed ones, is correct. A stout, 
translucent, simple bristle (Plate LXXVIII, fig. 9a) tapered dis tally, occupies the dorsal 
margin, and apparently another ventrally. Toward the tip of the tail the ventral 
bristles and their terminal pieces are longer and more slender. The foot in this region 
has a single, powerful, translucent spine, the sharp point of which projects beyond the 
tissues. 

Reproduction. — Ehlers states that in July at Zurkowa the eggs were bluish- violet. 
Marenzeller describes the eggs as deep rose-red, and as filling eighteen segments behind 
the fifty-second. 

In the structure of the bristles this form leans to Pionosyllis, and it may in the 
meantime be placed in that group. 

Grube considered that this species approached 8. moniliformis, and he states that the 




Fig. 54. — Transverse section of the body- wall of Pionosyllis (Trypanosyllis) hyalina, Grube ; d. m. dorsal 
longitudinal muscle ; d. v., dorsal vessel ; v. v., ventral vessel ; s. o., segmental organ. Other letters as 
before. After Malaquin. 

proboscis (Russel) reached the tenth segment (in spirit), whilst the stomach extended 
between this and the nineteenth foot. 

Claparede, in 1864, describes the same species (as 8. simillima) from Port-Vendres, 
and states that the eggs are bluish. He gives an amended figure in his later work on 
Neapolitan Annelids (1868), showing ciliated grooves on each side of the head posteriorly, 
and he points out that the tips of the spines are truncated, and the tips of the bristles 
bidentate. 

Ehlers described, also from the Adriatic, this form under the name of Syllis pellucida, 
though he pointed out its near approach to Grube's Syllis hyalina. The differences in the 
shape of the ventral cirrus and in the length of the proventriculus are not material, and 
Grube had overlooked the details of the armature of the proboscis. 

There is doubt concerning the Syllis borealis of Malmgren (1867), for while there is 



TRYPANOSYLLIS ZEBEA. 169 

a general resemblance in outline of the body and processes, lie figures the bristles with 
simple tips. Unless the artist overlooked the bifid nature of these organs, Malmgren's 
form may be different. 

This form was described by Marenzeller (1874) as a new species from the Adriatic, 
where Grube originally found it. His example had somewhat shorter cirri. He gives the 
foot two spines and five or six bristles. The tail has a rudimentary cirrus between the anal 
cirri. He saw only eleven of the anterior flat papillse in the proboscis, and the dorsal 
tooth. The cseca of the gut are not mentioned. He compares it with Syllis hyalina, 
Grube, and S. pellucida, Ehlers, but, on insufficient grounds, thought it different. He 
figures a bristle with a short tip, but shows no serrations on the convex margin of the 
dilated end of the shaft; indeed they are not readily seen. 

Langerhans (1879) cleared up the synonymy of this species. He also found it bud- 
ding in November, the female stolon having violet ova, and the swimming-bristles 
developing. In a male the sperms were roseate, and it had small (developing) swimming- 
bristles. He gives a figure of two free female buds with ova, and in both a median and 
two lateral tentacles are present. The frontal incurvation is small. He is not, however, 
quite sure of their relationship, and they differ from the Tetraglene-buds of De St. 
Joseph. 



Genus L. — Trypanosyllis, Glajoarede, 1864. 

Head with palpi well-developed as in Syllis. Eyes four, large ; tentacles large, long, 
and distinctly articulated. Body proportionately large, with large dorsal cirri, alternately 
long and short. Anal cirri two, with a median process. Ventral cirri lanceolate 
(pinniform). Proboscis with an anterior row (in extrusion) of twelve flat papillae, and 
behind a hook-like series of horny teeth. A single dorsal tooth. Foot truncate, bilobecl; 
bristles with rather large terminal pieces which are boldly bifid. Reproduction by 
stolons. 

This genus was established by Claparede (1864) for a species (identical with that 
previously described by Grube as Syllis zebra) which he found .at Port-Venclres. 



1. Trypanosyllis zebra, Grille, 1860. Plate L, figs. 9 and 10— head and tail; Plate LI, 
fig. 1; Plate LXX, fig. 8— foot; Plate LXXVIII, fig. 18— bristle. 

Specific Characters. — Head rounded trapezoidal, slightly narrower posteriorly where a 
forward incurvation and a median hollow occurs. Eyes four, somewhat large, placed 
anteriorly at the angles of a trapezium; the anterior slightly larger and wider apart than 
the posterior. The median tentacle has about thirty articulations ; lateral a little shorter. 
Palpi of moderate size, diverging anteriorly. Upper tentacular cirri very long with about 
forty articulations. Body comparatively large, attaining a length of 5 cm., and about 
1 — 2 mm. in breadth (De St. Joseph). Segments 190 — 215; slightly tapered anteriorly and 

79 



170 TRYPANOSYLLIS ZEBRA. 

more distinctly so posteriorly, where it ends in two caudal cirri with numerous articulations. 
The dorsum is somewhat rounded anteriorly, the ventral surface flattened. Anteriorly 
the dorsum is beautifully banded with madder-brown or violet-brown bars— two in each 
segment, the longer touching the bases of the opposite cirri, which are pale, and the 
shorter following closely behind. Both bands can be traced to the posterior third, 
becoming considerably paler, however, behind the middle, the shorter bar then disappearing, 
and by-ancl-by the longer also fades, leaving the tip of the tail pale yellowish— from the 
intestine. The ventral surface is dull yellow. Dorsal cirri characteristically large, and all 
are distinctly articulated and alternately long and short, the latter having about twenty 
articulations, the former about double the number. They are occasionally coloured violet 
by the corpuscles (De St. Joseph). Setigerous region blunt and bilobed with two flat, 
conical papillas below the spines. Bristles translucent, with traces of serrations on the 
convex margin of the distal end of the shaft ; terminal pieces rather shorter and broader 
than in Pionosyllis, boldly bifid at the tip and spinous on the edge. Ventral cirrus ovato- 
lanceolate ; extends a little beyond the setigerous region. 

Synonyms. 

1860. Syllis zebra, Grrube. Arch. f. JSTaturges., Bd. xxvi, p. 86, Taf. iii, f. 7. 

1861. „ „ idem. Ausflug Triest, p. 143, Taf. iii, f. 7. 

1864. Trypan osyllis Krolmii, Claparede. GUanures, p. 38, pi. vii, f. 2. 

1865. Syllis zebra, De Quatrefages. Annel., II, p. 28. 

1874. „ „ Marenzeller. Sitzb. k. Akad. Wiss. Wien, Bd. lxix, p. 40, Taf. v, f . I. 

1875. „ KroJinii, Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., torn, ii, p. 35. 
1879. „ zebra, Langerlians. Zeitsch. f. w. Zool., Bd. xxxii, p. 556, Taf. xxxii, f. 17. 

1885. „ „ Oarus. Fauna Medit., II, p. 230. 

1886. ,, Krohnii, De St. Joseph. Ann. Sc. Nat., 7 e ser., torn, i, p. 180, pi. ix, f. 51—56. 
1904. „ „ Allen. Journ. Mar. Biol. Assoc, n.s., vol. vii, p. 220. 

Habitat— Between tide-marks, Herm, 1868. Common in dredgings from Millbay 
Channel and Asia shore, as well as from Yealm, Plymouth (Dr. Allen). 

Cherso (Grube). Port Vendres (Claparede). Bay of Muggia in the Adriatic 
(Marenzeller) . Marseilles (Marion and Bobretzky) . Madeira (Langerhans) . Common in 
dredgings 10—15 metres, at St. Dinard, France (De St. Joseph). 

Head (Plate L, fig. 9) rounded trapezoidal, slightly narrower posteriorly where a 
forward incurvation at the median depression occurs. Eyes four, black, rounded or ovoid, 
rather large, placed anteriorly, the posterior pair slightly the smaller. Median tentacle 
arising between the anterior eyes, broad at the base and tapering distally, with about 
thirty articulations. Lateral tentacles somewhat shorter. Palpi of average size. Upper 
of the first pair of tentacular cirri very long (forty articulations). 

Body comparatively long (50 to 76 mm.), large, rounded dorsally and flattened 
ventrally ; segments 190 ; slightly tapered anteriorly, more distinctly so posteriorly where 
it ends in two caudal cirri with numerous articulations (Plate L, fig. 10). The dorsal 
cirri are large, and alternately long and short, the former having about forty articulations, 
the latter about half the number. 

In front it is ornamented by violet transverse bands which occur on each side of the 



TRYPANOSYLLIS ZEBRA. 171 

segment- junctions, and the dorsal cirri are violet from corpuscles. An alternation of 
longer and shorter cirri is noted by De St. Joseph, and they are present anteriorly in 
the example figured. 

The proboscis occupies the area from the sixth to the eighteenth segment, and the 
proventriculus the following eighteen (De St. Joseph). 1 The stomach has no lateral ca?ca. 
Anteriorly the proboscis has ten soft papilla?, and ten hook-like horny denticulations, 
accompanied by a single large conical tooth on the dorsal arch. 

The foot (Plate LXX, fig. 8) anteriorly has dorsally the long and large cirrus alternately 
with a shorter one after the first few segments, the longer cirri having about forty and 
the shorter about twenty articulations. As transparent objects the cirri show a median 
groove towards the base. 

The setigerous region forms a blunt or truncated bilobed process only slightly 
rounded at the inferior angle. Dorsally the upper angle is pierced by two of the three 
spines, and below them are two flat, conical papilla?, the bristles closely following — to the 
number of a dozen or thereabout — in the groove beneath. They extend to the margin 
beyond the rounded ventral angle. Each (Plate LXXVIII, fig. IS) has a translucent shaft 
with the usual dorsal convexity, a few longitudinal stria? in the middle, and a more distinct 
series of slightly oblique striations in the enlarged distal region which has traces of 
serrations on its convex margin. The terminal piece is rather shorter and broader than 
in Pionosyllis, with a few oblique striations on the blade, boldly bifid at the tip and 
spinous along the free (upper) edge. 

The ventral cirrus is ovato -lanceolate, and extends a little beyond the setigerous 
region. 

The feet in front of the tail become much less, but they preserve a similar shape. 
The bristles are more slender, and the ventral cirrus is somewhat longer and more 
tapered at the tip. The simple bifid bristle mentioned by De St. Joseph as occurring 
in this region has not been seen. 

In a form procured in August at Herm the eyes were larger, and the pigment-bars 
occurred — the one in the middle of the segment and the other at the segment- junction. 
The dorsal cirrus was somewhat shorter than in the type. 

The foot presents a long and often gracefully coiled dorsal cirrus of about twenty 
segments, best marked towards the tip. The conical setigerous region has an irregular 
outline at the tip. Superiorly the sharp tips of the two spines project, then a 
prominent papilla appears, below which is the fascicle of bristles in their furrow ; the 
translucent shafts have the usual curvature towards the tip, which is dilated and bevelled 
to an acute point. Oblique stria? occur in this region of the bristle, and its front edge 
shows a trace of serrations. The terminal piece agrees with the type. 

This beautiful species was first found by Grube (1860), and his description and 
figures are good. 

Claparede (1864) described it as a new species, and instituted for it the genus 
Tri/panosylUs. This acute and enthusiastic zoologist first found the species budding, 
the nurse-stock having seventy-two segments and the stolon twenty-two. He gives a 

1 This has not been verified. 



172 TRYPANOSYLLIS ZEBRA. 

diagnostic coloured figure of the head and anterior region, and shows the true structure 
of the pharynx in extrusion, with its anterior circle of rounded papillae and its posterior 
of chitinous, hook-like processes. He describes the feet as cylindrical and trilabiate, but 
his figures of these and of the bristles need improvement. He pointed out the caeca of 
the intestine, which he thought comparable with those of Aphroditaceans, but it dis- 
appeared in the stolons. Its blood was of a fine pale rose colour. 

Marenzeller (1864) states that the number of the papillae of the proboscis is twelve, 
and he thinks that it differs from the Tryjpano syllis Krolmi of Claparede — amongst other 
reasons because that form has only ten teeth (papillae). 

Langerhans (1879) mentions an additional pair of eye-specks on the ventral surface, 
and a median process between the anal cirri. He also describes and figures the intestinal 
caeca anteriorly. In the last six to twelve segments is a simple bristle. 

Marion and Bobretzky (1875) found at Marseilles, in February, buds of a yellowish 
and rose colour, 20 mm. long. Their dorsal cirri were more cylindrical and shorter 
than that of the nurse-stock. Two large eyes occurred on the head with two lateral 
antennae and two minute palps. Swimming-bristles were present. The nurse-stock 
regenerated its posterior segments. These buds have the form of Tetraglene. 

De St. Joseph (1886) gives a careful account of this species, mentioning that in the 
terminal segments are simple birostrate bristles. He describes ten soft papillae at the 
end of the proboscis, followed by ten corneous ones, and a single large conical tooth. 
The intestine has for some distance lateral caeca, He adds further information as to its 
reproduction by budding either by male or female stolons which have the form of 
Tetraglene, Grube, and shows that such are not to be confounded with Chxtosyllis, Mgrn., 
as Langerhans does, since they differ in the configuration of the head, which is devoid of 
tentacles. Besides this type of head in the stolons of the Syllideans, there are (1) the 
form seen in Syllis arnica, De Quatref. ; (2) the form of Mela, Johnston; (3) the form of 
Chdetosyllis, Mgrn.; (4) the form of Sacconereis ; and (5) the form of Polybostrichus. The 
male buds of Trypano syllis zebra are reddish-brown, the dorsal cirri brownish- violet, with 
14 — 16 articulations, and with two kinds of bristles, viz. compound as in the nurse-stock, 
and from the third segment a superior fascicle of swimming-bristles. Moreover the feet 
are long. The head shows a deep median cleft and carries on each side a pair of very large 
eyes, the inferior (ventral) being usually the larger, and having in addition only the dorsal 
and ventral cirri of an ordinary segment. It thus corresponds with Grrube's Tetraglene. 
The body is filled with sperms. The female stolon has also four large eyes, but the head 
is less deeply incised in front. The dorsal cirri are nearly of equal size and have sixteen 
articulations. The feet are much shorter than in the male, and the entire body is 
distended with eggs, the intestine, as in the male, being rudimentary. It also has dorsal 
swimming-bristles from the third segment. After the separation of the buds from the 
nurse-stock, regeneration of the posterior segments occurs in the latter with the 
development of caudal cirri. 

The Syllis maculosa of M. Edwards 1 does not appear to diverge much from this 
species, though the author observes that the tentacular cirri are absent, and shows the 



1 c 



Regne Anim. lllust./ pi. xv, fig. 1. 



EUSYLLIS TUBIFEX. 173 

distal piece of the bristle simple. A re-investigation may clear np doubts, especially as 
to the fusion of the palpi at the base. 

The Trypanosyllis Bichardi of Gravier 1 (1900) from the Red Sea comes very near 
this form, the chief distinction being the greater length of the dorsal cirri. The variation 
in this respect may be greater than is at present supposed. 

The small size of the head of the bud in Trypanosyllis and its absence in Syllis 
sjpongicola inclines Calvin Mensch 2 to Huxley's view that the stolon amongst Syllids is not 
so distinctly individualized as would appear from the earlier observations of Alex. 
Agassiz on Autolytus comutus. 



Genus LI. — Eusyllis, Malmgren, 1867. 

Head with a ciliated collar; palpi soldered at the base; general structure as in 
Syllis. Proboscis in extrusion armed with a single conical tooth, with a double row of 
papillas (circa ten) incomplete on the dorsal margin. Compound bristles falcate ; terminal 
pieces short and bifid. Dorsal cirri short and indistinctly articulated. Ventral cirrus 
broadly lanceolate. Median and posterior segments with or without long capillary 
(swimming) bristles. Two anal cirri. 

Reprodmtim.— Epigamy occurs in this genus (Malaquin), and the genital glands are 
developed immediately behind the proventriculus, viz. from the twelfth to the fifteenth 
segment. The development of the swimming-bristles takes place as in schizogamy. 



1. Eusyllis tubifex, Gosse (1855). Plate LIX, figs. 11 and 13— head; Plate LXX, fig. 9 

—foot; Plate LXXIX, ^g. 1— bristle. 

Specific Characters.— Read somewhat ovoid, with a pair of comparatively small red 
eyes on each side, the anterior and larger being the wider apart. Palpi of considerable 
length, ovoid, soldered at the base, richly ciliated internally, and with palpocils in front. 
Median tentacle longer than the lateral. The base of these organs is comparatively 
smooth, whilst the distal region is moniliform. Body of about fifty-six segments, and five 
eighths of an inch in length. Pale and translucent anteriorly, the proboscis having a bluish- 
white iridescence and a brown speck or two. Proventriculus is opaque-white, and dotted. 3 
Behind the latter is a pale region of the body, then the intestine gives it a pale- 
brown hue, which increases in depth towards the tail which has its tip pale. A series of 
white grains is grouped on the dorsum from the proventriculus backward. The tail has 
a median lanceolate process and two long moniliform cirri with palpocils. Cirri of the 
first four segments longest and most distinctly moniliform, the latter character diminishing 
posteriorly. Proboscis has a denticulated edge and a single large pyramidal tooth. Foot 

1 'Xouv. Arch. Mus./ 4 e ser., t. ii, p. 168, pi. ix, figs. 12 and 13. 

2 ' Amer. Naturalist/ vol. xxxiv, p. 165 (1900). 

3 Haswell, Eisig, and De St. Joseph demur to these being termed glands. 



174 EUSYLLIS TUBIFEX. 

has dorsally the tapering cirrus, the distal crenations being better marked in the anterior 
than in the posterior feet. Setigerous region conical, with a fan-shaped series of bristles, 
the curves of which increase from the dorsal to the ventral edge. The terminal piece is 
comparatively short and bifid, the hooks being directed upward. A well-developed tuft 
of cilia occurs between the dorsal cirrus and the setigerous lobe. Ventral cirrus is ovate, 
with a bluntly conical tip which does not reach the end of the setigerous region. 

Synonyms. 

1855. Syllis tubifex, G-osse. Ann. Nat. Hist., ser. 2, vol. xvi, p. 31. 

1865. „ „ De Quatrefages. Annel., II, p. 24. 

1869. Eusyllis „ Mcintosh. Trans. R. S. Edin., xxv, p. 414, pi. xv, f. 21. 

1874. , ; „ idem. Ann. Nat. Hist., ser. 4, vol. xiv, p. 198. 

1875. „ „ Mcintosh. Invert, and Fishes St. Andrews, p. 121. 
1891. „ } , Hornel. Trans. Liverp. Biol. Soc, vol. v, p. 244. 
1901. „ „ Whiteaves. Rep. Geol. Surv. Canada, No. 722, p. 81. 

Habitat. — In vast numbers in membranous tubes on the blades of Laminaria saccharina 
tossed on shore by storms. It is thus common both in the laminarian region and in deep 
water amongst alga3 and zoophytes. When the debris is placed in sea-water the annelids 
congregate at the water-line of the vessel. Stomach of the haddock, St. Andrews (E. M.). 

Le Havre Bank, Nova Scotia ('Challenger'). Madeira (Langerhans). Canada (W.C.M.). 

Head (Plate LIX, fig. 13) somewhat ovoid, with a pair of comparatively small 
red eyes on each side in the usual oblique position, the anterior being slightly larger and 
wider apart than the posterior. Palpi of considerable length, ovoid, joined at the base in 
front of the snout, and richly ciliated internally, besides having in front some motionless 
microscopic palpocils. The pair in outline resemble the sole of a ruminant's foot. They 
are capable of various movements of extension and contraction. The median tentacle is 
longer than the two lateral. One palpocil in each segment of the tentacle is generally 
prominent, though sometimes there are two. The three tentacles are often gracefully 
coiled, and when the animals are vigorous the base is comparatively smooth, while crena- 
tions and a moniliform condition occur distally. 

Body of about fifty-six segments, and about five eighths of an inch in length. It is pale 
and translucent anteriorly, the pharynx having a bluish- white iridescence and a brown speck 
or two. The proventriculus is opaque-whitish and dotted. Behind the latter is a pale 
portion, then the intestinal region is pale brownish, increasing in depth towards the tail, 
which, however, occasionally has its tip pale. A series of opaque-white grains is grouped 
on the dorsum from the proventriculus to the tip of the tail. A faint, reddish streak 
occurs just behind the former. All the cirri are pale. The body is little tapered anteriorly, 
but diminishes rapidly posteriorly where it terminates in two long moniliform cirri with 
palpocils. A short and somewhat lanceolate process marks the end of the body between 
the cirri. The cirri of the first four segments are longest and most distinctly moniliform. 
The sixth is shorter and is generally carried horizontally outward, and the rest are about 
the same length, but with less evident crenations towards the tail, where they become a 
little longer and more slender. All have palpocils — most developed towards the tip. The 



EUSYLLIS TUBIFEX. 175 

ceratophore, or basal process, is ciliated, and the large and powerful cilia extend to the 
setigerous region. 

The proboscis (Plate LIX, fig. 11), which extends to the sixth segment, has a 
denticulated edge, though more than a third of the circumference is only minutely 
crenated; and it is furnished with a small pyramidal tooth situated on the smoother part 
of the edge. The sides of the organ are thrown into prominent wrinkles. Several elon- 
gated papillse occur in front of the anterior edge of the proboscis — some directed forward, 
others backward (the soft ring under compression). The proventriculus is studded with 
minutely granular dots, and stretches to the thirteenth segment. This organ would seem 
to act as a compressor, for its two halves occasionally move against each other. The 
glandular intestine is rendered moniliform by the septa, and terminates posteriorly in a 
ciliated rectum. Occasionally ciliary action is observed just behind the proventriculus 
and apparently extending into the lateral casca as well as along the gut. 

The foot (Plate LXX, fig. 9) presents dorsally the tapering cirrus which has more 
distinct crenations anteriorly than posteriorly. After an interval is the somewhat conical 
setigerous region, with the tips of the spines pointing at the upper angle. The bristles 
(Plate LXXIX, fig. 1) form a fan-shaped series, with the convexity of the curve 
directed dorsally, and the curvature increasing from the dorsal to the ventral edge of the 
series. The bevelled distal end of the shaft is minutely spinous. The terminal piece is 
comparatively short and bifid, with the hooks directed upward so that they would be useful 
to the annelid in clinging to its tube or other surface. There is little difference between 
the anterior and the posterior bristles. The ventral edge of the fan of bristles has more 
slender bristles, but the terminal pieces are longer than the superior. The ventral cirrus 
is ovate with a bluntly-conical tip which does not extend so far outward as the setigerous 
region. In the posterior feet this lobe is smaller and more pointed. A well-developed 
tuft of long cilia occurs between the dorsal cirrus and the setigerous lobe, and another on 
the dorsal edge of the latter. 

Reproduction. — This species perhaps sometimes develops directly, since comparatively 
large ova occur in the ordinary forms, yet in these, between the dorsal cirrus and the 
setigerous lobe of the foot, indications of a spine are observed, though none of the natatory 
bristles are developed. 

Ovigerous buds with long natatory bristles supported by a spine also occur, and the 
ordinary bristles are somewhat longer both in shaft and terminal piece. Such buds are 
found on the blades of the tangles in April. 

Habits. — They rapidly secrete a translucent tube in confinement, and thus protect 
themselves. If incautiously interfered with their bodies are ruptured, and the anterior 
end wriggles out of the tube. 

On being irritated with the forceps, the annelid emits a fine green light from the 
ventral surface of each foot and as if from many minute pores at the area. It flashes 
along both sides posterior to the point of irritation, and fades, a faint trace remaining for 
a few seconds. On severe irritation it remains luminous behind the injured point for 
nearly half a minute, and the surface of granular light on each segment is larger than 
usual, and in a few instances is connected across the ventral surface by a few granular 
phosphorescent points. For some time it was sufficient to shake the vessel to cause a 



!?6 BUSYLLIS BLOMSTRANDI. 

repetition of the brilliant green flashes. Even when undisturbed a point of green light 
appeared here and there. After immersion in spirit a moniliform band of green 
phosphorescence occurred on each side and continued for some time, the tail being 
brightest. For five minutes the body was faintly luminous, and yet hernias of the tissues 
had occurred at the injured points. 

This species was first clearly described by Mr. Grosse (1855), who found it not 
uncommon in his jars containing Hydroids, from Ilfracombe. It is probably, however, 
the Nereis cirrhigera of Viviani. 1 

The Eusyllis pliosphorea of Verrill 2 comes very near this species. 

De St. Joseph thinks that all the representatives of Eusyllis reproduce without 
alternation of generations. He does not think that the Syllis tubifex of Gosse — which is a 
stolon of a Eusyllis — is to be confounded with my Eusyllis tubifex — which apparently he 
had not found at Dinar d. 



2. Eusyllis blomsteandi, Malmgren, 1867. 

Specific Character s.— Head subrectangular with the angles rounded. Eyes four, 
forming a rectangle (with occasionally two oculiform points in front). Palpi somewhat 
ovate. Median tentacle longer than the lateral, and the dorsal of the first pair of 
tentacular cirri exceeding the length of the tentacle. All are indistinctly articulated, and 
the tips of these and of the first dorsal cirri are coloured brownish. Malmgren states that 
the long tentacular cirri are twice or thrice the diameter of the body. Body five eighths 
of an inch long, of an orange colour and having the typical shape. The dorsal cirri 
become shorter than the breadth of the body after the fourth. Two caudal cirri occur 
at the tapered posterior extremity. Proboscis with a single tooth at the tip in extrusion, 
and a series of papilla behind it. Proventriculus with fifty-five rows of points, and the 
lateral pouches of the ventricle are small. Foot has dorsally the somewhat short, 
unjointed cirrus, then a bluntly-conical setigerous region. Bristles have slightly curved 
shafts with serrations on the bevelled region at the tip, and on the convex edge of the 
dilatation. Terminal piece of moderate length and nearly equally bifid at the tip. 
Ventral cirrus is somewhat ovate, and does not quite reach the tip of the setigerous 
region. 

Synonyms. 
1867. Eusyllis Blomstrandi, Malmgren. Annul. Polych., p. 40, Tab. vi, f. 43. 
1869 - » „ Mcintosh. Trans. P. S. Edin., vol. xxv, p. 415. 

1885. „ „ Cams. Fauna Medit., p. 229. 

18 87. „ „ De St. Joseph. Ann. Sc. Nat., 7 e ser., p. 171, pi. viii, f. 39. 

!892. „ „ Marenzeller. Polychast. Ostspitzberg, p. 411, Taf. 19, f. 3. 

Habitat— Dredged in the Minch, August, 1865. Plymouth? (Allen). 
Mediterranean (Cams), Madeira (Langerhans), Dinarcl, France (De St. Joseph), 
Spitzbergen (Marenzeller). 

1 ' Phosphorescentia Maris/ p. 9, Tab. iii, figs. 1 and 2 (1805). 

2 ' Trans. Conn. Acad./ iii, p. 39, pi. vii, fig. 2. 



EUSYLLIS BLOMSTRANDL 177 

Head sub-rectangular, with the angles rounded. Eyes four, forming a rectangle, and 
according to De St. Joseph, with a pair of oculiform touches in front. Palpi somewhat 
ovate. Median tentacle longer than the lateral, indistinctly articulated. Tentacular 
cirri of similar shape, the dorsal of the first pair exceeding the length of the tentacle. 
At the base of the head is a patch of cilia on each side. The tips of the tentacles, the 
tentacula, and the first dorsal cirri, are coloured brown (De St. Joseph). 

Body of an orange hue, about five eighths of an inch long, but incomplete in the 
examples, and having the typical shape. The dorsal cirri become shorter than the 
breadth of the body after the fourth. Two caudal cirri at the tapering posterior end. 
Proboscis with a single tooth at the tip in extrusion, and a series of papillse behind it. 
The proventriculus has fifty-five rows of points, and the lateral pouches of the ventricle 
are small. 

The foot has dorsally the short cirrus, then a bluntly-conical, setigerous region, with 
the tips of the spines l pointing to the upper angle. The bristles (Plate LXXIX, fig. 2) 
have slightly curved shafts, with traces of serrations on the bevelled region at the tip 
and the convex edge of the dilatation. The terminal piece is of moderate length and 
nearly equally bifid at the tip. Slight variations in the length of the terminal piece occur 
in different specimens from the Minch. The ventral lobe is somewhat ovate and does not 
quite reach the tip of the setigerous region. 

De St. Joseph 2 (1887) found an abnormal example with three proventriculi. The 
median was normal in position, whilst the two lateral were posterior to the former and in 
the situation of the lateral cseca, which in this case were minute. If no misinterpretation 
occurred the condition is remarkable. 

An elaborate account of this form is given by Marenzeller (1894) from examples 
procured in the Ostspitzbergen Expedition of Kukenthal and Walther in 1889. He 
figures in some of these the enlarged distal end of the shaft spinous along the bevelled 
edge, and in others hispid with small spikes over the dilated region. 

A fragmentary specimen from the South- West of Ireland (Royal Irish Academy's 
Expedition, 1885) appears to correspond with the foregoing species. The dilated and 
bevelled ends of the shafts of the bristles, however, show the minute spines less distinctly 
either from their removal by friction or otherwise. This also is a female laden with ova, 
and swimming-bristles are developing below the dorsal cirrus. 

Eusyllis monilicomis of Malmgren is a very closely allied form, and further examina- 
tion may explain certain points of divergence, as, for instance, in the description of 
De St. Joseph, for it differs from the type in Malmgren's paper. 

Malaquin, though he did not ascertain the precise details of the early history of the 
eggs and larvse of Eusyllis monilicomis, found young forms abundant on the tufts of 
Polyzoa (Membranijoora), showing no movements, devoid of cilia, and enveloped in a 
vitelline membrane. Each presents two eyes, and a larval pharynx abutting on a 
vitelline mass differentating into an intestine. These are of interest in relation to 

1 De St. Joseph gives these a curved tip (as in Sylline), but such was not seen in the 
foregoing examples. 

2 De St. Joseph, op. cit., p. 172, pi. viii, fig. 39. 

80 



178 ODONTOSYLLIS FULGURANS. 

Eusyllis Blomstrandi from the close connection of the forms. As development proceeds, 
the tentacle and additional pair of eyes posteriorly, the appearance of the adult pharynx 
behind the larval and the gradual recrudescence of the latter and the completion of the 
former, the growth of the proventriculus behind and in rear of it, the lateral cseca and 
the intestine with its anus, the growth of the palpi subsequent to the tentacles, as lateral 
buds, 1 the fusion of the base as they increase in size and pass to the front with their rich 
coating of cilia and palpocils, and the completion of the alimentary canal, are all figured 
and described by the author. There is much in common with the growth of Autolytus, 
but the cephalic band of cilia does not appear till there are seven or eight bristled seg- 
ments, at which time the dorsal ciliated bands also occur posteriorly. 

The author points out that he thus differs from Viguier, who described the develop- 
ment of the proboscis as from behind forward, and who considered the larval pharynx as 
the proventriculus. 



Genus LIL — Odontosyllis, Claparede, 1863. 

Palpi short or moderately elongated, more or less separate, or fused at the base. 
Tentacles (three) and dorsal cirri filamentous, short. They become longer in the mature 
(sexual) state. Nuchal organ has a central pit (Malaquin), and an occipital gibbosity occurs. 
Tentacular cirri two pairs. Ventral cirrus present. Proboscis with a series of pointed 
cuticular horny papillae, the points curved backward. 2 Ventricle (stomach) short and 
devoid of T-shaped caaca. Bristles with terminal piece simple or bifid (Claparede 3 ). 



1. Odontosyllis fulgurans, And. and JEW., 1833. Plate XLIX, fig. 5; Plate LIX, figs. 
15 and 15 a and b — head and tail ; Plate LXX, fig. 11— foot ; Plate LXXIX, fig. 4— 
bristle. 

Specific Characters. — Head rounded, a Y-shaped groove dividing it into three regions. 
Eyes four, large, the anterior with lenses, occasionally nearly forming a square. Palpi 
moderately elongated. Median tentacle, which arises in front of the eyes, longer than the 
two lateral. Body fully an inch in length with 60 to 100 segments, slightly tapered 
anteriorly and more distinctly so posteriorly. A prominent papilla in the mid-dorsal line 
occurs behind the head, giving gibbosity to the region. Tail with two median cirri and a 
slight median process between them. Of an orange hue, or pale yellow in front and reddish 
over the intestinal region behind. Whitish grains on the dorsum. Proboscis short, 
with seven denticulations. Proventriculus long (extending to the sixteenth segment). 
Foot distinctly bifid at the tip. Dorsal cirrus of moderate length, unjointed. The 
dorsal edge of the setigerous lobe passes with a slight curvature to the spine, a deep 
groove splitting the foot beneath. The dilated distal region of the shafts of the larger 

1 Malaquin compares these with the condition in Nerilla, Schmidt, op. cit., p. 412. 

3 Ehlers observes that the opening of the oesophagus has two transverse thickenings like teeth. 

3 Claparede mentions a bifid bristle (like that of Syllis gracilis) in a large specimen from St. Vaast. 



ODONTOSYLLIS FULGITrlANS. 



179 



bristles is smooth. The terminal process is comparatively short, ends in a strong hook, 
and after an interval has a smaller process about the middle of the curve. Ventral cirrus 
ovate, its bluntly- conical tip being less prominent than the setigerous region. 



Synonyms. 

1756. Noctiluca marina, Adler. Amoenitates Acad., p. 202 (with fig. in text). 

1761. Nereis noctiluca, Linnaeus. Fauna Suecica, II Edit., pp. 509 and 2098. 
1767. „ „ idem. Syst. Nat., ed. 12, i, pt. 2, p. 1085. 

1787. „ „ Pennant. Brit. Zool., IY, p. 46; and edit. 1812, IV, p. 93. 

1791. „ „ G-melin. Linn. Syst, Nat., ed. 13, i, pt. vi, p. 3115. 

1806. „ „ Turton. Linn. Syst. Nat., IV, p. 86. 

1807. „ „ idem. Brit. Fauna, p. 134. 
1817. „ „ Stewart. Elem., I, p. 390. 

1833. Syllis fulgurans, Audouin and Edwards. Ann. Sc. Nat., t. xxix, p. 229. 

1834. Nereis noctiluca, idem. Annel., p. 209. 
„ Syllis fulgurans, idem. Ibid., p. 207. 

1851. „ „ G-ruhe. Fam. Annel., p. 62. 

1853. Nereis noctiluca, Daly ell. Pow. Creat., ii, p. 161. 

1864. Odontosyllis fulgurans, Claparede. Grlanures, etc., p. 95, pi. viii, f. 1. 

1865. ,, „ De Quatrefages. Annel., ii, pp. 24 and 648. 

1874. „ virescens, Marenzeller. Adriat. Annel., April, 1874, p. 41, Taf. iv, f. 2. 

1875. „ fulgurans, Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., t. ii, p. 40, pi. iv, 

f. 11, 11a. 
1879. „ „ Langerhans. Zeitsch. f. w. Zool., xxxii, p. 554. 

„ ,, „ Webster. Annel. Yirg. Coast, Trans. Alb. Instit., ix, p. 220 (sep. 

copy, 20). 
idem. 32nd Eep. N. Y. Mus. Nat. Hist., p. 109. 
1885. „ „ Cams. Fauna Medit., p. 231. 

1887. „ „ De St. Joseph. Ann. Sc. Nat., 7 e ser., t. i, p. 175. 

1890. „ „ Giard. Bull. Sc. Fr. Belg., t. xxii, p. 79. 

„ „ „ Malaquin. Annel. Boulon, p. 36. 

1904. „ „ Allen. Journ. M. B. A., n.s., vol. vii, p. 219. 



Habitat. — Under an Ascidian fixed to a littoral stone between tide-marks, Herm. 
It was observed in the dnsk, when tearing off the Ascidian, by its bright green phos- 
phorescence. Dredged by Dr. Gwyn Jeffreys in St. Magnus Bay, Shetland, in 100 
fathoms in 1867. Dredged at Plymouth (Dr. Allen). 

Mediterranean (including the Adriatic) and the Atlantic. Amidst tubes of Dasychone, 
Marseilles (Marion and Bobretzky). Common at Dinard (De St. Joseph). Virginian 
Coast, dredged in 6 — 12 fathoms on shells and stones; pelagic male on surface, New 
Jersey (Webster). 

The specimen is unfortunately imperfect, so that only a brief account can be given. 
In the general outline of the fragment the body resembles Autolytus, the dorsal cirri being 
smooth, and they are slightly longer than in A. prolifer, from which they differ also in 
the absence of the distinct basal joint (ceratophore). The basal region is somewhat 
constricted, and it dilates a little beyond this and again slightly tapers towards the tip. 



180 ODONTOSYLLIS FULGURANS. 

Moreover, on its convex edge it presents several wart-like papillae, but whether these are 
due to imperfect preservation is uncertain. 

Head (Plate LIX, fig. 15) rounded, provided with two moderately elongated palpi, 
which in the Zetlandic specimen seemed to be short, and three tentacles, the median, 
which arises in front of the eyes, being the longest. Eyes four, large, purplish, the 
anterior with lenses. In the Zetlandic example from deep water they nearly formed a 
square. A Y-shaped groove divides the head into three regions. 

Body fully an inch in length with upwards of sixty segments, slightly tapered 
anteriorly and more so posteriorly. A prominent papilla behind the head in the mid- 
dorsal line. Of an orange hue throughout. De St. Joseph gives it 100 segments and states 
that it is pale yellow in front and behind, reddish over the intestinal region, and with 
whitish grains arranged along the dorsum. 

Proboscis with seven denticulations ; proventriculus long (to the' sixteenth segment). 

Posteriorly the body terminates in two caudal cirri with a slight median process 

between them (Plate LIX, fig. 16 a). When the tail is injured the two cirri in front of 

the truncated extremity bend backward and become somewhat longer than the others 

(Plate LIX, fig. 156). 

The feet are more prominent than in Autolytus and are distinctly bifid at the tip 
(Plate LXX, iig. 11). The dorsal edge 1 passes outward with a slight curvature to the 
two spines which just pierce the upper and outer angle, a deep groove occurring 
beneath so as to split the tip vertically. The greater part of this lobe is occupied by the 
strong, translucent bristles which have a slight curvature below the dilatation at the tip 
of the shaft (Plate LXXIX, fig. 4), which is bevelled as in allied forms, and ends in a 
point. The edge appears to be almost smooth. The terminal process is articulated as in 
Autolytus, and shows a strong hook, and after a distinct interval, so as to be about the 
middle of the curve, a smaller process. The bristles are proportionally larger than in 
Autolytus, and their structure is diagnostic. 

The inferior lobe of the foot is more or less ovate, its bluntly-conical tip being less 
prominent than the upper lobe. 

Claparede found this species at Port-Vendres, Pyrenees Orientales, and describes the 
beautiful emerald lustre of its phosphorescence in daylight. A band of green shone along 
each side of the body, and by-and-by broke up into points grouped at the bases of 
the feet. 

Marenzeller's 0. virescens from the Adriatic, as De St. Joseph states, approaches this 
species very closely, indeed, the structure of the head, the foot, pharynx, and other parts, 
is almost identical, but he did not observe the second tooth on the terminal piece of the 
bristles or notice the phosphorescence. 

De St. Joseph points out that the pharynx (proboscis) has ten teeth and ninety rows 
of points in the proventriculus, and that the terminal pieces of the bristles are bifid, and 
arranged as his excellent figure shows. He observed a transparent gland in the feet of 
the middle region of the body, the function of which is undetermined. He found that it 
developed directly, and that the eggs were violet. Moreover, after a precursor of a 

1 In one preparation what seemed to be a band of cilia coursed along the upper border of the 
foot. 



ODONTOSYLLIS FULGURANS. 181 

small, curved, dorsal bristle on the twenty-seventh, the long natatory bristles 
extended from the twenty-eighth to the fifty-second segment. In another example the 
sexnal elements and the capillary bristles occurred from the twenty-fifth to the fortieth 
segment. He found gregarines in the intestine of one ; whilst in another the head was 
regenerated, but without proboscis or proventriculus. Its phosphorescence is so bright as 
to show the parts under the microscope. 

The specimen from deep water in St. Magnus Bay showed the structure of the 
bristles very clearly, and the terminal hook of the distal piece was perhaps a little less 
curved than in the form from the Channel Islands. 

It would be difficult to frame a theory explanatory of the phosphorescence of this 
annelid under a littoral ascidian in the Channel Islands, and at the depth of 100 fathoms 
in the Zetlandic seas, just as it was long ago pointed out how uncertain the theory is 
which endows, on the one hand, phosphorescent animals with the faculty of alluring others 
which fall a prey to them, and, on the other, of attracting the attention of some to their 
own destruction. 

This species has been a favourite subject of study with observers from De la Voie 
(1666) onwards, the memoirs of Vianelli, Claparede, and Panceri on its phosphorescence 
being especially noteworthy. The last-named author locates the luminosity in certain 
glands at the base of and in the dorsal cirri, but other tissues may not be excluded under 
the influence of the nervous system. The figures of this author are of much interest. 
Viviani probably refers to this form under his Nereis radiata. 1 Adler, again (1756), gives 
his Noctiluca marina a round head, two eyes, two tentacles (figure shows three), and a 
body of twenty segments. It is two lines long, and is found amongst the marine algas. 

The Nereis noctiluca, of 0. F. Miiller 2 (1806) is apparently a female bud of a form 
approaching this and resembling the coloured figure of a male Autolytus. In the figure 
of the segments the foot is inverted. He adds " Corpus totum purpurascens." 

It is possible that the Syllis fulgurans of Audouin and Edwards, which was sent to 
them by Duges from the shores of the Mediterranean, is this species. 

A very closely allied form is the Odonto syllis dugesiana of Claparede, 3 who described 
it as having nine teeth in its proboscis, but Langerhans 4 has given it only six (as in 
0. fulgurans), and the compound bristles are similar. 

Claparede 5 shows a young Odonto syllis of three bristled segments and three eyes on 
each side. The bristles are very long and have tapering terminal pieces. 

Webster found a large adult male with swimming-bristles commencing on the 
twenty-first segment and extending over forty-two, in Great Egg Harbour, New Jersey. 

Malaquin 6 describes a young Odontosyllis in the dipharyngeal stage with five bristled 
segments and lateral ciliated pits. 

1 ' Phosphor. Maris/ p. 11, Tab. iii, figs. 5 and 6. 
3 ' Zool. Dan./ iv, p. 31, Tab. cxlviii, figs. 1—3. 

3 ' Grlanures/ p. 97, pi. viii, fig. 2. 

4 ' Zeifcsch. f. w. Zool./ Bd. xxxii, p. 554, Taf. xxxii, f. 15. 

5 ' Beobacht./ p. 81, Taf. xii, fig. 15, 1863. 

6 Op. cit., p. 429, pi. xiv, fig. 29. 



182 ODONTOSYLLIS CTBNOSTOMA. 



2. Odontosyllis ctenostoma, Oldparede, 1868. Plate LXX, fig. 12— foot; Plate 

LXX1X, figs. 5 and 6 — spine and bristle. 

Specific Characters. — Head rounded, with short palpi in front, and ventrally pro- 
jecting separately in the preparations. Eyes four, large, the anterior with lenses and 
distinctly separated from each other, nearly forming a square. Median tentacle slightly 
longer than the lateral. None of the appendages articulated. Body like that of 
0. fiulgurans, and about an inch in length. The segments are banded transversely with 
greyish granulations which also occur on the gibbosity, and the general colour is green, 
like that of Bryopsis, amongst which it occurs (Marion and Bobretzky). Six teeth occur 
in the proboscis anteriorly. The proventriculus commences at the eighth bristled 
segment. The foot forms a short cone slightly bifid at the tip. Dorsally is the 
moderately long cirrus, which is slightly tapered toward the tip. The setigerous region 
has two pale spines with a beak and crown (spike) at the tip. The paler bristles issue 
between the blunt papilla at the tip, are curved distally, dilated at the end of the shaft, 
and the convex edge and tip are finely serrated. Terminal piece short, with a simple 
boldly-curved hook at the tip. The front edge is concave and finely spinous. In 
Claparede's figure of the compound bristle a tooth occurs on the anterior margin of the 
terminal piece near its base, as in Odontosyllis fiulgurans. Langerhans, however, shows 
none, and points out that the proboscis (his pharynx) is longer than in the latter species. 
Ventral cirrus short and broadly lanceolate, the tip falling short of that of the setigerous 
region. 

Synonyms. 

1868. Odontosyllis ctenostoma, Claparede. Aimel. Nap., p. 202, pi. xii, f. 4. 

viresceus, Marenzeller. Sitzb. k. Akad Wiss. Wien, Bd. lxix, p. 41, Taf. iv, f. 2. 

Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., torn, ii, p. 42, pi. iv, f. 12. 

Langerhans. Zeitsch. f. w. Zool., Bd. xxxii, p. 555, Taf. xxxii ; f. 16. 

Cams. Fauna Medit., ii, p. 230. 

Allen. Jonrn. M. B. A., n.s., vol. vii, p. 219. 

Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 323. 

Be St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 185. 

Habitat. — Plymouth, between tide-marks, under stones, and amongst sea- weeds and 
sponges (Allen). 

Naples. Marseilles (De St. Joseph, Marion and Bebretzky). Adriatic (Marenzeller). 
Common at Cannes. 

Head, like that of 0. fidgurans, rounded in front, and with four distinct, brownish eyes 
nearly in a square, the anterior pair having lenses. They are distinctly separated from 
each other, yet Claparede states that they are almost coalescent. The palpi form short 
projections — best seen from the ventral surface. Tentacles and tentacular cirri similar to 
those of the species mentioned. 

Body like that of 0. fiulgurans, and ranging from 18 mm. to 25 mm. or even to 1 cm. 



1874. 


„ vires 


1875. 


1? > 


1879. 


;> > 


1885. 


)> } 


1904. 


J5 } 


1905. 


)) ) 


1906. 


T) } 



ODOXTOSYLLIS G-IBBA. 183 

(Claparede), slightly diminished anteriorly, and more distinctly tapering posteriorly. It 
is bright green at Dinard, with a transverse band of greyish granulation in each segment 
(De St. Joseph). Yellowish-green (Marenzeller). Body terminated by two cirri. 
Marenzeller and Marion and Bobretzky observe that the armed region of the proboscis is 
found in the fifth setigerous segment, and it has six teeth under a glandular mass of the 
fibrous region of the organ. The proventiculus occurs in the eighth setigerous segment, 
and it is twice as long as the former. 

The foot (Plate LXX, fig. 12) is comparatively short and bluntly bifid at the tip, the 
bristles issuing between the papillae. Two pale spines pierce the tip superiorly and their 
ends are peculiarly bent so as to form a short, blunt beak anteriorly, whilst a kind of heel 
projects posteriorly in some, as apparently first indicated by Marenzeller. The tip is 
indeed somewhat T-shaped, especially in the posterior segments (Plate LXXIX, fig. 5). 
The bristles are translucent, with the shafts slightly curved, dilated, and bevelled at the 
tip, which is finely spinous. The terminal piece (Plate LXXIX, fig. 6) is short, with a 
bold and sharp hook at the tip, and a concave, spinous edge in front. 

The ventral cirrus is broadly lanceolate, and does not reach the tip of the setigerous 
region. 

Reproduction. — An example captured on April 23rd at Plymouth is laden with 
nearly ripe sperm-cells. The segmental organs are green (Fage 1 ). 

The habits of this form would seem to resemble those of O.fulgurans, and Marion 
and Bobretzky found it brilliantly phosphorescent. 



3. Odontosyllis gibba, Claparede, 1863, var. Robertian^. Plate XLIX, figs. 3 and 4; 
Plate LIX, fig. 14 and Ua— head; Plate LXX, fig. 13— foot; Plate LXXIX, 
fig. 7 — bristle. 

Specific Characters. — Head small, rounded or bluntly conical, with two reddish eyes, 
confluent or separated, on each side, the anterior the wider apart. Two short anterior 
tentacles and a median with a dark tip in front of the eyes. Tentacular cirri dark. Body 
about half an inch long, most distinctly tapered posteriorly. Segments 43 — 48. Of a 
pale brown or fawn colour, with a whitish stripe interrupted by three or four brownish 
touches along the dorsum, and a transverse white belt extending to the bases of the feet 
on the seventh segment. Ventral surface of a pale brown hue with dark touches in front. 
Gribbous segment succeeding the head is somewhat semicircular, and bears two clavate 
cirri each with a white ring on the dilated region. Terminal segment has two short and 
somewhat clavate cirri with a white touch at the tip. Foot uniramous in so far as there 
is only one spine. Dorsal region elevated, giving rise to a clavate cirrus, dilated towards 
the tip, which has a white ring, and with a slender terminal process. The cirrus is simply 
clavate or fusiform in other examples. Setigerous region separated by a sulcus, its dorsal 
margin convex, and terminating externally in two conical papillse. -Bristles long, slender, 
and translucent, dilated distally, and with a shoulder and a bevelled edge. Terminal 
process short, very slightly diminished at the tip, which has a simple hook. 

1 ' Ann. Sc. Nat./ 9 e ser., iii, p. 316. 



184 ODONTOSYLLIS GIBBA. 

Synonyms. 

1863. Odontosyllis gibba, Claparede. Beobacht., p. 47, Taf. xii, f. 7, 8. 

„ Byllis brevicomis, Grube. Arch. f. Naturges., Bd. xxix, p. 44, Taf. iv, f. 7. 
1872. Umbellisyllis fasciata, M. Sars and Gr. 0. Sars. Remark. Forms JSTorweg. Coast, p. 41, p]. i v 

f. 12—18. 
1875. Odontosyllis brevicomis, Marenzeller. Sitz. Akad. Wiss. Wien, Bd. lxxii, p. 32, Taf. iv, f. 1. 
„ „ gibba, Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., t. ii, p. 38, pi. iii, f . 10 and 10 b 

pi. iv, f. 10 a and 10 c. 
1885. „ „ Cams. Fauna Medit., II, p. 230. 

1904. „ „ Allen. Journ. M. B. A., n.s., vol. vii, p. 220. 

1905. „ „ G-raeife. Arbeit. Zool. Stat. Triest, xv, p. 323. 

1906. „ „ De St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 185. 

Habitat. — Dredged on dead shells in 15 fathoms off St. Peter's Port, Guernsey, July, 
1868. It also occurs at Kingstown Harbour where it was procured by Professor Haddon. 
Dredged at Plymouth (Dr. Allen) ; St. Andrews Bay. 

Shores of France — Normandy and Marseilles (deeper coralline region). Adriatic 
(Grrube and Marenzeller). 

Head (Plate L, fig. 5, and Plate LIX, figs. 14 and 14 a) small, rounded, or bluntly- 
conical, with two reddish eyes on each side, the anterior pair the wider apart. Two short 
anterior tentacles, and a median in front of the eyes with a dark tip. The tentacular 
cirri have a darker colour than the general fawn of the region. The flap-like process at 
the posterior part of the head moves to and fro in life. 

Body fully half an inch long, most distinctly tapered posteriorly, somewhat rounded 
dorsally and flattened ventrally ; segments 43 — 48. Of a pale brown or fawn colour 
with a whitish stripe interrupted by three or four brownish touches along the dorsum, 
and a transverse white belt on the seventh segment extending to the bases of the feet. 
The gibbous segment succeeding the head is somewhat semicircular or half shield-shaped 
and bears two clavate cirri with a white ring on the dilated region. This prominent 
shield-shaped region overlaps the head and is very conspicuous from its lighter hue. The 
four succeeding segments are somewhat smaller than those which follow, and each has a 
whitish median patch in continuation of the dorsal belt, and the last (fifth) of these has, 
in addition, three white clots before and behind its white touch. The sixth segment has 
a whitish patch at the bases of the feet and two brown dots in the centre, and it commences 
the somewhat larger region of the body. The rest of the segments are nearly alike except 
that they diminish in size towards the tail. The terminal segment has two short and 
somewhat clavate cirri with a white touch at the tip. Under surface with some dark 
touches in front. 

The southern examples, as described by Marion and Bobretzky, have each segment 
ornamented by a double transverse line of violet which reaches the base of the dorsal cirri. 
A similar line occurs at the segment-junctions. An irregular touch of yellowish orange 
is distributed irregularly on the lateral regions. Chalky touches occur on the palpi, the 
median tentacle, and here and there on the dorsal surface posteriorly. Many of the dorsal 
cirri have touches of violet towards their extremities ; others again have this replaced by 
the chalky (white) pigment. 



ODONTOSYLLIS GIBBA. 185 

In the example from Plymouth the colouration consists of numerous isolated brownish 
specks (Plate XLIX, fig. 4) which here and there form a band. 

A figure of the proboscis is given by Marion and Bobretzky, showing a series of 
denticulate papillaB anteriorly. 

The foot (Plate LXX, fig. 13) is uniramous in so far as there is only one spine. The 
dorsal moiety of the foot, however, may be held to be represented by the elevated region 
at each side from which the dorsal cirrus springs. The cirrus is clavate — somewhat like 
that of Lepidonotus — with a dilated region towards the tip, followed by a slender 
terminal process. The anterior cirri have the largest dilatations, the posterior being 
more slender, though still rendered characteristic by the white ring towards the tip. The 
setigerous region is separated from the foregoing by a marked sulcus, the dorsal margin 
of the lobe being convex, and it terminates externally in two short conical papillae, while 
beneath the lobe is a larger conical process, which may represent the ventral cirrus, 
unless that is altogether absent. It does not extend quite so far outward as the upper papillge. 
The bristles (Plate LXXIX, fig. 7) have a long, slender, translucent shaft, curved towards 
the dilated distal end where it has a shoulder and a bevelled edge. From the shoulder 
passes off the terminal blade, which is short, very slightly diminished at the tip, and ends 
in a simple hook, a feature not clearly indicated by Marion and Bobretzky. The bristles 
form a diminishing series from above downward both as regards the length of the shaft 
and the terminal pieces. The ventral cirrus forms an elongated conical process. 

This species is Hesione-like in its active movements and irritability. When at rest the 
anterior six segments of the body are elevated. Marion and Bobretzky's examples swam 
freely to and fro like Nereids. It differs from the form described by Claparede and Marion 
and Bobretzky in the arrangement of the eyes, the anterior and posterior pairs of which 
are more widely separated ; and still more in the shape of the dorsal cirri, which instead 
of forming fusiform or gently clavate organs are somewhat cylindrical with a rather 
sudden bulbous dilatation distally, from which a slender filiform tip projects. On the 
whole, however, in the present state of our knowledge it will suffice to indicate the variety, 
especially as the bristles do not appear to differ materially in structure. 

A similar form (young?) comes from the tow-net in St. Andrews Bay, 1894. The 
setigerous process of the foot seems to differ slightly in its relation to the ventral cirrus, 
but this may be due to youth. 

Claparede and Grrube described and figured this form within the same year, and it 
appears to be extensively distributed. 

Malaquin 1 (1893) observes that the protoplasmic " setting " of the muscular fibres of 
the proventriculus has totally disappeared, only the longitudinal fibrils of the column 
being present. Each column has transverse striation as in Syllis hyalina. 

So far as can be made out the Umbellisyllis fasciata of M. Sars 2 appears to be this 
species, though the large size (24 mm.) gives a margin for variation, and the presence of 
an additional minute pigment-speck in front of the anterior pair is novel. Whatever 
may be the condition as regards specific identity, the Norwegian form is an Odontosyllis 
from deep water — viz., 100 to 300 fathoms. 
1 Op. cit., p. 231. 

3 ' On some Remarkable Forms of Animal Life, etc./ 1872, p. 41, pi. iv, figs. 12 — 18. 

81 



186 SYLLINE KUBROPUNCTATA. 



Genus LIII. — Sylline, Grube, 1860. 

Head as in Autolytus, but the palpi, though coalesced at the base, are separated 
distally. Tentacles and tentacular and dorsal cirri not articulated. Body as in Autolytus. 
Feet with a smooth, tapering dorsal cirrus ; setigerous region rather short, and having 
beneath it a broadly lanceolate ventral cirrus. Spines with a hammer-like tip ; bristles 
with a short terminal piece ending in a single boldly-curved hook. 



1. Sylline rubropunctata, Grube, 1860. Plate XL VIII, fig. 5 ; Plate LXX, fig. 26- 
foot; Plate LXXIX, fig. 26— bristle. 

Specific Characters. — Head as in the type, only the short palpi are clearly separated 
by a median fissure. Eyes two on each side, the anterior being the larger and wider apart ; 
both with lenses — indeed, the anterior show two. Median and lateral tentacles long. 
Tentacular cirri as in Procersea (Marion ,and Bobretzky). Body about 20 mm. in length, 
and with 120 or more setigerous segments. Dorsally the translucent body is marked by 
a series of orange touches — two or four in number — longitudinally arranged, the buccal 
segment, however, having only the two outer. Minute, clear corpuscles occur in the skin, 
forming a transverse line in the segment (Marion and Bobretzky). 1 

Eoot rather prominent, with a considerable dorsal cirrus tapered distally Cirri of 
the first fourteen feet alternately longer and shorter. The setigerous region is short, 
with the long slope inward ventrally. Two spines pierce the upper angle of the region, 
their tips having a shoe-like modification, or shaped like a hammer. The bristles are 
slightly yellowish, curved distally, and bear a short, simply-hooked terminal piece. The 
enlarged end of the shaft is spinous on the convexity and sides of the peak. 

Synonyms. 

1860. Sylline rubropunctata, Grube. Arch. f. Naturges., Bd. xxvi, p. 87, Taf. iii, f. 8. 

1861. „ „ idem. Ausflug Triest, p. 144, Taf. iii, f. 8. 

1875. Autolijtus (Procersea) ornatus, Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., torn, ii, p. 44, 

pi. v, f. 14. 
1879. Procersea rubropunctata, Langerhans. Zeitscli. f. w. Zool., Bd. xxxii, p. 579, Taf. xxxii, 

f. 30. 
1890. Autolytus rubropunctatus, Malaquin. Annel. Boulonn., Rev. Biol. Nord Fr., t. iii, p. 179. 
1904. „ „ Allen. Journ. M. B. A., n.s., vol. vii, p. 220. 

Habitat. — Dredged on Queen's Ground, Plymouth (Allen). 

Porto re in the Adriatic (Grube) ; Marseilles, from the deeper waters inhabited by 
Gorgonia (Marion and Bobretzky) ; Madeira (Langerhans). 

Head ovoid, with a median, longitudinal sulcus ; the fused palpi are separated in front 
by a median fissure. Eyes black, two on each side, the anterior the larger and wider apart. 

1 Yiguier questions if a dorsal cirrus is present on the second bristled segment, but it seems to 
occur in the British specimen. 



SYLLIDEA (PROPER). 187 

Both have lenses, the specimen, moreover, showing in the anterior eyes a larger lens and 
a smaller in front of it. The median and lateral tentacles are long, the former being 
frequently carried backward. All have palpocils and "corpuscles brillants " (Marion and 
Bobretzky). Tentacular cirri two pairs, the dorsal in each being as long as the median 
tentacle. 

Bod}/ about 20 mm. in length, a little tapered anteriorly, more distinctly tapered 
posteriorly, and ending in two caudal cirri. The pale body is marked by two touches of 
orange on the first segment, by four on the succeeding, and has a series of minute 
corpuscles often arranged as a transverse band anteriorly. The first dorsal cirrus is as 
long as the median tentacle. 

Foot (Plate LXX, fig. 26) well marked, with a smooth dorsal cirrus about equal to 
the diameter of the body, and the comparatively prominent setigerous region has the 
form of a truncated cone, with two or three spines piercing the upper angle, each of these 
having a modification of the tip, as in every case this is bent as well as blunt. The 
bristles are slightly yellowish, and form a considerable group below the spines. The shaft 
has a ventral curve distally (Plate LXXIX, fig. 26), is enlarged and bevelled, with 
distinct spines along the convex (upper) border and peak, and a short terminal piece with 
a single bold terminal hook. The concavity below the hook is crescentic, and an abrupt 
edge bounds it inferiorly. The type of bristle thus differs essentially from that 
characteristic of Autolytus, since that is bifid. Moreover, the ventral cirrus is free 
distally, forming a broadly lanceolate process. 

Grube (1860), who did so much patient and excellent work in the group, first 
described this form as a Sylline, as its frontal lobes (palpi) had coalesced, its cirri were 
not articulated, and the ventral were altogether absent. His description, both in regard 
to structure and colouration, is easilv recognized, though the ventral cirrus is not absent. 

Marion and Bobretzky fifteen years later (1875) published an account of the same 
annelid under the title of Autolytus (Procersea) ornatus, as a new species. They gave a 
good figure, but did not add much to Grube's original description. 

Langerhans (1879) found large examples of the same form at Madeira, and describes 
the four dorsal touches as red, the lenses of the eyes as somewhat cylindrical, the pharynx 
(proboscis) with thirty teeth, and the proventriculus with thirty rows of points. He 
shows two kinds of terminal pieces to the bristles, viz. bifid and slender simple. So far 
as can be ascertained, the former would appear to refer to the bristles seen on edge. 
Females carried white and reddish eggs. 

Malaquin (1890) regards this species as synonymous with the Autolytus (Procersea) 
ornatus, De St. Joseph, 1 a view adopted here. 



Group III.— SYLLIDEA (PROPER). 

Syllida? provided with ventral cirri. Palpi free throughout their entire length. 
Tentacles and cirri moniliform, as if of independent articulations. Reproduction by 

1 ' Ann. Sc. Nat./ 1885, p. 220, pi. x, figs. 98—99. 



188 SYLLIS ARMILLARIS. 

stolons (schizogamy). Malaquin divides them into two primary gronps according to the 
presence or absence of a branchial appendage, and he separates the genera by the armature 
of the proboscis. Ventricule (stomach) well-developed, with lateral T-shaped pouches. 
Apparently only one genus of this group occurs in Britain. 



Genus LIV. — Syllis, Savigny, 1820. 

Head with three tentacles ; two pairs of tentacular cirri — all moniliform. Palpi 
separate throughout. Body more or less elongate, with distinct segments. Opening of 
the pharynx (proboscis) only with papilhe ; armature a single anterior tooth dorsally. 
Proventriculus short, two T-shaped glands behind. Feet with moniliform dorsal and 
filiform or pinniform ventral cirri. Falcate bristles with a short terminal piece ending in 
a single hook, or with simple bristles partially, rarely throughout. Tail with two 
elongated cirri. 

Reproduction normal, or by alternation of generations. Sexual individuals furnished 
only with two tentacles and devoid of tentacular cirri. Sexes similar. 



1. Syllis armillaris (Muller), (Ersted, 1842-3. Plate LX, fig. 1— head, fig. 2, Ioida— 
head; Plate LXX, fig. 14— foot; Plate LXXIX, figs. 8, 8 a, 9, and 9 a— bristles. 

Specific Characters. — Head rounded, about as long as broad, four reddish eyes obliquely 
placed, the anterior pair being the wider apart. Palpi ovate. Median and lateral tentacles 
moniliform. Body two or three inches long, slightly diminished in front and tapering to a 
lanceolate point posteriorly between the caudal cirri. Straw-yellow, streaked transversely 
by two lateral bars and a central bar of greyish brown in each segment, so that anteriorly 
there are three rows of dark touches. Proboscis with a single conical tooth, and ten papillse 
on its margin. 1 The foot has dorsally a moniliform cirrus of 10 — 12 segments, barely 
equalling the diameter of the body, and carried alternately horizontal and erect. 
Setigerous region bluntly conical — bearing spines and a group of strong bristles, which in 
the anterior segments have a short spinous terminal piece with a simple hook at the tip, 
which points upward in the normal position of the foot. Ventral cirrus lanceolate. 

Synonyms. 

1776. Nereis armillaris, 0. F. Muller. Zool. Dan. Prodr., p. 217. 

1780. „ „ 0. Fabricius. Fauna Grroeul., p. 294. 

1791. „ „ G-melin. Linn. Syst. Nat., i, pt. vi, p. 3115. 

1800. Die geperlte Nereide, O. F. Muller. Xaturges. einiger Wurm-Arten, p. 150,, Tab. ix, f. 1 — 5. 

1806. Nereis armillaris, Turton. Linn. Syst. Nat., iv, p. 86. 

1820. Lycastis armillaris, Savigny. Syst. Annel., p. 45, pi. iv, f. 3. 

1824. Nereis armillaris, Bruguiere. Tabl. Encycl. Meth., Vers, I, p. 134, Tab. Iv, f. 13 — 17. 

1825. „ „ De Blainville. Diet. Sc. Nat., xxxiv, p. 436, pi. xiv, f. 2. 
1828. Nereisyllis monilaris (and ornata), idem. Ibid., p. 473, t. lvii. 

1 Vide body-wall in Fig. 50, p. 139. 



SYLLIS AHMILLARIS. 189 

1830. Nereis armillaris, Bosc. Vers, ed. 2, i, p. 168. 

1834. Syllis „ Audouin and Edwards. Annel., p. 205, pi. ivb, f. 1 — 5. 

1840. Ioida macrophthalma, Johnston. Ann. Nat. Hist., iv, p. 231, pi. vii, f. 5, and p. 343. 

1842-3. „ „ GErsted. Kroyer's Nat. Tids., p. 118. 

„ Syllis armillaris, idem. Ibid., p. 117. 

1843. „ „ idem. Annul. Dan. Consp., p. 24, figs. 27, 90, 94, 102. 
„ „ „ idem. Gronl. Annul. Dorsib., p. 181. 

1844. „ „ Johnston. Ann. Nat. Hist., xiii, p. 438. 

„ „ „ Thompson. Eep. Brit. Assoc., 1843, p. 273. 

1844-5. „ „ (Ersted. Kroyer's Nat. Tids., p. 408. 

1845. „ „ Johnston. Ann. Nat. Hist., xv, p. 145, pi. ix, f. 1, 2. 
1851. „ „ Grnbe. Fam. Anuel., pp. 61 and 132, 

„ Sars. Nyt Mag., vi, p. 209. 

1865. „ arnica, De Quatrefages. Annel., ii, pp. 20 and 87. 

,, ,, danica, idem. Ibid., p. 21. 

„ „ armillaris, Johnston. Cat. Brit. Mus., p. 191, pi. xiv a, f. 1, 2. 

„ Ioida macrophthalma, idem. Ibid., p. 197, pi. xiv a, f . 5. 

1867. Syllis armillaris, Malmgren. Annul. Polych., p. 42, Tab. vii, f. 46. 

„ „ „ Parfitt. Trans. Devon. Assoc., ii, pt. i, p. 236. 

1869. „ „ Mcintosh. Trans. R. S. Edin., xxv, p. 414. 

1874. „ „ idem. Ann. Nat. Hist., ser. 4, vol. xiv, p. 198. 
„ Ioida macrophthalma, idem. Ibid., p. 198. 

„ Syllis armillaris, Malm. Op. cit. Goteb., p. 82. 

1875. „ „ Mcintosh. Invert, and Fishes St. Andrews, p. 121. 
„ Ioida macrophthalma, idem. Ibid., p. 122. 

„ Syllis armillaris, Mobius. Jahresb. Comm. Deutsch., p. 169. 

1879. ,, „ Tauber. Annul. Danic., p. 94. 

„ „ „ Langerhans. Zeitsch. f. w. Zool., Bd. xxxii, p. 535. 

1883. „ „ Levinsen. Vidensk. Meddel. Foren Kjobenh., p. 245. 

1886. „ „ Harvey Gibson. Proc. Lit. Phil. Soc. Liverp., vol. xi, p. 153. 

1890. Ioida macrophthalma, Giard. Bull. Sc. Fr. Belg., t. xxii, p. 79. 
„ Syllis armillaris, Malaquin. Annel. Boulon., p. 35. 

1891. „ „ Hornel. Trans. Liverp. Biol. Soc, vol. v, p. 245. 
1893. „ „ Levinsen. Vidensk. Ud. "Hauchs," p. 330. 

Habitat. — A comparatively common species, under stones near low water-mark, 
St. Andrews, and ranging to the coralline ground ; in the interstices of Filigrana implexa 
from the Bell Rock ; under roots of Laminaria in rock pools beyond Maiden Hock ; amongst 
debris from deep sea fishing ; dredged in the laminarian region ; under stones between 
tide-marks, Lochmaddy, North Uist, and in muddy fissures of rocks, near low water-mark 
in tracks. 

Faroe (Willemoes-Suhm) ; shores of Norway and Sweden; Greenland ((Ersted), and 
Behrings Sea (Marenzeller). 

Head (Plate LX, fig. 1) rounded, about as long as broad, with the reddish eyes 
obliquely placed on each side, the anterior and larger pair being the wider apart. Palpi 
somewhat ovate. Median and lateral tentacles moniliform, the former the longer. 

Body two or three inches long and of sixty to seventy segments, slightly diminished in 



190 SYLLIS ARMILLARIS. 

front, tapering to a lanceolate point posteriorly, and furnished with two cirri. It is 
straw-yellow or slightly skin- coloured, streaked transversely by two lateral bars and a 
central bar of greyish-brown in every segment, the effect being that anteriorly three 
rows of dark touches are visible. 

The proboscis has a single conical tooth anteriorly, and about ten papillse on its 
margin. 

The foot (Plate LXX, fig. 14) has dorsally a moniliform cirrus of ten to twelve 
segments tapering towards the tip, barely equalling the diameter of the body, and 
alternately carried horizontal and erect. Beneath is a short, bluntly-conical, setigerous 
region bearing a series of stout spines, a point piercing the surface at the upper angle, 
and a group of strong bristles (Plate LXXIX, figs. 8 and 8 a) with the shafts curved 
towards the tip, which is bevelled, and bears a terminal piece of considerable length in the 
anterior segments, a simple hook at the tip, and an edge covered with spikes (though 
these are often worn). In the posterior segments the terminal piece is shorter; indeed, in 
the anterior feet the upper bristles in each foot have longer terminal pieces than the 
lower. The hooks of the bristles point upward. Ventrally is a lanceolate lamella (cirrus). 

In some a tuft of extremely fine, translucent bristles, above the jointed forms, 
indicates the formation of the sexual stage. 

Habits. — It is a hard, stiff form, and comparatively sluggish, hiding under debris in 
a vessel, or crawling slowly over the bottom. 

H. Rathke's Syllis tigrina 1 comes very near this form, and as subsequent authors have 
not identified it, in all probability it refers to a variety of S. armillaris. 

The sexual form of this species is the Ioida macrojphtlialma of Dr. Johnston (1840) 
and subsequent authors. The following example appears to pertain to the same form 
as Dr. Johnston's, but there are certain discrepancies, so that further investigation is 
necessary. 

Habitat. — On a laminarian blade covered with Obella from the West Sands, St. 
Andrews, after a storm, October 25th, 1864. 

Head (Plate LX, fig. 2) small, with a slender median and two lateral tentacles, and 
two large dark eyes. 

Body about three quarters of an inch long, with the segments and feet well marked. 

The foot in the first six segments presents no capillary bristles. Dorsally is the 
moniliform cirrus of twelve or thirteen segments, tapering to the tip, then a considerable 
interval, with the tip of a spine projecting from its surface. The conical setigerous region 
has a group of jointed bristles, with simple hooked tips (hook directed upward) as in S. 
armillaris, these tips being longer in the anterior bristles than in the median (Plate LXXIX, 
fig. 9 a). A ventral lobe occurs beneath of an elongate conical form, thus differing from 
the lobe in Syllis armillaris. 

This apparently is the form described by Dr. Johnston, but there is doubt, since the 
long swimming-bristles are stated to be inferior and the jointed superior. This is 
probably due to a misprint, 

1 < Beitr. Fauna Norweg./ p. 165, Taf. vii, f. 9—13 (1843). 



SYLLIS CUCULLATA. 191 

The feet (Plate LXX, fig. 15) bearing the long swimming-bristles (Plate LXXIX, 
fig. 9) have the region between the cirrus and the setigerous lobe more prominent, and 
the strong translucent and long bristles piercing a central eminence supported by a spine 
of a more slender type than those of the inferior lobe. These long bristles are beautifully 
iridescent. The conical setigerous lobe follows, its strong spine piercing the tip above the 
bristles, which have the hooks of the terminal piece directed upward (as usual). The 
ventral lobe agrees with that in front. 

This is probably the Nereis scolojpendroides of Montagu (MS. Drawings, Linn. Soc, 
Plate XXXVIII, fig. 1, 1808). 

De Quatrefages treated this form as pertaining to a special genus. 

An Ioida (Plate LIX, figs. 16 and 16 a) procured at Plymouth by Dr. Allen in 
September, appears to differs from any procured in the north, and probably pertains to 
a southern species. 

The head is deeply bilobed and furnished with two very large, somewhat ovoid eyes 
on each side. The foot shows a crenate dorsal cirrus of considerable length. Swimming- 
bristles occur dorsally. The setigerous region bears a series of bristles, but as the 
specimen was lost, their structure is unknown. The tail has no median process. The 
softness of the crenated cirri would suggest a connection with Pionosyllis prolifera or 
a nearly related form. 

The genus GhdetosyUis of Malmgren (1867) refers to a nearly allied, though probably 
different, sexual form distinguished by the great length of the tip of the falcate bristles. 

The Tetraglene rosea of Grube, 1 from the Adriatic, is apparently the loida-stage of 
a Syllis, having a moderately long slender tip to the compound bristles. Small simple 
bristles occur in the dorsal division, and long swimming-bristles of the typical form are 
found in the ventral division of the foot. 



2. Syllis cucullata, n.s. Plate LX, fig. 3 — head; Plate LXX, fig. 17 — foot; Plate 

LXXIX, ^g. 11— bristle. 

Specific Characters. — A somewhat massive species, with four rather large eyes nearly 
in a square. Palpi of considerable size. Tentacles and tentacular cirri of moderate length 
and crenated — especially towards the tip. Foot with a bluntly-conical setigerous region 
and two hooked spines. Bristles rather stout, translucent, dilated and bluntly bevelled 
at the end of the shaft, and furnished with spines. Terminal piece short, simply and 
boldly hooked. 

Habitat. — Under stones between tide-marks, Luccomb Chine, Isle of Wight. 

Head (Plate LX, fig. 3) rounded, with two considerable palpi in front. Eyes 
moderately large, the anterior pair only very slightly the wider apart, so that they nearly 
form a square. Tentacles and tentacular cirri of moderate length — slightly moniliform. 

Body comparatively firm and broad, little tapered anteriorly, but diminishing to a 
slender tail posteriorly. Proboscis exserted. 

1 < Arch. f. Xaturges./ Bd. xxix, p. 42, Taf. iv, fig. 6 (1863). 



192 SYLLIS KROHNII. 

The foot (Plate LXX, fig. 17) lias the dorsal cirrus crenate rather than moniliform, 
and tapered. It arises from the dorsum without a distinct basal region. The setigerous 
lobe is bluntly conical, with the trend of the distal margin downward and slightly inward, 
and having the usual furrow for the bristles. The two spines reach the surface at the outer 
and upper angle and have distinctly hooked tips which point upward. The bristles (Plate 
LXXIX, fig. 11) have strong translucent shafts with dilated and bluntly-bevelled ends 
without evident serrations, though two spikes occur at the point. The terminal piece is 
short, ending in a simple hook, occasionally with a minute spike on the edge. The ventral 
lobe is broadly ovate and does not quite extend to the tip of the setigerous region. 



3. Syllis Krohnii, Elders, 1864. Plate XLIX, fig. 6; Plate LXXIX, fig. 12— bristle. 

Specific Characters. — Head notched in front, somewhat narrowed behind, and with 
two prominent, tapering, ciliated palpi. Eyes four, reddish, placed obliquely in pairs, the 
anterior being the wider apart. Median tentacle shorter than the lateral. It arises between 
the eyes, and is speckled with white dots. The dorsal tentacular cirri are longer than 
the lateral. All are slightly annulated and have white specks and whitish tips. Body 
about an inch and a half long, little diminished in front, but tapering posteriorly to a slender 
tail, with a median papilla and two lateral cirri. Dorsum pale anteriorly, with about a 
dozen madder-brown belts, the anterior being double. Ventral surface pale anteriorly, 
pale orange behind, and translucent towards the tail, which has two long cirri. Dorsal 
cirri alternately large and small. Proboscis with a denticulated edge and a prominent 
tooth. The setigerous region has a series of translucent bristles with a trace of serrations 
on the convex edge of the distal enlargement of the shaft. Terminal piece of moderate 
length with a simple hook at the tip and a finely spinous edge — the spine next the tip 
being so prominent as to give a pseudo-bifid appearance to the tip. Ventral cirrus of an 
elongated conical form. 

Synonyms. 
1864. Syllis Krohnii, Ehlers. Borstenw., I, p. 234, Taf. x, f. 1—4. 
1805. ,, „ De Quatrefages. Anneles, ii, p. 31. 

1869. „ „ Mcintosh. Trans. E. S. Edin., vol. xxv, p. 415, pi. xvi, f. 14. 

1881. Typosyllis Krohnii, Langerhans. Can. Annel., Nov. Act. Leop.-Car. Deutsch. Akad., xlh, 

p. 102 (?). 
1885. Syllis „ Cams. Fanna Medit., p. 225. 

1906. Syllis (Typosyllis) Krohnii, De St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 180. 

Habitat. — In a tube of sand under a stone in a rock-pool near low water at Paible, 
North Uist. It may have crept accidentally into the tube, for it generally prefers to hide 
itself under debris in a vessel. Between tide-marks, Herm. Benham 1 states that it is 
abundant under stones, and forms tubes, but gives no locality. It is not common in 
the author's experience. 

i f Camb. Nat. Hist./ ii, p. 307. 



SYLLIS KEOHNIL 193 

Adriatic (Ehlers) ; Mediterranean ; Canaries (Langerhans) ; shores of France (De St. 
Joseph). 

Head somewhat narrowed behind, notched in front and with two prominent 
tapering palpi, which are ciliated. Eyes four, reddish, placed obliquely in pairs, the anterior 
pair being the wider apart. The moniliform median tentacle is less elongated than the 
lateral cirri, and projects upward from its attachment between the eyes. It is speckled 
with opaque white dots. The lateral tentacles are shorter, but of similar shape to the 
median. The buccal segment has two cirri, the dorsal being longer than the lateral 
tentacles. All these organs are slightly annulated, and have white specks and whitish tips. 

Body about an inch and a half long when extended, linear, very little tapered in front, 
but diminishing posteriorly to a slender tail which has a median papilla and two cirri. 
Dorsum pale anteriorly, marked transversely by more than a dozen madder-brown bands, 
the anterior being double. The rest of the dorsum is opaque yellowish white (from the 
viscera), paler at the sides and posteriorly. The under surface is pale anteriorly, pale 
orange throughout the greater part of its length, and translucent towards the tail. 

The proboscis has its anterior edge denticulated, and stretches from the second to 
the sixth segment. Opposite the fourth segment of the body the organ has a prominent 
tooth. The proventriculus extends from the sixth to the tenth segment. The region of 
the canal behind shows the lateral cseca, which, springing about the twelfth, pass 
forward to the ninth, and backward to the thirteenth. The gut is ciliated internally and 
of a brownish colour. 

The segments throughout are distinctly marked, each furnished with a large dorsal 
cirrus alternately with a smaller one. The former are a third larger, more opaque, 
speckled with distinct white dots, and instead of passing transversely outward as the 
others do, they are borne upward in a graceful manner, the tips usually curved inward or 
coiled in various ways. The other cirri are smaller, paler, also speckled with white dots, 
and longer than the diameter of the body. Both are somewhat tapered towards the tips, 
and at the base of the anterior cirri is a brown speck. The tail terminates in two long 
cirri. 

The setigerous region of the foot bears a series of translucent bristles (Plate LXXIX, 
fig. 12), the shaft of which is slightly curved distally and the tip dilated and bevelled, a 
trace of serrations occurring on the convex edge. The terminal piece is of moderate 
length with' a simple hook at the tip, and the edge of the process is finely spinous, the 
first spine being so prominent as to give a pseudo-bifid aspect to the tip. 

The specimens were unfortunately so softened by decay after death that the outlines 
of the foot could not be safely drawn. The ventral cirrus forms an elongated cone. 

In Ehlers' figure the alternate cirri are club-shaped, a feature not observed in the 
healthy specimen from North Uist; whilst, again, the cirri of the third and fourth 
segments were nearly equal, whereas Ehlers shows a clavate pair. All the cirri are much 
more distinctly articulated than in the British example. 

Reproduction. — A large example, procured in August under a stone between tide- 
marks at Herm, had a bud at its posterior end. It agreed with the coloured figure for 
the most part, every alternate dorsal cirrus being long and generally carried with a 
coiled tip. Behind the madder-brown bars in front is a pale yellowish part and then a 

82 



194 SYLLIS MACEOCEBAS. 

region of considerable extent marked by brownish pinnae from the intestine, and speckled 
with white. 

The bud had a salmon-tint and was full of ova, and next day it became detached. It 
is wider than its nurse-stock, has a small head with four large red eyes, only two of which, 
however, are visible from the dorsum, though seen when the head is viewed obliquely. 
They subserve ventral vision. The cirri of the bud are nearly equal. 

Habits. — The species is hardy. The specimen captured in August at Paible was 
carried to Lochmaddy, and after a few days across the Minch to Dunvegan, and thence to 
Murthly, where it was figured by my sister. It lived for five months in confinement fifty 
miles from the sea, in a tumbler of sea- water, the only change being an increase in 
translucency and a diminution in the size of the cirri. The sea-water was rarely changed. 

When placed in a flat vessel it generally makes for the side, where it remains 
quiescent. It sometimes coils both larger and smaller cirri upward in repose, but 
generally the smaller remain extended transversely. It is fond of twisting itself and 
turning over and over, and occasionally coils itself into a ball, ventral surface outwards. 
The movements of the cirri are many and graceful, and render the species one of the 
most beautiful of the British Syllicls. 

4. Syllis macroceras, Orube, 1857. Plate LXX, fig. 18— foot; Plate LXXIX, fig. 13 and 

13 a — bristles. 

Specific Characters. — Head about as long as broad, with two long palpi. 1 Eyes four, 
dark reel, arranged nearly in a rectangle, though the anterior pair are wider apart. 
Median tentacle longer than the lateral, but shorter than the third from the front. These 
processes are not articulated. Body about half an inch in length, and with 100 segments. 
Colour dull orange, paler in front and at the tail which has a pair of cirri. Foot with a 
long, tapering, smooth dorsal cirrus ; the setigerous region is conical, but the inferior 
slope trends away more rapidly than the superior. Spines with a blunt, curved tip. 
Bristles translucent, little modified in each group, with a short terminal piece which shows 
a well-marked hook at the tip and no trace of spines. The dilated distal region of the 
shaft is short and has indications of serrations on the bevelled edge. Ventral cirrus large, 
broadly ovate, and does not reach the tip of the setigerous region. 

Synonyms. 
1857. Syllis macroceras, Grube. Annul. (Ersted (Nat. Foren. Vidensk. Meddel.)., iii, p. 28. 
1865. „ „ De Quatrefages. Anneles, II, p. 28. 

1869. „ macrocera, Mcintosh. Trans. R. S. Edin., xxv ; p. 416, pi. xv, f. 12. 

Habitat. — Procured under a stone near low water-mark, Lochmaddy, North Uist; 
in the Grouliot Caves, Sark, under stones ; and in the tidal region at Whitecliff Bay, Isle 
of Wight. 

A widely distributed form — ranging to St. Croix, Christiansted (Grube). 

Head about as long as broad, with two long palpi in front. Median tentacle longer 
than the two lateral, but shorter than the third from the front. These organs are not 

1 Grube states that they are connate to the middle, but they were not so in these examples. 



SYLLIS CITXMNGHAMI. 195 

articulated. Eyes four, dark red, arranged nearly in a rectangle, though the anterior pair 
are wider apart. 

Body about half an inch in length, of a dull orange-yellow colour, paler in front 
and at the tail, which has a pair of cirri. The segments are about 100. 

Foot (Plate LXX, fig. 18) with a long, tapering, smooth, dorsal cirrus. The seti- 
gerous region is conical, the inferior slope trending away more rapidly than the superior. 
The spines, which pierce the upper angle, often have a blunt curved tip. The trans- 
lucent bristles (Plate LXXIX, figs. 13 and 13 a) have a very short terminal piece with a 
well-marked hook at the apex, whilst the abbreviated edge shows no traces of spikes. 
These terminal pieces do not vary to the same degree as, for instance, in Syllis armillaris. 
The dilated distal region of the shaft is short with indications of serrations on the 
bevelled edge. The ventral cirrus is large, broadly ovate, and does not reach the tip of 
the setigerous region. 

The long simple (swimming-) bristles are developed in the posterior region of one 
from Lochmaddy, indicating the formation of a bud with the reproductive elements. 

In transverse section the ventral longitudinal muscles are separated by the nerve- 
cords, which are comparatively large and have the basement-tissue, hypoderm, and cuticle 
externally. 

This form seems to agree with Grube's description in regard to the arrangement of 
the eyes and the form of the head, but the palpi were not connate. The author points out 
the shortness of the terminal pieces of the bristles, which he describes as almost equilateral, 
and the absence of articulations in the cirri, thus differing from all except Syllis longo- 
cirratus, in which, however, the tip is dilated. His example came from St. Croix, 
Ohristiansted. 

5. Syllis Cunningham!, 1 n. s. Plate LXX, fig. 19 and 19a — foot; Plate LXXXIY, 

fig. 6 — bristle. 

Specific Characters. — Head apparently typical, though eyes not seen. Tentacles and 
cirri articulated. Body normal. Attached posteriorly was a male bud considerably larger 
in diameter than the parent-stock. Foot anteriorly has a somewhat stout tapered cirrus 
of about seventeen segments. Setigerous region rather narrow and elongate when seen 
from above ; obliquely conical in lateral view. Xext the spines are two bristles larger 
than the others and apparently devoid of terminal pieces. The smaller bristles beneath 
are of average size, and have short terminal pieces — spinous on the edge and with a 
simple hook at the end. The ventral cirrus is narrow and elongate, but does not reach 
the tip of the setigerous region. 

Habitat. — In the tracks of Spio in fissures of the hard rocks between tide-marks. 
Cobo Bay, Guernsey, 1868. 

Head apparently typical, but the eyes had disappeared before it was critically 
examined. The tentacles and tentacular cirri are of average length and articulated. 

Body about three quarters of an inch in length and normal. The posterior region 
1 Xamed in honour of my late valued and eloquent colleague. Principal John Cunningham, of St. 
MaiVs College, St. Andrews. 



196 SYLLIS BREVICIRRATA. 

was occupied by an opaque bud distended with male elements, and much more bulky than 
the caudal region of the adult in front. 

The foot (Plate LXX, fig. 19) anteriorly shows a somewhat stout tapered cirrus of 
about seventeen segments. The setigerous region is rather narrow and elongate when 
seen from above, obliquely conical in lateral view. The two spines, which have a blunt 
tip, pierce the upper angle. Next these are two bristles much longer than the others, 
and apparently devoid of terminal pieces. The group of smaller bristles of the average size 
beneath show short tips with a simple hook and a spinous edge (Plate LXXXIV, fig. 6). 
The ventral cirrus is narrow and elongate, but does not reach the tip of the setigerous 
region. 

The bud was nearly ready for separation. The foot (Plate LXX, fig. 19 a) has a dorsal 
cirrus of about thirteen articulations with a tuft of slender, tapering, swimming-bristles 
beneath. The setigerous region has a similar outline to that of the nurse- stock, the spines 
end in a probe-tip; the two upper bristles are larger and devoid of terminal pieces, and the 
inferior bristles correspond with those in the adult. The ventral cirrus is also similar. 

This species presents an intermediate condition to that of Syllis gracilis and Syllis 
sjpongicola, in which a further differentiation of the tip of the shaft of the enlarged bristles 
occurs. Subsequent investigation may show a closer relationship with either of these than 
at present can be made out. The occurrence of these enlarged bristles anteriorly is 
noteworthy. 



6. Syllis breviclkrata, n. s. Plate LXX, fig. 20 — foot; Plate LXXIX, figs. 14 and 

14 a — bristles. 

Specific Characters. — Head resembling that of Syllis gracilis. Foot with a short 
dorsal cirrus of about nine segments. Setigerous region bluntly conical, the tips of the 
spines slightly dilated and bevelled. Bristles stout, terminal pieces comparatively short, 
with a spinous edge and a simple hook at the tip. Ventral cirrus tongue-shaped, scarcely 
projecting so far as the tip of the setigerous region. 

Habitat. — Between tide-marks, Herm, August, 1868. 

Head apparently similar to that of Syllis gracilis. 

Body about three eighths of an inch in length, tapered to a delicate tail furnished 
with two caudal cirri posteriorly, but only slightly diminished towards the head. The 
colour has disappeared. 

The foot (Plate LXX, fig. 20) has a short and rather thick dorsal cirrus of about 
nine segments. The setigerous region is bluntly conical with the tips of the spines, which 
are slightly dilated and bevelled, projecting at the upper angle. The bristles (Plate 
LXXIX, figs. 14 and 14 a) are somewhat stout, with curved shafts and dilated and 
bevelled tips. The terminal pieces are comparatively short and have a simple hook at the 
tip with a spinous edge below. The tongue-shaped ventral cirrus scarcely projects so far 
as the tip of the setigerous lobe, and thus is shorter than in S. gracilis. 

Posteriorly the same type of bristle is found in the feet, but further examination 
may show that certain sexual forms of S. gracilis, to which this form is evidently allied, 
may vary in this respect. 



SYLLIS SPONG-ICOLA. 197 

The relationships of this Syllis appear to be with such forms as Trypanosyllis zebra 
and perhaps 8. gracilis. 



7. Syllis spongicola, Grube, 1855. Plate LI, fig. 4; Plate LXX, fig. 21 — foot 

of bud; Plate LXXIX, fig. 15— bristle. 

Specific Characters. — Head comparatively small, with two long and large palpi 
tapered distally. Eyes four, small, the anterior the larger, accompanied or not by two 
pigment-dots anteriorly. Tentacles and tentacular cirri of moderate length, articulated. 
Body of about 76 — 100 segments, pale orange, translucent yellowish or sand-coloured, 
translucent bluish towards tip of tail, and from an inch to an inch and a half or more in 
length. Anal cirri short. Proboscis with ten soft papillae and a tooth. Proventriculus 
about the tenth segment, and occupying seven segments. Foot with a dorsal cirrus of 
25 — 30 segments. Bristles simple, of a peculiar hamate character, shaped somewhat 
like an abbreviated bill-hook. Ventral cirri leaf-shaped, not reaching the tip of the 
setigerous region. 

Synonyms. 

1829. Nereis teticola, Delle Cliiaje. Mem., pp. 175, 195, Tav. lxvi, f. v (?). 

1841. „ „ idem. Descrizione, v, p. 103. 

1851. Syllis tethycola, Grube. Fam. Annel., p. 62. 

1855. „ spongicola, Grube. Arch. f. Naturg., p. 104, pi. iv, f. 4. 

1864. „ „ idem. Insel. Lussin, p. 84. 

1865. „ spongicula, De Quatrefages. Annel., II, p. 88. 
1868. „ hamata, Claparede. Annel. Nap., p. 195, pi. xv, f. 2. 

1870. „ oligochseta, Bobretzky. Annel. de la Mer Noire, f. 51 and 52 {fide aut.). 
1875. „ spongicola, Marion and Bobretzky. Ann. Sc. Nat. 6 e ser., t. ii, p. 24. 

1878. „ violacea-flava, Grube. Ann. Semp., p. 115. 

}} „ hamata, Langerhans. Zeitsch. f. w. ZooL, xxxiv, p. 128. 

1879. „ spongicola, Marion. Ann. Sc. Nat., 6 e ser., t. viii, art. 7, p. 19, f. 4 — 4 e. 
1881. Haplosyllis hamata, Langerhans. Nov. Act. Leop. -Carol. Akad., p. 102. 

„ „ aurantiaca, Eisig. Mitt. Zool. Stat. Neap., II, p. 270. 

1885. Syllis hamata, Cams. Fauna Medit., p. 225. 
„ „ spongicola, idem. Ibid., p. 228. 

1886. Haplosyllis spongicola, Albert. Mitt. Zool. Stat. Neap., Bd. vii, pp. 1 — 20, pi. i, f. 1 — 17. 

i} „ (Syllis) hamata, Langerhans. Zeitsch. f. w. Zool., xxxii, p. 527, Taf. 31, 

f . 1 a and 1 b. 

1887. Syllis (Haplosyllis) „ De St. Joseph. Ann. Sc. Nat., 7 e ser., i, p. 142, pi. vii, f. 5 — 8. 
1893. Haplosyllis hamata, Marenzeller. Zool. Ergebn., II, p. 7. 

1895. „ „ De St. Joseph. Ann. Sc. Nat., 7 e ser., torn, xx, p. 185. 

1900. Syllis (Haplosyllis) djiboutiensis, Gravier. Nouv. Arch. Mus., 4 e ser. ii, p. 147, pi. ix, f. 3 (?). 

1905. „ spongicola, Willey. Eep. Pearl Fish. Ceylon, IV, p. 269. 
9i „ „ Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 323. 

1906. „ (Haplosyllis) hamata, De St. Joseph. Ann. Sc. Nat., 9 e ser., t. hi, p. 178. 



198 SYLLIS SPONGICOLA. 

Habitat. — Procured at Jersey by Mr. Hornel. Millbay Channel, Plymouth Sound 
(Dr. Allen). 

It was first found by Professor Grube at Triest ; whilst Claparede got it at Naples ; 
Bobretzky in the Black Sea; the same author and Marion at Marseilles. Shores of 
France (Baron de St. Joseph). Ceylon (Willey). Canaries; Madeira — between tide- 
marks amongst algae — rarely (Langerhans). 

Head comparatively small, of greater transverse than antero -posterior diameter, and 
with four eyes, the anterior and larger being the wider apart. An additional pigment-dot 
occasionally occurs anteriorly. The median tentacle is as long, if not longer, than the 
longest cirrus and moniliform like all the cirri. The lateral are shorter. Palpi rather 
large, often tapered anteriorly. Tentacular cirri considerably shorter than the median 
tentacle, the upper longer than the lateral tentacles, the ventral shorter. 

Body of the typical form, segments 76 — 90, considerably tapered posteriorly, and 
ending in two cirri with a median papilla. The general hue is translucent yellowish with 
a brownish tinge from the intestine posteriorly, the tail, however, being translucent bluish. 1 
The proboscis occupies the first eleven segments, the proventriculus commencing in the 
twelfth, and extending for seven segments behind it. 

The first foot has the bifid strong hooks with occasionally a simple one (Claparede). 
The typical foot presents the comparatively short tapering cirrus. Beneath is the short 
obliquely-conical setigerous region with the spines passing to the upper angle. The 
bristles are simple, of a peculiarly hamate character (Plate LXXIX, fig. 15) — somewhat 
like a bill-hook, often bifid at the tip, the distal spikes apparently being longer in those 
uninjured by wear. The spines generally present a curved tip. The ventral cirrus 
is rather slender and conical, and its tip does not reach the extremity of the setigerous 
region. 

In a pelagic female bud the foot (Plate LXX, fig. 21) has the simple bristles in the 
setigerous region somewhat longer, whilst a long tuft of swimming-bristles occurs beneath 
the dorsal cirrus. A spine, moreover, supports the base of these. 

In a new Syllid recently described by Gravier 2 from Madagascar the bristles some- 
what resemble the foregoing, each being simple, and terminating in an enlargement 
obliquely bevelled, as if the distal piece had been lost. 

Body. — Marion and Bobretzky note that the colour is variable, the young being very 
transparent, whilst the adults are of a fine orange, sometimes opaque. 

Reproduction. — De St. Joseph 3 found that at the reproductive period the eggs fill 
segments 28 — 81, and that natatory bristles appear, whilst the segmental organs are 
indicated by two black points. The latter organs open on the one hand into the 
body-cavity, and on the other at the base of the feet, the apertures in each case being 
ciliated. The ova give the specimens a violet hue. He quotes Albert as having seen a 
bud without a head, detached, which swam away. Malaquin 4 refers to the same 
quotation, and calls the bud acephalous, and he has followed its further history. 

1 Claparede states that its pigment is continuous. 

3 ' Compt. Bend. 6 e Congres internat. ZooL, Berne/ 1904, p. 372, fig. 5. 

3 < Ann. Sc. Nat./ 7 e ser., p. 143. 

4 Op. cit., 1893, p. 333. 



SYLLIS SPONGICOLA. 199 

After swimming about it settles on a stone or on the bottom, and gradually the anterior 
end develops a head, first after the Tetraglene type, then that of Ghdetosyllis, thereafter a 
median and two lateral tentacles appear as in Syllis arnica. The Ioida stage follows, with 
moniliform tentacles and cirri. 1 Moreover, he found that the head, if removed, could be 
reproduced. The bud swims actively about and by-and-by discharges the sexual elements. 
The long natatory bristles are moved by muscles — as in Autolytus. Malaquin found 
only male buds in what he calls Syllis spongicola, and large female buds in 8. hamata. 
Langerhans describes the sperms as reddish, and the eggs as violet. Lo Bianco 2 states 
that Neapolitan specimens are ripe in September. In the sexual forms a pair of pigment- 
spots occurs on each segment, and Malaquin considers that they are veritable eyes, with 
refringent granules and rods, vitreous bodies and pigment. 

This is one of the many species science owes to the laborious Grube of Breslau, who 
in 1855 described it from an orange-coloured sponge, at Triest. He recognized most of 
its important features. 

The Nereis teticpla of Delia Chiaje (1829) is either this or an allied species somewhat 
approaching Syllis armillaris, he says, in general outline. 

Claparede (1868), in the specific description of this form (his S. hamata), observes that 
the dorsal cirri are slender and short, not attaining the diameter of the body. The form 
of the hooks he associated with its habits in tubes amongst Balani. He was uncertain 
whether it bored such tunnels or took advantage of those of Polydora and other forms. 
He thought it different from Grube' s S. spongicola, though the grounds for his opinion do 
not now seem to be sufficient. He mentions glands with transparent processes (boyaux) 
near the bases of the dorsal cirri. 

The Syllis setubalensis of the ' Challenger' 3 is probably only a variety of this species. 

A variety (tentacnlata) was found by Marion (1879) at Marseilles, having rather long 
tentacles. He gives a goocl figure of the extended proboscis with ten soft papillae and the 
median tooth, which seems to be large. 

Verrill figures a Syllis spongiphila in the collections of the ' Albatross,' which seems 
to approach this species, but no description has yet been studied. 

Langerhans (1879) considers this a representative of one of the oldest and simplest 
forms amongst the Syllidse. 

Marion and Bobretzky pointed out 4 the identity of Syllis hamata, Claparede, with 
Grrube's species. They mention that the young are transparent (translucent ?), whereas 
the adults are opaque orange. 

Albert (1886) has given a detailed account of the budding of this form at Naples, 
the stolons occurring posteriorly from the thirtieth to the sixty-third segment, the bud 
being broader than the nurse-stock, furnished with a pigment speck (ocular ?) at the base 
of the dorsal cirrus, and containing either ova of a violet hue or sperms. The bud is 
thrown off without a head but with long swimming-bristles. The author notes the 
degeneration of the muscular walls of the body, the shrinking of the alimentary canal, 

1 All these are but stages of one process. 

2 ' Mitt. Zool. Stat. Neap./ xiii, p. 486. 

3 ' Annel "Challenger," ' p. 195, pi. xxx ; figs. 5 and 6; pi. xxxiii, fig. 6 ; and pi. xv a, figs. 16, 17. 

4 Op. cit. 



200 SYLLIS CORNUTA. 

the differences in section of the ordinary and thefswimming-bristles, and the structure of 
the pigment-spots and glands in the sexual bud. 

In 1893 De St. Joseph observed that he had found specimens of thirty-eight and 
forty-six segments, with orange segmental organs, and filled with rose-coloured sperms, 
and acephalous buds of twenty-eight and seventeen segments loaded with sperms and 
having swimming-bristles. A spot of violet pigment occurred at the base of some of the 
dorsal cirri. He did not consider the Syllis hamata of Claparede identical with Syllis 
spongicola, Grube. 

The structure of the proventriculus of this species is described by Malaquin 1 as 
similar to that of Syllis hyalina, Grube, which is^mtermediate between the Autolyti and 
the Eusyllides. The intimate structure of the muscular columns is, however, more complex, 
for four or five transverse striations are present. 

The same author 2 describes and figures the ovaries of this species — in the lateral 
regions of the body — as having a large cgecal blood-vessel in their midst. Each is sur- 
rounded by a follicular membrane. 

In Gravier's 8. djiboutiensis, which resembles this form in general aspect, the dorsal 
cirrus is shorter, and the ventral still more so, whilst the bifid tip to the distal end of the 
bristle is noteworthy. 



8. Syllis cornuta, H. Rathke, 1843. Plate LXX, figs. 22-22 b— feet ; Plate LXXIX, 

figs. 16-16 b — bristles. 

Specific Characters. — Head transversely ovoid, with four eyes in a semicircle. Palpi 
elongate ovoid, soldered at the base. Median and lateral tentacles and the tentacular cirri 
long and moniliform. Dorsal cirri in the middle of the body have 17 — 20 or more articu- 
lations. Body moderately elongated. Foot with a dorsal cirrus gently tapering from a 
point a little above the base to the tip. Setigerous region conical, bearing one or more very 
large spines, and a series of falciform bristles, the bevelled ends of the shafts being 
smooth. The upper bristles have very long narrow terminal processes, which are slightly 
bent, like a curved bistoury, in lateral view, and minutely bifid at the extremity. The 
lower bristles have shorter hooked tips spinous along the edge, the last spine being the 
secondary process beneath the hook. The ventral cirrus is sausage-shaped with a tapering 
tip extending as far outward as the setigerous region. 

Synonyms. 
1843. Syllis cornuta, H. Rathke. Beitrage z. Fauna Xorw., p. 164, Taf. vii, f. 12. 
1851. „ „ Grube. Fain. Annel., pp. 61 and 132. 

1865. „ „ Johnston. Cat. Brit. Mus., pp. 192 and 342, 

„ „ „ De Quatrefages. Annel., II, p. 23. 

1867. „ „ Malmgren. Annul. Polych., p. 43, Taf. viii, f. 45. 

1869. „ „ Mcintosh. Trans. E. S. Edin., xxv, p. 415, pi. xvi, f. 15. 

1874. „ „ Malm. Op. cit. Goteb., p. 82. 



1 'Mitt. Zool. Stat. Neap./ pp. 228 and 229. 
3 Op. cit., 1893, p. 377. 



SYLLIS CORNUTA. 20J 

1875. EMersia sexoculata, Marion and Bobretzky. Ann. Sc. Nat., 6 e ser. t. ii, p. 20. 
1879. ,, „ Marion. Ibid., t. viii, p. 18, pi. xv, f. 3. 

„ Syllis cornuta, Tanber. Annul. Danic, p. 94. 
1885. EMersia cornuta, Langerhans. Zeitsch. £. w. Zool., Bd. xl, p. 247. 

„ ,, „ Carus. Fauna Medit., p. 228. 

1890. „ sexoculata, Griard. Bull. Sc. Fr. Belg., t. xxii, p. 78. 

1902. „ cornuta, Mcintosh. Ann. Nat. Hist., ser. 7, vol. ix, pp. 297 and 298. 

1903. „ ,, idem. Mar. Invert. S. Africa, vol. iii, p. 37. 

1904. „ (EMersia) cornuta, Allen. Journ. M. B. A., n.s., vol. vii, p. 219. 

„ „ cornuta, Soulier. Trans. Inst. Zool., Univ. Montpellier, 2 e ser., Mem. 14, p. 39. 

1906. EMersia sexoculata, De St. Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 181. 



Habitat. — Dredged off the Hebrides by Dr. Grwyn Jeffreys in June and July, 1866, 
and in 100 fathoms, St. Magnus Bay, Shetland, in the same months in 1867 ; deep-sea 
fishing off St. Andrews (E. and R. M.) ; off Bundoran, Donegal, and Berehaven, Ireland 
(R. I. A. Exped.). By < Porcupine,' 1869, 20—25 fathoms, No. 18. 

Mediterranean, shores of France. Finmark and Norway. South Africa. Spitzbergen. 
Madeira. 

Head transversely ovoid with the four eyes in a large semicircle, or as Rathke puts it 
in a single series. Palpi long, soldered at the base. Median and lateral tentacles long 
and monilified. Tentacular cirri likewise long. 

Body about an inch long and having about 100 segments. The colour is yellowish- 
white. 

The foot (Plate LXX, figs. 22, 22 a, and 22 b) in the anterior region bears dorsally a 
longer cirrus than posteriorly, the former having about seventeen articulations, the latter 
about twelve. In both the organ is tapered. The setigerous lobe is a truncated cone, 
bifid at the tip, with the spines (one is large) piercing the upper angle. Of these about 
four are present in front and two posteriorly. The bristles (Plate LXXIX, figs. 16, 16 a, 
and 16 b) are characteristic. Whilst the shafts correspond with the type the tips of the 
dorsal series quite differ, being elongate, tapered, slightly curved, and ending in a minutely 
bifid tip. There are three or four of these in each foot. The edge is minutely spinous. 
In the posterior feet these processes are shorter and stouter. The other bristles have a short 
terminal piece, minutely bidentate, and with a spinous edge (Plate LXXIX, figs. 16 a and 
16 b). Slight differences exist between those procured in Donegal Bay (fig. 16 b) and those 
from Shetland (fig. 16), the bifid condition of the bristles in the latter being most distinctly 
marked, as it also is in the posterior bristles of the southern forms. 

The inferior lobe is sausage-shaped anteriorly, and has a downward curve, its con- 
vexity being dorsal. The tip extends as far outward as the fleshy part of the setigerous 
region. In the posterior feet the lobe is more pointed. 

The characteristically large spines of this species are a feature of moment, especially 
towards the posterior region, where one is so prominent. In certain varieties the 
elongation of the ventral cirrus is noteworthy. . 

It is doubtful whether the form mentioned by Dr. Johnston was this species. In Dr. 

83 



202 SYLLIS COLLINGSIL 

Malmgren's figure 1 the dorsal cirrus is furnished with double the number of segments, 
and the bristles are not characterized by the minutely bidentate apex ; moreover, only a 
profile or linear view of the elongated kind is given. It is not always easy to make out 
the bidentate condition of the bristles, and there can be little doubt that the structure of 
the bristles, especially in the earlier papers, was imperfectly known. Indeed, it requires 
special care to observe that the bifid condition is simply due to the well-developed nature 
of the last spine of the row along the edge of the terminal blade. The condition is less 
distinct, for instance, than that shown in the figure of Marenzeller. 

The Norwegian examples collected by Canon Norman are smaller than those pro- 
cured by the ' Porcupine,' and the characters of the bristles more minute. 

This species has certain relationships with Syllis prolifera, Krohn (S. lussinensis, 
G-rube, and 8. Armandi, Claparede). The presence of the simple bristle in the anterior 
(twenty-one) segments of Krohn's species, however, differentiates it, but further 
investigation may clear up doubts. 

There seems to be variety as to the number of segments in the dorsal cirri. 

De Quatrefages (1865) constituted a genus Mlersia for the Syllis sexoculata of 
Ehlers from Martin sica, which has six eyes, three tentacles, and four tentacular cirri, with 
long articulated dorsal cirri, and which may fairly be included under Rathke's species. 

Giard (1890) is of opinion that the Typosyllis altemosetosa of De St. Joseph, is a 
variety of this form. 

De St. Joseph (1906) is inclined to consider the Syllis sexoculata of Ehlers as this 
form, and there is much to be said in favour of such a view. 



8 a. Syllis coenuta (?) var. Collingsii. 2 

Habitat — Caught in the surface tow-net off the Island of Sark, July, 1869, by 
Mrs. Collings, of Sark. 

Head with the general aspect of Syllis cormda ; eyes comparatively small, arranged 
in a semicircle, that is, the anterior and larger pair lying obliquely to the exterior of the 
posterior. In an example two small posterior eyes were present on one side. The 
tentacles and tentacular cirri are moniliform, but shorter than in S. armillaris. 

Body little tapered anteriorly, more so posteriorly, where it terminates in two short 
cirri, which are larger than those adjacent, and of nine or ten segments. The longest 
dorsal cirri are in front. 

The foot (Plate LXX, fig. 16) presents dorsally a moniliform tapering cirrus, which 
according to its position has its segments ranging from ten to twenty. The setigerous lobe 
forms an elongated cone with a bevelled tip, the two spines passing to the upper border. 
The bristles (Plate LXXIX, fig. 10) have slender shafts, curved, dilated, and bevelled at 
the tip, while the terminal piece is comparatively long and has a minutely bifid tip and 
the edge finely spinous. The ventral cirrus is lanceolate and elongate, with a slight 
swelling above its base. 

1 < Annul. Polych./ p. 43, Taf. viii, fig. 45 c. 

2 Named after Mrs. Collings, of Sark, the kind friend of every zoologist landing on the island. 



SYLLIS GRACILIS. 203 

The somewhat long terminal process of the bristles and the presence of a minutely 
bifid tip in this form, which may be a young example, would show its relationship with 
Syllis comiita, H. R., but the upper bristles did not attain the length of the ordinary 
examples of that species, a feature, however, which may be due to age. Moreover, the 
prominent spines of the typical species were not seen. It has been thought desirable, 
however, for the present to regard it as a variety of Syllis cornuta. 



9. Syllis gracilis, Grube, 1840. Plate LI, fig. 3; Plate LXX, figs. 25 and 25 a — foot; 

Plate LXXIX, figs. 17-17 c— bristles. 

Specific Characters. — Head broader than long, with four small black eyes, the anterior 
pair the larger and wider apart. Palpi slightly tapered and transversely striated. Median 
tentacle has about eighteen segments, the lateral twelve, and the tentacular cirri eighteen. 
Body of numerous segments (eighty or more) and of a brownish hue, paler anteriorly, the 
intestine giving a dark brownish colour to the central region from the proventriculus 
backward. The buds are purplish. The anterior cirri have from twenty to twenty-five 
segments, but they soon diminish to fifteen or sixteen. 

Two anal cirri of moderate length (thirteen segments) with a distinct median papilla 
ventrally. The anterior border of the proventriculus is at the twelfth segment, and it 
reaches to the seventeenth. The foot in the anterior third has a dorsal cirrus of sixteen 
segments, tapering towards the tip. Setigerous region forms a short cone. The enlarged 
distal ends of the shafts of the bristles are smooth. The distal piece is long, spinous on 
the edge, terminating in a well-marked hook with a minute tooth beneath. The distal 
pieces diminish in length from above downward. The ventral cirrus is long and lanceolate, 
projecting considerably beyond the tip of the setigerous region. The middle region of 
the body has two greatly-developed bifid bristles, the strongest of which has no inter- 
mediate teeth in the fork, whilst the other may have an intermediate spike or two. In the 
posterior third a tendency to the growth of a separate spur at the tip appears in one of 
these. In the posterior fourth a single strong bristle occurs, the end of the shaft being 
bevelled, and a stout terminal piece appended, its tip usually being indistinct. It is 
flanked on each side by a bristle of the type seen in front, viz., with a long terminal piece 
and a bifid tip. The stout bristles gradually disappear so that in the last twelve segments 
or thereabout only the falciform bristles remain. 

Synonyms. 

Grube. Akt. Echin., etc., p. 77. 

Delle Chiaje. Descriz., iii, p. 95; v, p. 101. 

Grube. Fam. Annel., pp. 61 and 132. 

Claparede. G-lanures, p. 77, pi. v, f. 3. 

De Quatrefages. Annel., II, pp. 24 and 644. 

Claparede. Annel. Nap., p. 193, pi. xv, f. 3. 

Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., t. ii, p. &3. 

Webster. Trans. Albany Inst., ix, p. 17, f. 6. 



1840. i 


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1851. 


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1864. 


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1890. 


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204 SYLLIS GRACILIS. 

Webster. 32nd Rep. N. Y. Mus. Nat. Hist., p. 109. 

Langerhans. Canar. Annel., Nov. Acta Leop.-Car., p. 105. 

Cams. Fauna Medit., p. 228. 

De St. Joseph. Ann. Sc. Nat. 7 e ser. i, p. 158. 

Malaquin. Annel. Boulonn, p. 35. 

De St. Joseph. Ann. Sc. Nat., 7 e ser., xx, p. 190, pi. xi, f. 4—7. 

G-ravier. Nouv. Arch. Mus., 4 e ser., ii, p. 150, pi. ix, f. 4 — 6. 

Mesnil. Comp. Eend. de Soc. Biol., t. liii, p. 268. 

Soulier. Trav. Inst. Zool. Mont., 2 e ser., Mem. 14, p. 31. 
{Ehlersia) gracilis, Allen. Journ. M. B. A., n.s., vol. vii, p. 219. 
gracilis, Willey. Kept. Ceylon Pearl F., iv, p. 269. 

De St. Joseph. Ann. Sc. Nat. 9 e ser. t. iii, p. 182. 

Habitat. — Under stones in the Gouliot caves, Sark, July, 1868. Common elsewhere 
in the Channel Islands. Deep-sea fishing, St. Andrews (E. M.). Millbay Channel, 
Plymouth Sound (Allen). 

Marseilles (Marion and Bobretzky). Port-Vendres (Claparede). Shores of France 
(De St. Joseph). Naples (Claparede). Canaries (Langerhans). Black Sea (Czerniavsky). 
Red Sea (Gravier). On oysters at low water on the Virginian coast (Webster). Ceylon 
(Willey). 

Head broader than long, with four small black eyes, the anterior pair the wider apart. 
The palpi are slightly tapered and transversely striated, as Claparede states, and this is 
probably due to muscular bands. The median tentacle has about eighteen segments, the 
lateral twelve, the tentacular cirri about eighteen, and the dorsal cirri of the anterior region 
twenty to twenty-five, those of the posterior about nine (Claparede gives twelve to fifteen). 

Body has 150 segments, twenty-eight belonging to a sexual stolon of nearly a mille- 
metre in diameter (Claparede). The colour is brownish, whereas that of the bud is 
purplish. The brownish hue is due to the presence of fine, interrupted strise on the 
dorsum, best marked in the anterior twenty-five segments. De St. Joseph gives the 
proventriculus seventy-four rows of points. 

The foot (Plate LXX, fig. 25) anteriorly has a moderately elongated dorsal cirrus, 
tapering towards the extremity, moniliform throughout and consisting of about sixteen 
segments. The setigerous region forms a short cone, the spines passing to a papilla at 
the upper angle. The bristles are stout, translucent, curved below the dilated end of 
the shaft, which is bevelled to an acute point but not spinous (Plate LXXIX, fig. 17). 
The distal piece is long, spinous on the edge, and ends in a well-marked hook with a minute 
tooth beneath, as in Pionosyllis. The terminal piece diminishes in length from above 
(fig. 17) downward (figs. 17 a, 17a 1 ). The ventral cirrus is long and lanceolate, pro- 
jecting considerably beyond the tip of the setigerous region. 

The posterior feet differ from the foregoing in the shorter and thicker dorsal cirrus, 
which has only about nine segments, in the shorter cone of the setigerous region which 
superiorly has three strong spines which do not pierce the surface superiorly, and two 
strong bristles (Plate LXXIX, figs. 17 b, 17 c) with a slightly bifid tip, which may be a 
modification of the dilated end of the shaft in the ordinary form. Their great strength 
points to a special function— possibly amongst the Balani so common in the Gouliot caves. 



SYLLIS GRACILIS. 205 

The ventral lobe is also shorter and distinctly tongue-shaped, being narrower at the base 
and tip than in the centre. 

The middle region of the body is characterized by the great development of the strong 
bifid bristles, 1 two of which are present in each foot (Plate LXX, fig. 25 a), the jointed 
bristles having disappeared. The strongest of these have no intermediate teeth in the 
fork, others present a single oblique spike, while a few have more than one point. Many 
have no distinct points though the fork is slightly rough. These two strong bristles 
continue to the posterior third, where a tendency to the development of a separate spur- 
like process, instead of a limb of the fork, occurs. This is generally better marked in one 
than in the other (of the two in each foot) . 

In the posterior feet (about the posterior fourth) a single strong bristle occurs in the 
centre, the end of its shaft is bevelled, and a stout terminal piece is appended, the tip of 
which is generally indistinct. It is flanked on each side by nine bristles of the type of 
those in front, viz., with a long terminal piece bifid at the tip. The stout bristles of the 
feet behind the foregoing segments gradually disappear, the first and then the second, 
until only jointed bristles remain, one of which, however, is stouter than the others, 
and has a shorter terminal piece. Finally in the last twelve or more segments only 
the slender bristles with the elongated and bifid terminal pieces are found. 

In an example from Plymouth eight segments, and the cirri of the tail, were in 
process of reproduction. 

Claparede (1864) 2 describes the spines as having a button-like tip as in $. Armandi. 
He found a papilla between the caudal cirri, as in certain other species, and mentioned 
that the blood, usually colourless in Syllids, is in this species of a beautiful rose colour. 

This author 3 mentions that the term Syllis gracilis was first used by Delle Chiaje, but 
as the species is indeterminable Grube's name stands. The same term was adopted by 
Schmarda for a species from the Cape after Grube had employed it. He drew special 
attention to the glands with coiled contents near the dorsal cirri, and to the coiled 
structures in the latter. 

Langerhans (1879) adds an interesting note on the arrangement of the bristles 
(simple, mixed, and compound) in the various segments of this species, but it is not con- 
stant in the several examples. He thought the stolons had the form of Ioida. 

De St. Joseph (1886) found his example at Dinard in the midst of Cynthia glomerata. 
He describes the proboscis as elongated, stretching from the second to the eleventh 
segment, the conical tooth in front with a poison canal. The proventriculus extends from 
the twelfth to the fifteenth segment. He criticises Czerniavsky, who made four species 
out of it in the 'Fauna of the Black Sea ' (Mare Ponticum). In his later note (1895) he 
adds further information. He observes that the first twenty-seven segments have bristles 
with a long bidentate terminal piece, whereas from the twenty-eighth to the forty-fourth 
segment these are replaced by two stout bristles with a forked point (ypsiloide), with 

1 Claparede states that the ypsiloid bristles commence from the nineteenth to the twenty-sixth 
segment. ( Annel. Nap./ p. 194. 

2 This author thinks the species approaches Syllis oblong a, Keferstein, and S. tigrina, Rathke,, 
though they differ in palpi and bristles. ( Beobach./ pp. 52 and 82. 

3 ' Annel. Nap./ p. 193, etc. 



206 SYLLIS BUSKIL 

three minute points between. In segments 45 — 53 stout bristles with, a simple terminal 
piece are present. In segments 54 — 62 are bristles like those in front (with long 
slender terminal pieces), besides two simple bristles, the one with an entire point, the 
other with a bidentate tip. 

In large examples (5 cm. long and 125 segments) from Villerville there were in front 
twenty-seven segments with long bidentate tips to the bristles ; the next fifty-six segments 
with two ypsiloide bristles ; the twenty-five following with one ypsiloide bristle, and one 
bristle with a short unidentate tip ; and the last sixteen with a long bidentate tip to the 
bristles, as in the anterior segments. He also met with an example in which regeneration 
had taken place, but to a less degree than in Mesnil's example. 

He dredged 1 an epitokous female of 32 segments, somewhat resembling his Syllis 
altemosetosa, at Cannes. It had a rounded head, with four enormous eyes, and three 
slightly articulated tentacles. The first segment bears a dorsal cirrus of ten articulations, 
a ypsiloide bristle, a bidentate one, and a ventral cirrus. The bristles vary posteriorly, 
and swimming-bristles with hooked points were present from the second to the thirtieth 
segment. A median papilla occurred between the anal cirri. 

Mesnil 2 relates a case of regeneration of the head with four eyes furnished with 
lenses, and the anterior region with complete pharynx, proventriculus, and other organs. 
The posterior part of the new region, that is, where it joined the old, presented very 
indistinct segmentation. He notes that about the seventh regenerated segment anteriorly 
a simple bristle, curved and bifid at the extremity, occurred, whereas he had not found 
such in the ordinary or unregenerated example. 

Gravier (1901) pointed out the presence of a third kind of bristle, viz., one with a 
spike on the dilated end of its stout shaft, and a short distal piece. Such may indicate a 
stage in the modification of the stronger forms devoid of the terminal piece. This author 
also describes and figures a female stolon of thirty-five segments and 10 mm. in length. In 
this the usual conditions are found. The long natatory bristles are tapered towards the 
extremity and hooked. The stout ventral bristles show a tooth between the bifid ends 
and minute spikes in the hollow. 

A female bud, with four enormous eyes, and greyish eggs in the body-cavity, has 
more recently (1906) been mentioned by De St. Joseph. 



10. Syllis Buskit, 3 n. s. Plate LXX, fig. 23— foot ; Plate LXXIX, figs. 18 and 18 a— 

bristles. 

Specific Characters. — Head apparently typical and with large palpi. Eyes had dis- 
appeared in the preparation. Tentacles and cirri articulated and of considerable' length. 
Body of comparatively large size, 1-g- in. long, the general aspect being that of Syllis 
armillaris. Foot presenting dorsally a comparatively long articulated cirrus of 24 — 28 

1 ' Ann. Sc. Nat./ 9 e ser., t. in, p. 183 (1906). 
3 < Compt. rend. Soc. Biol./ t. liii, p. 269 (1901). 

8 Named after Professor G-. Busk, who encouraged others, and also did much able work in the 
Polyzoa and other groups. 



SYLLIS ABYSSICOLA. 207 

or more segments. Setigerous region conical. Bristles strong, translucent, with 
indications of serrations on the distal dilatation of the shaft. Terminal piece of 
moderate length, with a simple hook at the end, and just below it the spinous edge ends 
in a somewhat stronger spike, so that a pseudo-bifid aspect is produced. Ventral cirrus 
elongate ovate or broadly lanceolate, and does not reach the tip of the setigerous region. 

Habitat. — Under stones between tide-marks, at Guernsey and Herm. 

Head apparently typical, with large frontal palpi, but the eyes had disappeared before 
a critical examination was made. 

Body of comparatively large size and 1-J- in. in length, the general aspect being that 
of Syllis armillaris. The tail in all is incomplete. 

The foot (Plate LXX, fig. 23) presents dorsally a comparatively long articulated cirrus, 
having twenty-four to twenty-eight or more segments. The setigerous region is conical, with 
the spines piercing the upper angle. The bristles (Plate LXXIX, figs. 18, 18 a) are strong, 
translucent, and with the usual curve towards the dilated end of the shaft. The con- 
vexity of the dilatation has indications of serrations. The terminal piece is of moderate 
length with a hook at the tip, and just below it another minute tooth at the end of the 
spinous edge. The latter tooth is easily overlooked. 

The ventral lobe is elongate-ovate or broadly lanceolate, and does not reach the tip of 
the setigerous region. It is somewhat diminished posteriorly. 

Posteriorly the body has many large ova, and a developing tuft of simple swimming- 
bristles appears between the cirrus and the setigerous region. A bud was thus forming. 
In this region the cirrus is shorter, about a dozen articulations only being present. 

This species somewhat approaches Tyjpanosyllis zebra, but is readily distinguished 
from it by the colouration, and the structure of the foot and bristles, the terminal pieces 
being shorter and broader proportionately, and the secondary process beneath the terminal 
hook is closer to the latter and more minute. 



11. Syllis abyssicola, Elders, 1875. Fig. 55. 

Specific Characters. — Head somewhat triangular, bluntly pointed anteriorly, with 
elongate-ovoid palps. Lateral tentacles of fourteen segments; median of seventeen segments. 
No eyes. Body deep yellow : first segment very short, with a pair of cirri on each side, 
the dorsal with thirteen segments. Proboscis has ten papillae, and a single tooth behind 
them. Foot has dorsally a cirrus of nine to eleven segments ; setigerous region cylindrical ; 
two or three of the upper bristles have long tapering tips with a simple hook. The lower 
bristles have much shorter tips. Ventral cirrus is somewhat cylindrical and pointed. 

Synonyms. 

1875. Syllis abyssicola, Ehlers. Zeitsch. f. w. Zool., Bd. xxv, p. 45, Taf. ii, f. 18—20. 
1879. Ehlersia „ Langerhans. Ibid., Bd. xxxii, p. 237. 

Habitat— Dredged at Station 30, 56° 24' X., 11° 37' W., in the < Porcupine' Expedi- 
tion of 1869, at a depth of 1380 fathoms amongst dark-coloured mud. Langerhans says 
1483 fathoms. 



208 SYLLIS BKEVICOLLIS. 

Head (fig. 55) somewhat triangular, bluntly pointed anteriorly, with distinct, elongate, 
ovoid palpi (in spirit). Paired tentacles of fourteen segments : median of seventeen seg- 
ments. JSFo eyes, a rare condition in the Syllids. 

Body incomplete posteriorly, 5*5 mm. long and 0*5 mm. in breadth ; segments thirty, 
and of a deep yellow colour. First segment very short, apparently overlapped by the 
head, bearing on each side a pair of cirri, the dorsal having thirteen segments and being 
somewhat longer than the ventral. 

The proboscis has ten soft papillae, and behind them a single tooth. It extends to 
the ninth segment, the proventriculus being in the next seven segments. 

The foot has dorsallya cirrus of nine to eleven segments, thus being comparatively short, 
a cylindrical setigerous region bearing bristles, two or three of which dorsally have 
very long tapering tips with a slight hook, and a group with much shorter tips, also 
simply hooked ventrally. The ventral cirrus is simple and more or less cylindrical and 
pointed at the tip (Ehlers). 




Fig. 55. — Head of Syllis abys&icola, Ehlers. (After Ehlers.) 

Ehlers considers that in the absence of the eyes this species approaches Syllis 
variegata, Grube. 1 



12. Syllts beevicollts, Ehlers, 1875. 

Specific Characters. — Head resembling that of Syllis armillaris, but the median 
tentacle is longer. The lateral tentacles have fifteen segments. Body normal. First 
segment with two tentacular cirri, the dorsal very long. Proboscis with a single tooth, 
and reaching to the thirteenth segment. Foot has a long dorsal cirrus of forty to fifty 
segments. Setigerous lobe short and blunt ; bristles have a somewhat long terminal piece 
with a simple hook. Ventral cirrus short and clavate, a little longer than the setigerous 
lobe. 

Synonyms. 

1875. Syllis brevicollis, Ehlers. Zeitsch. f. w. Zool., Bd. xxv, p. 44, Taf. ii, f. 16 and 17. 
1879. Typosyllis (Syllis) brevicollis, Langerhans. Ibid., Bd. xxxii, p. 533. 

Habitat. — Dredged on the 15th July in the ( Porcupine ' Expedition of 1869 at Station 
47, 59° 14' 1ST., 7° 18' W., in 650 fathoms (Langerhans says 1380 fathoms), on Holtenia 
ground with mud and Grlobigernia-ooze, mingled with sand. 

1 "Beschrieb. neuer oder wenig gekaunter Annel.," ' Arch. f. Naturges./ Jahrg. 26 (1860), p. 85, 
Taf. iii, fig. 6. 



AUTOLYTUS. 209 

Head somewhat like that of 8. armillaris, except that the median tentacle is longer 
(forty-four segments) and the palpi seem to be closer. The paired tentacles have fifteen 
segments. 

The incomplete body has sixty-four segments and is 18 mm. long, the breadth being 
1*5 mm. The colour of the dorsum (in the spirit-preparation) is brownish-yellow. The 
first segment bears two tentacular cirri, the dorsal very long. The proboscis (pharynx) 
has a single tooth and reaches to the thirteenth segment. 

The foot has a long dorsal cirrus of forty or fifty segments. The setigerous region is 
short and blunt — carrying bristles with a somewhat long terminal piece and a simple hook 
at the tip, and thus resembling those of 8. armillaris. The ventral cirrus is short and 
clavate, a little longer than the setigerous region (Ehlers). 

How far the cirri vary in length is still an open question, so that the species may yet 
be linked on to others. 



Group IV.— AUTOLYTEA. 

Syllideans deprived of ventral cirri. Palpi little developed, soldered on the ventral 
surface of the cephalic lobes. Cirri not articulated. Reproduction generally by stolons 
(schizogamy), rarely direct (epigamy, Malaquin). 



Genus LV. — Autolytus, Grube, 1851. 

Animal surculare. — Head rounded in front; palpi obsolete (coalescent), but separated 
by a line ventrally. Proboscis sinuous, with a crown of teeth. Stomach little developed. 
Tentacles three, smooth, and little tapered. Two tentacular cirri on each side of the 
buccal segment. Dorsal cirri of the first two segments considerably longer than the 
others. Ventral cirri absent. Falcate bristles with short bifid tips, the lower point or 
hook being the larger. The sexual generation budded from the foregoing shows dissimilar 
males and females (Polybostrichus and Sacconereis), but both always have a median 
tentacle, and the paired tentacles are often present. 

In a transverse section of a female (Fig. 56) the muscular system is as powerfully 
developed as in Syllis, the dorsal longitudinal muscles widening from the median raphe 
to the inferior border. The ventral longitudinal muscles are curved, and separated by 
the large nerve-cords. The large ova lie in the cavities of the feet and in the peri- 
visceral space around the gut, which in the example had thick glandular walls. 

The type of circulation in this group is simple, a dorsal and a ventral vessel united 
only in the anterior region, as seen in Autolytus and Myrianicla, are present. The nurse- 
stock in the latter has a dorsal vessel fixed to the wall of the body by the mesentery, and 
in a fold of the intestine, though quite free from it (the intestine). The ventral vessel is 
intimately united with the intestine. In the anterior region the vessels are independent 
of the proboscis dorsally and ventrally. The dorsal vessel is only contractile anteriorly 
(Malaquin) . 

84 



210 



AUTOLYTUS. 



Grube, in his ' Familien der Anneliden ' (1851), made a new Family, the Amytidea, 
to include Polybostrichus longisetosus of (Ersted, Nereis prismatica of 0. F. Muller, Nereis 
bifrons of 0. F. Midler, Photocharis cirrigera of Bhrenberg, Nereis corniculata of 0. F. 
Muller, and Nereis clavicomis of Sars — all sexual forms connected with Autolytus prolifer 
and its immediate allies. 

Malaquin (1893) takes Autolytus Edwardsi, De St. Joseph, as the type of reproduction 
in the group, and describes various phases in separate individuals. In the first instance 
an adult develops genital glands in the posterior segments, this region subsequently 
separating by fission and having an independent existence for a time. A segment in front 
of the genital region, between the fortieth and forty-fifth segment, is thickened and becomes 
the formative segment — the head of the bud being developed from it and subsequent 
additional segments in the bud. In some cases a number of segments in front of the head 
of the bud bear genital glands. After the separation of the bud the nurse-stock develops 
new segments. 

In the second case the stolons of twenty to twenty-two segments have in front of them 




Fig. 56. — Transverse section of Autolytus prolifer to show the arrangement of the muscles. Letters as "before. 



a certain number of segments in process of development. The bud separates as usual, 
regeneration of the posterior segments of the nurse-stock ensuing. 

In the third case the genital segments posteriorly are separated as a bud with head 
and appendages. A formative zone in front of the bud actively develops segments which 
have genital products in front of them and the bud and its genital organs behind. 

In the fourth case several buds, generally males, are attached to the posterior end 
of the nurse-stock, the last separating as a bud as in the case of Myrianida. It is an 
example of rapid stolonization. 

The fifth case resembles the first and second. In a nurse-stock of forty segments 
the last fifteen, viz., from the twenty-fifth to the fortieth, are observed to have ovaries, 
a bud of four segments with cirri and small bristles, a formative zone, and a pygidium, 
the head being at the forty-first setigerous segment of the stock. 

In the sixth case the condition is the same as in the first, only the head of the bud is 
between the twenty-fifth and thirtieth segments. 



AUTOLYTTTS PICTITS. 211 

In the seventh case a nurse-stock of twenty-eight segments bears a chain of stolons 
from one to seven in number. Malaquin is of opinion that where the chain of buds is 
smaller than the nurse-stock it is a case of budding, whereas when the stolon is larger or 
as large as the nurse-stock it is a case of fission. 

Langerhans makes this his third tribe, and divides it into seven sub-sections, viz., 
flurysyllis, Ehlers; Anoplosyllis, Claparede; Heterosyllis, Claparede; Autolytus, Grrube ; 
Procerxa, Ehlers ; Myrianida, Aud. and Edwards ; and Virchoivia, Langerhans. 

The British forms fall under only three of these, viz., Autolytus, Procersea, and 
Myrianida; and if, as appears to be right, the first two ought to be conjoined, then only 
two genera are represented in our fauna. 

Langerhans distinguishes Procersea from Autolytus by the presence of elongated cirri 
on the first, second, and third segments of the body, the latter {Autolytus) only having 
these on the first and second. Moreover, he points out that the pharynx has twenty teeth 
(papillae) — ten larger and ten smaller. 

De St. Joseph included Procersea and Stephanosyllis under Autolytus, and his 
description of the genus corresponded for the most part with that now given. He also 
noted that one group, Autoly tides, Malaquin, has an unarmed proboscis. 1 

As Calvin Mensch 2 points out, only one bud is borne at a time in this genus and in 
Autolytus cornutus of Agassiz, who demonstrated the relationship of- (Ersted's Polybostrichus 
(1843) and J. Muller's Sacconereis (1853). 



1. Autolytus pictus, Ehlers, 1864. Plate XL VII, fig. 1— male bud; Plate XLIX, fig. 8— 
nurse-stock; Plate LXXI, fig. 2— foot; Plate LXXIX, fig. 21— bristle. 

Specific Characters. — Head somewhat small, rounded in front, long diameter antero- 
posterior. Eyes four, the anterior pair the wider apart, in some connate, and both have 
lenses. Two lateral and a longer and thicker median tentacle, all of a madder-brown hue. 
The tentacular cirri are less elongated but of the same colour. Body about one inch long, 
and of about 100 segments ; slightly narrowed anteriorly and more distinctly so posteriorly, 
where it terminates in two short, flattened, and curved cirri about twice the length of the 
last segment. Dorsum marked by a pale central band with numerous and regular transverse 
spurs, which cut the brown (sepia) pigment into oblong spaces. Beneath and external to 
the latter is a pale lateral belt into which the transverse spurs run. Just above the feet 
a dark brown belt passes from end to end, intersected here and there, opposite the pale 
transverse spurs, by narrow pale lines. First twelve or thirteen segments are darker and 
have the ground colour paler than the rest. Ventral surface pale or flesh-coloured. 
Foot carries a short dorsal cirrus (except anteriorly) which shows slight crenations when 
coiled. Setigerous process short and truncated, with comparatively short bristles, the 
shaft of which is stout, enlarged distally, bevelled at the edges, and spinous. Terminal 
piece short and broad, with a bifid tip, the lower hook being the larger. 

i < Ann. Sc. Nat./ 7 e ser., I, 1887. 

2 " On the Life-History of Autolytus cornutus and Alternation of Generations in Annelids/' 
< Americ. Nat./ vol. xxxiv, p. 165 (1900). 



212 AUTOLYTUS PICTTJS. 

Synonyms. 

1864. Stephanosyllis scapidaris, Claparede. Glanures, p. 107, pi. vii, f. 5. 
,, Procersea picta, Ehlers. Borstenw., I, p. 256. 

1865. Myranida ,, De Quatrefages. Annel., II, p. 63. 

1868. Procersea „ Claparede. Annel Nap., p. 219. 

1869. Autolytus pictus, Mcintosh. Trans. R. S. Edin., t. xxv, p. 415, pi. xv, f. 11. 

1874. Autolytus (Procersea) pictus, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 198. 

1875. „ „ „ idem. Invert, and Fishes St. Andrews, p. 121. 

1879. Procersea picta, Langerhans. Zeitsch. f. w. Zool., Bd. xxxii, p. 577, pi. xxxii, f. 28. 

1885. „ „ Carus. Fauna Medit., p. 235. 

1887. Autolytus (Procersea) pictus, De St. Joseph. Ann. S.c. Nat., 7 e ser., t. i, p. 222, pi. xi, 

f. 100—105. 
1890. „ „ „ Malaquin. Annel. Boulonn., p. 39. 

1904. „ „ „ Allen. Journ. M. B. A., n.s., vol. vii, p. 220. 

1906. „ „ „ De St. Joseph. Ann. Sc. Nat. 9 e ser., t. iii, p. 186. 

Habitat. — Under stones in rock-pools and near low water, St. Andrews ; under a stone 
in a rock-pool at Paible, North Uist ; on a zoophyte off Sark (Mrs. Collings) ; Plymouth 
(Allen). It is probably very widely distributed in Britain, and is not uncommon at St. 
Andrews, where it was first found by my sister, who made the coloured drawings. 

Abroad it occurs in the Mediterranean (Carus) and the White Sea; the Adriatic 
(Ehlers); Cannes (De St. Joseph); Madeira (Langerhans). 

Head somewhat small, rounded in front, the longer diameter being antero-posterior. 
Byes two on each side, the anterior pair the wider apart, but sometimes they are connate, 
both the anterior and posterior being furnished with lenses, the former, like the eye, being 
the larger, and De St. Joseph states that they lie in front, while the lenses of the posterior 
are behind. Two lateral and a longer and thicker median tentacle, all of a deep madder- 
brown colour, the median being darkest, though its ceratophore is paler. Their surface is 
supplied with numerous palpocils. The ocular region is richly ciliated, and the dark por- 
tion (probably De St. Joseph's fused epaulettes), at the sides of the succeeding ring, is 
also ciliated. A dark patch of deep brown is placed behind the median tentacle, and 
from the latter two diverging pale lines proceed backward. The tentacular cirri are 
less elongated than the tentacles, but have the same madder-brown colour. 

Body about an inch in length, and with upwards of 100 segments, slightly narrowed 
anteriorly, and more distinctly so posteriorly, where it terminates in two short, flattened, 
and curved cirri — ■ about twice the length of the last segment. The dorsum is prettily marked 
by a pale central band with numerous and regular transverse spurs which cut the sepia- 
brown pigment into oblong spaces. The spaces, however, are not solidly pigmented, but 
are striated by fine pale lines and grains (under a lens). By transmitted light some of the 
dark pigment appeared of a fine purplish hue. On each side below (and external to) these 
striated oblongs is a pale lateral band, into which the transverse spurs run. Just above 
the feet a dark brown belt passes from end to end, intersected here and there — opposite 
the pale transverse spurs — by narrow pale lines. The first twelve or thirteen segments 
are darker and the ground-colour is paler than in the rest. The lateral pale line within 
the feet is also less distinct in this region, but behind it becomes boldly marked though 



AUTOLYTUS PICTUS. 213 

less white. The ventral surface is pale or flesh-coloured. The tints are retained for 
a long time (eight months) in spirit. 

The proventriculus, according to De St. Joseph, has from forty-eight to sixty rows of 
brown points, and occurs in segments 10 and 11. The proboscis, he states, is furnished 
with ten large teeth, alternating with ten smaller. He also found in the sub-cuticular 
tissue greyish plates with two refringent hemispherical concretions. 

The foot (Plate LXXI, fig. 2) carries the dorsal cirrus, which in the fourth 
segment exceeds in length the tentacles and tentacular cirri. They are short posteriorly. 
All have palpocils, and show slight folds and crenations when coiled. As in Amblyosyllis 
they are stretched out on irritation. When falling through the water the long coiled 
processes in front bear the weight on touching the bottom. The setigerous process is 
somewhat short and truncated at the tip, though it varies in outline according to the 
development of the reproductive elements. It has several spines, which point above the 
bristles. The latter (Plate LXXIX, fig. 21) are comparatively short, with a stout, slightly 
curved shaft, the distal end of which is enlarged, and both sides of the bevelled region 
have minute spines, largest at the apex. The distal piece is short and rather broad, and 
the tip is bifid, the lower hook being the larger. Towards the tail only a single conspicuous 
spine occurs in each bristle-bundle, and finally a larger and a smaller spine alone exist. 
The terminal processes of the bristles vary much in length, being very short in the 
anterior region, then lengthening, and again becoming short as the bristles diminish in 
size towards the tail. The ventral cirrus seems to be fused with the setigerous lobe. 

In the thirty-fourth segment De St. Joseph met with a single dorsal bristle, " a article 
en alene," as figured by Langerhans, who found it on the fourth foot. 

He also found individuals bearing eggs, as Ehlers and Claparede did, but he thinks 
Claparede was hasty in concluding them to be Autolyti without alternation of genera- 
tions. He observed eggs in the thirteenth segment and in the fifteenth segment (with a 
developing head of a bud). The buds have from forty-five to' fifty-eight segments. In 
the males (Polybostriehus) the first region has six segments, the second thirty-one segments, 
and the third twenty-two segments. The first region has two spines in each foot ; in the 
second the compound bristles are accompanied by a strong spine, except the two first 
segments, which have two, and the natatory bristles are accompanied by two hooked 
bristles ; in the third region there is only one spine. 

No proboscis or proventriculus occurs in the buds, and the intestine is rudimentary. 

Malaquin observes that the first three setigerous segments (devoid of swimming- 
bristles) carry the male elements. The rest have no genital glands, but have long 
feet and swimming-bristles. The latter have specially-developed muscles — two superior 
and two inferior — for moving them. 

Reproduction. — The male buds (Plate XL VII, fig. 3) of this form are procured in May, 
and are composed of three regions. The head and two achaatous segments and all their 
appendages are pale. Of the great bifid palps the large inner division is ciliated and 
tapered, the outer slender and non-ciliated. Two minute rudiments occur in the central 
region behind these. The median tentacle is long. The eyes are large, and appear to lie 
on the ventral rather than on the dorsal surface, and are bright red. The anterior pair are 
the wider apart, much (more than twice) larger than the posterior, and nearly circular, 



214 AITTOLYTITS AURANTIACA. 

while the posterior are also nearly circular. The buccal segment bears two pairs of 
tentacular cirri, the dorsal being the longer. The succeeding acha3tous segment bears 
three long tentacles — -a median and two lateral. The long anterior cirri are often coiled 
like conical screws. 

The body reaches only about half the length of the nurse-stock, and its colouration 
differs. After the first two brown squares a considerable and wider portion of the 
dorsum has a central dark brown streak interrupted by pale bands at the usual intervals, 
and with the lateral region of the feet of a granular brown. Behind this the body 
tapers towards the tail and has the ordinary markings. The tail has two short cirri. 

Each of the feet of the middle region of the body bears a long tuft of simple bristles, 
and the divisions of the feet are split deeply towards the centre of the body, the inner 
portion being coloured, so as to add to the breadth of the central belt of brown. 

Habits. — The species lives well in captivity, and, like many others, constructs a tough 
and translucent tube or case on the side of the vessel at the water-line. Some lived for 
nine months at Murthly, which is about fifty miles from the sea. 



2. Autolytus aurantiaca, Claparede, 1868. Plate LXXI, fig. 3 — foot; Plate LXXIX, 

figs. 22, 22 a — bristles. 

Specific Characters. — Head small, rounded in front with a long median and two lateral 
tentacles. Eyes four, black ; anterior pair the wider apart, whilst those on each side are 
close together. Body about f inch long, tapered towards both extremities, of an orange 
colour somewhat clouded in the middle from the intestine and its contents. Foot forms a 
short blunt process, devoid of a distinct ventral cirrus. Dor sally is the short tapered 
cirrus, then the massive setigerous region with bristles resembling A. pictus, and bifid at 
the tip of the short terminal piece. 

Synonyms. 

1868. Procersea aurantiaca, Claparede. Annel. Nap., p. 219, pi. xv, f. 1. 

1874. „ luxurians, Marenzeller. Sitz. Akad. Wiss. Wien, p. 50, Taf. vi, f. 1, and Taf. vii, f. 1. 

1875. Autolytus (Procersea) aurantiacus, Marion and Bobretzky. Ann. Sc. Nat., 6 e ser., t. ii, p. 44. 
1879. „ „ ,, Langerhans. Zeitsch. f. w. Zool., xxxii, p. 578. 

1885. „ „ „ Cams. Fanna Medit., p. 236. 

Habitat. — In a tube under a stone in a rock-pool at Paible, North Uist. It may have 
been an accidental visitor to the tube. 

Madeira, amongst littoral plants, as well as in 20 fathoms (Langerhans). Shores 
of France (Marion and Bobretzky). Mediterranean (Claparede). Adriatic (Marenzeller). 

Head small, rounded in front, with a long median and two lateral pale tentacles. 
Eyes four, black, and about equal in size, the two anterior being wide apart, while those 
on each side are close together. 

Body about f inch long, and with 70 — 100 segments (Marenzeller), tapered towards 
both extremities ; of an orange colour, somewhat clouded in the middle from the intestine 



AUTOLYTUS PROLIFER. 215 

and its contents. Langerhans describes it as diffused red, darkest at the tips of the 
tentacles and cirri. From the tenth foot backward each has a small red speck (Langerhans). 
Small specimens are pale, and Marenzeller states that twenty denticulations occur at the 
opening of the proboscis into the gut. 

The feet (Plate LXXI, fig. 3) form short, blunt processes without a distinct ventral 
lamella (cirrus). Dorsally is the pale tapered cirrus, which is comparatively short. The 
massive setigerous region bears a group of strong, slightly curved bristles (Plate LXXIX, 
figs. 22 and 22 a) resembling those of Autolytus pictus, the bevelled portion beyond the 
dilated distal end of the shaft being spinous. The distal piece is short and bifid. On 
immersion in spirit the pale cirri assumed a greenish hue. 

Further investigation may show that this form is a well-marked variety of Autolytus 
pictus, but the opportunities for critical comparison have been so few that it has been 
deemed best to leave the subject as it stands. 

Claparede points out the muscularity of the tentacles and tentacular cirri in this group 
(e. g., Autolytus aurantiaca) and the presence of cilia on their surface. 



3. Autolytus pbolifer, 0. F. Midler, 1788. Plate XL VIII, fig. 6 (An. surculare; Plate 
LX, fig. 4 — early larva, and figs. 5 and 5 a — anterior and posterior ends of free bud ; 
Plate LXXI, fig. 4— foot of bud; Plate LXXIX, figs. 23 and 23 a— bristles. 
Plate L, fig. 11 — pelagic male (Polybostrichus) . Plate XLIX, fig. 7 — pelagic 
female (?), tinted like A. pictus, with ventral egg- sac. 

Specific Characters. — Head smoothly rounded in front ; eyes placed close together on 
each side, but obliquely, so that the anterior are the wider apart. A long tapering median 
cirrus and two lateral cirri. Two tentacular cirri on each side. All these processes are 
smooth. Body about half an inch long, slightly and rather abruptly tapered anteriorly, more 
distinctly tapered posteriorly, where it ends in two slender cirri. Colour pale yellowish, 
somewhat darker in the middle from the intestine, and with numerous small whitish specks 
over the latter. Ten papillae in pharynx (Langerhans). Foot with a comparatively short 
dorsal cirrus with a distinctly marked cirrophore. The setigerous region is bluntly conical, 
with short bristles abruptly dilated at the tip of the shaft, minutely spinose, and having a 
short bifid terminal piece, the second of the two hooks being the larger. A single simple 
bristle is generally present in each foot, the tip being bent and acutely pointed, as in 
Autolytus fallax, Malmgren. 

Synonyms. 

Animal Surculare. 

1788. Nereis prolifera, 0. F. Miiller. Zool. Dame, II, p. 15, Tab. lii, f. 5—9. 
1791. „ „ Gmelin. Linn. Syst. Nat., ed. 13, i, pt. 6, p. 3120. 

1806. „ „ Turton. Linn. Syst. Nat., iv, p. 90. 

1824. „ „ Bruguiere. Tabl. Bncycl. Meth., Vers, I, p. 134, Tab. lvi, f. 12—15. 

1825. „ „ De Blainville. Diet. Sc. Nat., xxxiv, p. 440. 
1828. Nereisyllis prolifera, idem. Ibid., lvii, p. 473. 

1830. Nereis prolifera, Bosc. Vers, ed. 2, I, p. 174. 



216 ATTTOLYTITS PROLIFER. 

1834. Nereis prolifer a, Audouin and Edwards. Annel., p. 209. 

» „ „ idem. Ann. Sc Nat., xxix, p. 231. 

1845. SylUs „ Johnston. Ann. Nat. Hist., xv, p. 146, pi. ix, f. 3, 4. 

1847. „ „ Frey and Leuckart. Beitr., p. 92, etc. 

1850. Autolybns prolifer 9 G-rube. Arch. f. Naturg., p. 310. 

„ „ Krohn. Ibid., xviii, 1, p. 66. 

1851. „ prolifera, G-rube. Fam. Annel., p. 62. 

1855. „ „ idem. Arch. f. Naturges, Bd. xxi, p. 105. 

1865. Syllis „ Carrington. Annel. Southport, p. 8. 

,, „ „ Johnston. Cat. Brit. Mus., p. 192, pi. xv a, f. 3, 4. 

„ Autolytus prolifer, De Quatrefages. Annel., II, pp. 45, 63, and 87. 

,, „ Agassizii, idem. Ibid., p. 44. 

„ prolifer, A. Agassiz. Journ. Bost. Soc. Nat. Hist., vii, p. 392. 

1866. „ „ Greef. Archiv f. Naturges., Bd. xxxii, p. 363. 

1867. „ „ Malmgren. Annul. Polych., p. 32. 
9 , „ fallax {?), idem. Ibid., p. 33, Tab. vi, f. 41. 

1868. „ prolifer, idem. Ann. Nat. Hist., 4th ser., vol. i, pp. 173 and 183 (translated). 
„ „ hesperidum, Claparede. Annel. Nap., p. 216, pi. xiv, f. 1. 

1869. „ „ Langerhans. Zeitsch. f. w. Zool., t. xxxii, p. 575. 

1874. ,, prolifer, Mcintosh. Ann. Nat. Hist., ser. 4, vol. xiv, p. 198. 
j, „ „ Malm. Op. cit. Gloteb., p. 82. 

1875. „ ff Mcintosh. Invert, and Fishes St. Andrews, p. 121. 
1879. „ hesperidum, Tauber. Annul. Danic, p. 95. 

1883. „ prolifer, Levinsen. Yidensk. Meddel. Foren. Kjobenh, p. 246. 

1885. „ „ Cams. Fauna Medit., p. 235. 

1886. „ ,, Viguier. Arch. Zool. Exper., 2 e ser., iv, p. 429, pi. xxvi, f. 13 — 15, and 

pi. xxvii, f. 1. 

1887. „ „ De St. Joseph. Ann. Sc. Nat., 7 e ser., i, p. 238, pi. xii, f. 118. 

1890. ,, „ Malaquin. Annel. Boulonn., p. 41. 

1891. ,, „ Hornel. Trans. Liverp. Biol. Soc, vol. v, p. 245. 

1892. „ ,, (eyes), Andrews. Journ. Morph., p. 186. 

1903. ,, „ Mcintosh. Mar. Invert. S. Africa, vol. iii, p. 35. 

1906. „ „ De St, Joseph. Ann. Sc. Nat., 9 e ser., t. iii, p. 186. 

$ Bud. 

1788. Nereis comiculata, O. F. Miiller. Zool. Danic, II, p. 15, Tab. Hi, f. 1 — 4. J. 
1806. „ „ Turton. Linn. Syst. Nat., iv, p. 87. 

1824. „ „ Bruguiere. Tabl. Encycl. Meth., Vers, I, p. 134, Tab. lvi, f. 8—11. 

1825. „ „ De Blainville. Diet. Sc. Nat., xxxiv, p. 438. 
1842-3. Polybostrichus longisetosa, QErsted. Kroyer's Nat. Tids., p. 119. 
1851. Biplocera comiculata, G-rube. Fam. Annel., p. 64. 

,, Polybostrichus, etc, idem. Ibid., pp. 63 and 64. 

1855. Sacconereis helgolandica $ , Max Miiller. Arch. f. Anat., p. 13, Taf. iii, f. 9 — 13. 

„ Crithidia thalassina, G-osse. Ann. Nat. Hist., 2nd ser., xv, p. 308. 

1862. Polybostrichus Miiller i, Keferstein. Zeitsch. f. w. Zool., Bd. xii, p. 113, pi. xi, f. 1 — 6. 
1865. „ longisitis, De Quatrefages. Annel., II, p. 72, 

„ Crithidia thalassina, idem. Ibid., II, p. 81. 

„ Diplocersea comiculata, idem. Ibid., p. 76. 



AUTOLYTITS PROLIFER. 217 

1891. Autolytus Alexandri, Hornel. Trans. Liverp. Biol. Soc, vol. v, p. 245 (?). 
1893. „ prolifer, Levinsen. Yidensk. Ud./' Hauchs/' p. 330. 

$ Bud. 
1778. Scolopendrina marina, Slabber. Natuur. Yerlus., pp. 51 and 83, Taf. x, f. 3 — 5. 
1855. Sacconereis helgolandica $ , Max Miiller. Arch. f. Anat., p. 14, Taf. ii, f. 4. 
1865. „ „ De Quatrefages. Aimel., II, p. 74. 

1875. „ „ Mobius. Jabresb. Oomm. deutscli., p. 170. 

Habitat. — On shells and other debris procured by the deep-sea lines of the fishermen, 
St. Andrews ; on stones near low water-mark (whence they are best procured by 
immersion in sea-water, as they by-and-by crawl from the stones and growths thereon to 
the margin of the water) ; it occurs also in tough translucent tubes under stones (gneiss) 
between tide-marks in Bressay Sound, Shetland, where buds were found in July, 1871. 
It may be a question whether it has not tubes at St. Andrews likewise, for it was generally 
collected at the margin of a vessel into which stones from low water-mark had been placed. 
Nurse-stock and buds were likewise found amongst tangle-roots in 8 fathoms in the same 
region (Bressay Sound). 

Mediterranean. Atlantic. English Channel. North Sea. South Africa. Shores 
of France. 

Head smoothly rounded in front; eyes placed closely together on each side, but 
obliquely, so that the anterior are the wider apart, and they have a small lens. A long 
tapering median tentacle, and two lateral. Two tentacular cirri occur on each side. All 
these processes are more or less smooth. 

Body about half an inch in length, of a pale yellowish hue — somewhat darker in the 
middle from the intestine, and with numerous small whitish specks over that. It is 
slender, elongated, with numerous distinct segments, slightly and rather abruptly tapered 
anteriorly ; more distinctly so posteriorly, where it ends in two slender cirri. The cirri 
of the foot behind the tentacular cirri are considerably longer than the tentacular cirri. 

The proboscis in transverse section presents a radiate arrangement of fibres from 
the firm inner lining to the outer coat. 

The foot (Plate LXXI, fig. 4) forms a bluntly-conical or rounded process with a 
groove at the spine. The dorsal cirrus is comparatively short, the basal region or 
cirrophore being of some size, and distinctly differentiated from the distal region. The 
bristles (Plate LXXIX, figs. 23 and 23 a) are somewhat short, curved backward towards 
the abrupt dilatation at the tip, and have a short terminal piece with two hooks, the 
second being the larger. The dilated part of the shaft has minute spikes. A single 
simple bristle is generally present in each foot, the tip being bent and diminished to a fine 
point after the manner of Autolytus fallax, Malmgren. 1 

So far as present observations go, this form would seem to frequent the lower margin 
of the tidal rocks, and probably also the laminarian region beyond, and from this to deep 
water, so that the area for the production of buds is extensive. These buds ( J and ? ), 
from their pelagic habits, scatter the reproductive elements very widely, and, as they occur 

1 ' Annul. Polycheeta/ p. 153, pi. vii, fig. 41. 

85 



218 AUTOLYTITS PBOLIFER. 

throughout the year continuously, Nature has therefore provided for any losses the 
species may encounter in its varied life-history. 

The buds ( $ and ? ) are found in the pelagic condition all along the shores of 
Britain, from Shetland to the Channel Islands. Well-developed forms are met with 
every month, and the younger examples, as might be expected, are equally distributed 
over the year. 

Habits. — The female buds with the ventral sac (Plate XLIX, fig. 7) are truly pelagic, 
and are frequently caught in the large mid- water net. So far as the records and specimens 
of those captured by the tow-nets (and those chiefly the bottom tow-nets) go, the majority 
occur in February, though a considerable proportion are found in March and April. An 
occasional form appears every month from June till November. The greatest number, 
however, are procured in the months first mentioned. From the nature of their origin 
there is little difference in size between those obtained in February and those in October 
and November, and the continuance of schizogamy throughout this extended period is 
noteworthy. 1 

In glancing at a series captured in the tow-net near the bottom, for example, on 
April 20th, the diversity in size of the ovigerous females is interesting, for they range 
from 2*5 mm. to about 14 mm. The small forms carry a small mass of ova, the larger a 
great mass, and there is a marked difference in the size of the ova in these respectively. 
Thus, on the date mentioned, those of the small mass attached to the small example 
measured '07 mm., whilst those of the larger were *15 mm. The respective larvse 
probably show the same disparity, and those from the larger eggs thus might be supposed 
to enter life with a better chance of survival. That, however, does not necessarily follow, 
for the smaller may be more inconspicuous. 

After the escape of the larvas the body of the female is diaphanous, and the 
ovigerous membrane forms soft wrinkled folds at each side. It is probably a cuticular 
structure, augmented by a secretion of the dermal cells, as in the instance of the tube, 
and in communication with the body-cavity of the annelid. They probably break up 
and disappear soon afterwards. . The fine elastic ovigerous membrane keeps the eggs 
in situ during the active pelagic life of the animal. 

The head of Saeconereis (the female bud) is rounded in front, comparatively small, 
and with a pair of large connate eyes, situated at the outer and posterior border of the 
head. The upper part of the pigment looks dorsally and externally, the lower commands 
the ventral surface as a large dark brown eye on each side, and more conspicuous than 
the dorsal pair. Each is provided with a lens. A long median tentacle occurs behind 
the eyes, and two tentacular cirri. 

Eleven bristled segments follow (in one in March only three, and yet the form 
carried eggs), and they have a dorsal cirrus and a setigerous region ; then about thirty 
segments bearing the long dorsal swimming-bristles succeed. Each of these has a 
deeply-cut foot, which increases in size from the first for some distance backward, and 
has dorsally the cirrus (Plate LXXI, fig. 4), which is of moderate length, and smooth. 

1 It is interesting that Greef, also in December, found a translucent pelagic Saeconereis off the 
Canaries, and he followed the development of the larvae to the appearance of the first pair of bristled 
feet (' Zeitsch. f. w. Zool.,' Bd. xxxii, p. 251, Taf. xiv, f. 31—36). 



AUTOLYTUS PROLIFER. 219 

Beneath is a bluntly-conical region, which is stiffened by a group of spines, several 
of which are curved at the tip and spread out for supporting a web, it may be that 
enclosing the ova. From this region the dense and long tuft of swimming-bristles 
projects. These (Plate LXXIX, fig. 23 a) are translucent flattened bristles, which 
increase in diameter towards the broad terminal part, which then diminishes a little, 
and ends in a curved tip. 

These spines and bristles are moved by considerable strands of muscular fibres, a 
conspicuous series being dorsal and another ventral. The inferior division of the foot 
consists only of a setigerous region, which is bifid at the tip and bears a series of the 
characteristic jointed bristles, the shafts of which are slightly curved towards the end, 
which is bevelled to a point and serrated on each side. The terminal piece is short 
and bifid as in the nurse-stock. Posteriorly a considerable portion of the tapering 
caudal region is devoid of the long swimming-bristles. In addition to the long 
swimming-bristles is the group of ordinary bristles with abruptly-dilated ends to the 
shaft and a short bifid terminal piece. In many is also a single modified bristle in each 
tuft, having a somewhat dilated end to the shaft and a lateral spike or process. 

The region bearing the ova is reddish and green, the former colour being chiefly 
at the segment-junctions. The region behind the sac is greenish, then pale posteriorly. 
The colours are similar but paler ventrally. The egg-sac, from a central constriction, 
has the shape of an hour-glass. The region behind the long bristles varies in the 
number of its segments — from ten to twelve or upwards (in the ovigerous examples), 
the tip being furnished with a pair of short symmetrical styles or cirri. Other 
specimens in the same month (March) show a larger number of segments anteriorly 
and posteriorly to the ovigerous region and the long bristles. 

The stages reached by the embryos in the sac of the pelagic Sacconereis are 
indicated in the following notes at different dates : 

April 19th, 1887. — Embryos in sac developed, and they swim in the water ; some have 
three segments, other examples have four, besides head and tail (Plate LX, fig. 4), with 
palpocils. Eyes four. Head of adult pinkish, and the eyes dark brown. Two appear 
to be dorsal, but the pigment is closely connected with the ventral. The ventral 
position of these eyes would certainly enable the animal to look downward. The eyes, 
indeed, form a large double mass, the anterior looking upward and outward, the other pair 
commanding the ventral surface. The tentacles are long and smooth. 

April 26th, 1887. — The young Autolyti have only two pairs of bristle-bundles, but the 
feet extend further outwards and are more distinct. They have a more elongated and 
pointed dorsal region, and a shorter and thicker ventral. The number of segments is the 
same. A process appears on the head — probably a tentacle. 

April 27th, 1887. — Numerous females in the bottom trawl-like tow-net. Some are 
dull yellowish or paler, others greenish. The ova thus differ in tint. 

May 9th, 1887. — The young have increased in size. The head has lost its palpocils. 
The four eyes are still nearly in a transverse line, the two median being dorsal; the two 
lateral are at the sides. The buccal segment has a large pharyngeal cavity. The next 
two segments have rudimentary feet with bristle-bundles, the tips being bifid as in 
Autolytus. The penultimate and the terminal segments have no bristles. The animal is 



220 ATTTOLYTUS PROLIFER. 

evidently feeding, and the alimentary canal is distended. May 11th, 1887. — The yonng 
are now pale greenish, with eyes of a lnstrons silvery bine. 

The male buds (Polybostrichus, Plate L, fig. 11) occur from February onward to 
November, and probably throughout the entire year, and are discriminated by their 
shorter bodies and structural differences. 

Head comparatively small, with two circular dark brown eyes (in spirit), reddish 
or blackish in life, which have a lens and which look laterally as well as dorsally, a 
long median tentacle arising behind them, and a large and long pair of cirri laterally (post 
cephalic). From the front of the head project the bifid palpi, which are undivided and 
massive at the base. The outer division is short, the inner long, tapering, and often 
thrown into graceful coils. These processes appear to develop considerably after the 
separation of the bud from the nurse- stock — just as the body increases in length and bulk. 
A smaller and more slender cirrus is ventral to the great tentacular cirrus, and a still 
smaller to the outer side of the eye. 

On the ventral aspect of the head a large pair of eyes occupies each lateral region, 
and each also shows a cuticular thickening or lens. These eyes chiefly look downward, 
though in a lateral view of the head a considerable portion of the pigment. of both dorsal 
and ventral pairs is visible. 

Body increasing in length, and probably in the number of its segments, with age; 
slightly narrowed in front and tapering to a slender tail. A median furrow runs along 
the dorsal surface, while ventrally the nerve-cord and the adjoining ventral longitudinal 
muscles form three central stripes. Behind the head are fourteen narrow segments 
provided with slender dorsal cirri, which are largest in front, and the ordinary bristles 
with bifid terminal pieces. 

In the next region the feet project more, so that the body is broader, and the 
dorsum of each has a tuft of the long swimming-bristles. These are longest in the 
anterior twelve or fourteen segments and diminish gradually in the succeeding. The 
feet so provided were twenty-eight in number in a fine example fully half an inch in 
length. In these the dorsal cirrus is shorter than in the anterior region. Each foot 
has also beneath the former a group of the ordinary bristles with the bifid terminal 
pieces. About thirty diminishing segments with short dorsal cirri, and the ordinary 
bristles with the bifid terminal piece, occur posteriorly, the last having two larger cirri, 
though they are not conspicuous. The peculiarly broad and flattened body of this form, 
and its long bristles, fit it, like the young Nerine, for example, for its pelagic life. The 
specimen from which the foregoing description was drawn up was captured on February 
1st. Most of those procured in the subsequent months were smaller. 

The stolons have neither proboscis nor proventriculus. They swim gracefully and 
vigorously, the sheen of the long lustrous bristles with their ever-changing hues being 
very beautiful. 

De St. Joseph gives an account of the changes in the muscular system in the second 
region of this form (male). 

Martin Slabber (1778) describes and figures under the name of Scolopendrina marina 
(Zeedurzendbeen) a female bud of this form, with swimming-bristles, in which the larvaa 



AUTOLYTITS PKOLIFEE. 221 

in the ventral sac are advanced and some have escaped into the surrounding water. They 
appear to have four or five post-cephalic segments. On the whole the figures are good. 

0. F. Miiller mentioned the male bud and the nurse-stock as separate species, but 
he correctly figures them, and assigns them different habitats, the former being pelagic 
with long bristles, the latter as only rarely occurring in the gulfs of Norway. He 
describes the nurse-stock and its female buds as mother and foetuses, and he recognized 
the ova in the buds. 

From this period onward the gradual unfolding of the life-history of Autolytus is one 
of the most interesting chapters in zoology — associated as it is with one of the very 
prominent examples of the so-called " alternation of generations." The nurse-stock and 
its divergent sexual bucls were each regarded as separate species by the earlier observers, 
such as 0. F. Miiller, (Ersted, and Max Miiller. It was not till Leuckart and Frey, and, 
more definitely, Krohn, had shown the relationships of Sacconereis and Autolytus, and 
De Quatref ages and he had studied the conditions in Syllis prolifera, that the true nature 
of the processes were elucidated. Further advances were made by Alexander Agassiz, 
Keferstein, Greef, Ehlers, Malaquin, and others. Thus the male and female buds, 
formerly known as Polybostrichus and Sacconereis, were linked to the nurse-stock 
Autolytus, and the structure and physiology of the parts elucidated. 

The genus Sacconereis was established by Johannes Miiller, 1 and his son Max Miiller 
(1855) gave good descriptions and figures of both male and female of Sacconereis 
helgolandica, the female carrying the ova in the sac and with swimming-bristles from the 
fourth foot backward. He shows that the sperms are found in the first three bristled 
segments of the male, and gives figures of four stages in the development of the young. 
The forms were at that time considered as independent. 

Alex. Agassiz 3 gives a careful account of the life-history of a species he terms 
Autolytus cornutus, which does not seem to differ in any essential respect from Autolytus 
prolifer (and in this Greef and others coincide) except in the structure of the compound 
bristles, which he figures as having a simple " sickle-shaped " tip, a feature at variance 
with the usual structure of Autolytus, and which may have arisen from misinterpretation. 
He furnishes an excellent description, with figures of the nurse-stock, male and female 
bucls, and for the first time the development of the eggs (borne on the ventral surface in 
a sac), and young issuing from these up to the stage of the nurse-stock, thus illustrating 
the alternation of generations in the species. 

The Polybostrichus Mulleri of Keferstein 3 (1863) seems to be a male form with much 
longer tentacles than usual, but it may be only a variety. The body is in two parts, the 
anterior having 19 — 22 segments, the posterior with long swimming-bristles. 

Claparede 4 describes and figures a new species under the name of Autolytus roseus, 
which bears a very close resemblance to a female Autolytus prolifer bearing the sac with 

1 "Ueber den allgemeinen Plan in der Entwickelung der Echinodermen/' 'Phys. Abh. Akad. 
Wiss. Berlin/ 1852, p. 52 (1853). 

2 ' Journ. Bost. Soc. Nat. Hist./ vol. vii, p. 392. 

3 ( Zeitsch. f. w. Zool./ xii, p. 113, pi. xi, f. 1—6. 

4 < G-lanures/ p. 106, pi. vii, fig. 4 (1864). 



222 ATTTOLYTUS PROLIFER. 

eggs. As he figures the bristle with a simple hook to the terminal piece it is possible 
that a misinterpretation has occurred. 

Malaquin (1893) states that the formation of the sac is always subsequent to the 
separation of the stolon, but apparently he has not seen it. The first and second setigerous 
segments carry the eggs, and they have no swimming-bristles. The rest of the segments, 
with the exception of a few of the last, are provided with these bristles. 

He gives a careful account 1 of the development of Autolytus from -^ mm. 
onward. The youngest stage is devoid of bristles, shows a head with four eyes, the 
anterior with lenses, and palpocils, a tail, and two constrictions, with cilia on the elevated 
regions between. The pharynx is indicated. In the next stage, also achsetous, the body 
is longer, shows constrictions, the cephalic and post-cephalic segments have cilia, the 
pharynx abuts on an intestine, but the latter is devoid of an anus. Rudiments of the 
lateral tentacles appear, and the proctodeum is formed. 

In the next stage the larval Autolytus is larger, the head is separated from the 
peristomial segment, and bristles appear in the first segment, the lateral tentacles are 
longer, and the median appears as a bud. Ciliated dorsal bands occur on the head and body. 

The pharyngeal apparatus is only completed when the young form has from twenty- 
two to twenty-five segments. It feeds by thrusting out its proboscis and engulfing prey. 
At the sides of the larval pharynx are glands. Moreover, Malaquin points out that at the 
stage of eight or ten segments the larva presents two pharyngeal bulbs, the posterior 
becoming the proventriculus. By-and-by the transformation, by disappearance of the 
anterior enlargement, takes place, and the pharynx acquires the sinuous adult characters, 
when there are about twenty-two segments with bristles. 

The Autolytus fallax of Malmgren (1867) does not seem to differ in any important 
point from this species. 

In connection with this group is a bud which may be termed Autolytus A. 

Habitat. — Clinging to an osculum of Halwhondria panicea under a stone in a pool at 
the East Rocks, St. Andrews, November 5th, 1863. 

Head rounded, small, with two large round black eyes. A comparatively short 
median and two lateral tentacles. 

Body about half an inch long, not much diminished anteriorly, but tapered 
posteriorly, and the segments are distinctly marked. 

The foot (Plate LXX, fig. 10) has dorsally a comparatively short and unjointed cirrus, 
the basal region (ceratophore) apparently not being differentiated. It is tapered to a 
blunt tip. The space between the cirrus and the setigerous region of the foot is short, 
and is supported by some slender spines which separate from those passing to the seti- 
gerous lobe. A tuft of very slender translucent bristles projects from this region about 
as far as the tip of the setigerous lobe. These may be developing swimming-bristles. 
The setigerous lobe is ovate, and has a series of slender spines in its centre. A careful 
search furnished one or two bristles (Plate LXXIX, fig. 3) of the form shown. 

This differs from Johnston's Ioida macrophthahna in the structure of the foot, since 
the cirrus is obscurely moniliform and twice the length of the foot, whilst there are two 

i Op. cit., pp. 392—404. 



AUTOLYTUS PPOLIFER. 



223 



bundles of bristles— the superior capillary, the inferior articulated. Unfortunately the 
bristles are imperfect, only a shaft (Plate LXXTX, fig. 3) having been obtained. The 
peculiar nature of the dilatation, together with the shape of the foot, would seem to point 
to its connection with Autolytus rather than either jEusyllis or other form. 

Autolytus ? Female Bud. 

A form captured by the tow-net in the Firth of Clyde differs from the ordinary 
female Autolytus carrying ova in several particulars, and I am indebted to Professor Graham 
Kerr, of Glasgow, for bringing it under my notice just as this part is passing through 
the press. The sole example (Fig. 57) is stained and mounted on a slide, so that only an 
imperfect description can at present be given. It appears to be fully a quarter of an inch 
in length, and is readily discriminated by the absence of the single ventral egg-capsule, 
the ova being borne in eight conspicuous globular or pear-shaped sacs. 



fi 



n ii! 




*L ' / 








ps\) 



1 !m\Y \H v \ 



I 



Fig. 57. — Autolytus (female bud) with, pyriform lateral sacs containing ova. 



The head resembles that of an Autoly ius -bud, with a long median and two somewhat- 
shorter lateral tentacles, a large rounded dorsal and a larger ovoid ventral eye occur 
on each side. Each appears to be furnished with a lens. Then follows the buccal 
segment with two tentacular cirri on each side, the base of the ceratophore being 
opaque, but apparently without differentiation into a rudimentary ovigerous sac, yet 
the preparation left uncertainty on this point. The second segment bears a group of 
typical compound bristles, though the minute bifid terminal pieces are absent, and 
behind the foot a rounded swelling resembling an egg. A similar condition exists in 
the third, fourth, and fifth feet, only these have longer dorsal cirri in the preparation. 
The sixth has the same arrangement on one side, but on the other an egg lies in front 
of the foot in the ccelom. The seventh has similar bristles, but on one side a large 
ovisac. The eighth foot has a lanceolate flap and a tuft of long tapering swimming- 
bristles, and the setigerous region with the compound bristles is larger and truncated 
distally. Altogether about fourteen segments (i.e., to the twenty-first) bear the long 
swimming-bristles, whilst the caudal region has about twenty-six segments, the body 



224 AMBLYOSYLLIS. 

gradually tapering and terminating in two short and slightly clavate caudal cirri. The 
great pear-shaped egg-sacs are arranged alternately, so as to give room for the distal 
globular region of each, and appear to occupy about ten segments. The dorsal cirri 
are longest in the ovigerous region, and diminish to short processes posteriorly. The 
alimentary canal is rudimentary, but is open in front. 

The relationships of this form have not yet been ascertained, though it is clearly a 
female bud of an Autolytus having this characteristic method of carrying the com- 
paratively large circular granular ova. Whether it has any connection with such forms 
as Procer&a, Bhlers, which is here included under Autolytus, remains to be seen. 

Professor Graham Kerr 1 noticed that this form quite differed from the ordinary 
female bud of Autolytus prolifer. 

The structure of the head, the arrangement and nature of the ovisacs, and other 
features, distinguish it at once from Kolliker's three forms, viz., Exogone (Erstedii, 
Exogone cirrata, and Gystonereis Edwardsii, 2 all of which bear ova in sacs. 



Genus LVI. — Amblyosyllis, Grube, 1857. 

Head short. Eyes, four pairs on each side, confluent. Tentacles and cirri long and 
slender. Palpi small, appearing as two bosses on the ventral surface. Two nuchal flaps. 

Pharynx long, sinuous, armed with a point (trepan, Malaquin) and a circle of 
papillge. Pharyngeal glands posteriorly, ventricule small, and with two minute cseca. 

Body of few segments; cirri thread-like, not articulated (Langerhans). Ventral 
cirrus lanceolate. Falcate bristles bidentate. Penultimate segment provided with two 
pairs of cirri (Malaquin). 

Grube's genus Amblyosyllis (18*57) has priority. 

Claparede 3 again established the genus Pterosyllis for this species, but the characters 
had probably been known previously in this country. He describes the proboscis as 
furnished with bifid cuticular papillse in P. formosa, which seems to be very closely allied 
to P. spectabilis. His Pterosyllis dorslgera 4 may have some connection with a female of 
the same species, though the moniliform condition of the cirri is noteworthy. Subse- 
quently Ach. Costa, 5 without having seen Claparede's accounts, described probably the 
same form under the name of Nicotia lineolata. The publication of the ' Catalogue of the 
British Museum' 6 added another generic name for the same type, viz. Gattiola. 

Malaquin notes that the nuchal flaps (ailerons occipitaux) receive two large nerves 
from the occipital lobes of the cephalic ganglia. Lang considers them organs of smell, 
but Malaquin doubts this interpretation. 

i 'Trans. Nat. Hist. Soc. Glasgow/ N.S., vol. viii, p. 4 (1905-6). 
3 ' Nouv. Mem. de la Soc. Helvetiqne/ Bd. vii, p. 14, Taf. iii. 

3 'Beobach u. Anat./ etc., p. 46 (1863), and ' Gdannres/ p. 100 (1864). 

4 ' Gdannres/ p. 100, pi. vi, fig. 1. 

5 < Ann. del Mns./ p. 160, 1864. 

6 'Cat. Brit. Mns./ p. 195. 



AMBLYOSYLLIS LINEATA. 225 



1. Amblyosyllis lineata, Grube, 1864. Plate XL VIII, fig. 4; Plate LXXI, fig. 1 — foot; 

Plate LXXIX, fig. 20— bristle. 

Specific Characters. — Head small, transversely ovoid, with a median and two lateral 
tentacles of considerable length (indistinctly articulated, De St. Joseph). Palpi small, 
appearing as two bosses on the ventral surface. Eyes four, red, the anterior slightly 
the wider, and the pairs on each side almost connate. Behind the eyes are the ciliated 
nuchal wings, which have a sepia-brown tint with a pale border. Body about half an 
inch in length ; segments sixteen, tapered in front and again posteriorly to a less degree, 
and ending in a bluntly-conical segment bearing two long cirri. General colour purplish 
brown, with definite yellowish patches. Feet thirteen pairs. Dorsally is the long 
slightly crenate cirrus. Near the base is a tuft of slender hair-like (swimming) bristles 
in certain females. Setigerous lobe stout with a conical papilla above the spine. Bristles 
a dense group, each bristle dilated and bevelled at the tip. Terminal piece longest in 
the superior bristles, tapering to a hook distally, and on the edge beneath is- a secondary 
process, so that it is bifid. Ventral cirrus of a curved lanceolate form, and it scarcely 
reaches the tip of the setigerous region. 

Synonyms. 

1863. Pterosyllis formosa, Claparede. Beobacht., p. 46, pi. xiii, f. 31 — 34. 

1864. „ „ Ehlers. Borstenw., p. 222. 

„ Nicotia lineolata, A. Costa. Ann. del Mus. Zoologico, p. 160, Taf. hi. 

,, Amblyosyllis lineata, Grube (?). Insel Lussin, p. 84. 

1865. Gattiola spectabilis, Johnston. Oat. Brit. Mus., p. 195, pi. xviA, f. 1—7, and p. 342. 
„ Pterosyllis formosa, De Quatrefages. Annel., II, p. 17. 

1867. Gattiola spectabilis, Parfitt. Trans. Devon. Assoc, ii, pt. 1, p. 236. 

1874. ,, plectoryncha, Marenzeller. Op. cit., p. 47, pi. v, f. 3. 

„ Stephanosyllis omata, Verrill. Proc. Americ. Assoc. So., 1873, p. 378, pi. iv, f. 1. 

1875. Pterosyllis lineolata, Marion and Bobretzky. Ann. Sc. Nat., 6 e ser, t. ii, p. 43, pi. iv, f. 13, 

and pi. v, f. 13. 
1879. Amblyosyllis madeirensis, Langerhans. Zeitschr. f. w. Zool., t. xxxii, p. 561, pi. xxxii, f. 19. 
1885. „ lineolata, Cams. Fauna Medit., p. 231. 

1887. Pterosyllis (Gattiola) spectabilis, De St. Joseph. Ann. Sc. Nat., 7 e ser, i, p. 187, pi. ix, 

f. 64—67. 

1904. Amblosyllis spectabilis, Allen. Journ. M. B. A., N.s., vol. vii, p. 220. 

1905. „ lineata, Graeffe. Arbeit. Zool. Stat. Triest, xv, p. 324. 

Habitat. — Under stones near low water-mark, in the creek to the south of the 
harbour (behind the first small promontory) at Lochmaddy, North Uist, and again near 
the foot of the Lee hills, under a stone, near low water-mark. Off Fermain Bay, Guernsey, 
in 10 fathoms ; and under stones in tidal pools, Herm. Off Plymouth, where it occurs in 
considerable numbers in sponges (Miss Florence Buchanan, Garstang, and Allen). 

Naples; Marseilles (Marion and Bobretzky). Eastern shores of America (Verrill) ?. 

Head small, transversely ovoid, with three long tentacles — a median and two lateral 

.all of considerable length (forty to fifty articulations, De St. Joseph). The palpi are 

86 



226 AMBLYOSYLLIS LINEATA. 

small and form two bosses on the ventral aspect of the snout. Eyes four, red, situated 
obliquely on each side, the anterior pair slightly wider than the posterior, and the pairs 
on each side connate, or almost so. Some specimens have a black point in the centre of 
each. Behind the eyes are the cephalic wings, which have a sepia-brown colour with a 
pale border. Tentacular cirri two pairs (one long and the other short). 

The cirri on the head and first segment are sometimes pinkish from the presence of 
transverse reddish bars, and De St. Joseph found palpocils on all. 

Body about half an inch or more in length (13 mm., De St. Joseph), segments 
sixteen to thirty (Marion and Bobretzky), deeply indented at the sutures, but not 
narrowed in front. It is tapered anteriorly, and again posteriorly, but to a less degree, 
and ends in a somewhat shield-shaped or bluntly-conical segment bearing two long cirri. 
The body is slightly rounded dorsally, flattened ventrally, and witli a median ventral 
groove in the preparations. The general colour is purplish brown, with peculiar and 
definite yellowish patches. A dark brown bar, with an angle directed backward and 
thinned off at either end, crosses the segment in a line with the origin of the lateral cirri. 
Behind is a narrow, pale band with a somewhat cordate mark in the centre, succeeded by 
a broad brown band which runs into the dark belt first mentioned, and has on each 
side a pale and somewhat triangular patch which diminishes anteriorly as the brown of 
the general surface slopes to meet the first dark belt. Considerable variation in the 
arrangement of the pigment-bands occurs — De St. Joseph, for instance, describing only 
narrow, straight transverse bars. The last segment is dark brown. The penultimate 
segment is compressed and has no setigerous process, only two elongated cirri on each 
side, the longer inferior. 

When the male is fully distended with ripe sperm the body recalls the condition in 
the Queen termite. 

De St. Joseph states that the proboscis has six large teeth, each subdivided into 
three divisions (tricuspid). The proventriculus has eighteen rows of points, and instead 
of the two horny pieces of other Syllids has two horny valves. 

Feet (Plate LXXI, fig. 1) thirteen or more pairs, increasing in size from the front 
nearly to the posterior end. They vary much in appearance, according to the condition 
of the reproductive elements, as likewise does the body. Dorsally is the long cirrus 
which arises from the posterior part of the lobe. It is slightly crenate when living, and 
has minute grains or specks of white pigment. Near the base of the dorsal cirrus in 
certain female examples is a small tuft of slender hair-like simple bristles, which readily 
become curved (from lack of stiffness). These probably represent the pelagic bristles of 
other forms. They were not present in a male distended with nearly ripe sperms. 

The setigerous lobe is stout, with a bevelled tip, and superiorly is a conical papilla 
above the exit of the spine, which is supported by four or five others. The bristles 
(Plate LXXIX, fig. 20) form a dense group, the longer occurring superiorly, the shorter 
inferiorly, so that the outline of the brush slopes from above downward and inward, as 
indicated in Plate LXXI, fig. 1, where the foot is observed from above; the translucent 
and slightly curved shaft of the bristle is dilated at the tip which is bevelled to a point. 
The terminal blade is longest superiorly, shortest inferiorly, and tapers to a hook at the 
tip, while on the edge beneath is another process, less distinct, for instance, than in 



AMBLYOSYLLIS LINEATA. 227 

Pionosyllis, so that the blade is bifid. The edge is probably minutely serrated, but 
this could not be satisfactorily determined. 

The ventral cirrus arises from the posterior edge of the foot, is of a curved, 
lanceolate form, and scarcely reaches the tip of the fleshy part of the setigerous region. 
The occurrence of the ventral cirrus distinguishes this genus from both Autolysis and 
Myrianida, and is another instance of the plastic nature of the appendages in the 
Annelids. 

Reproduction. — The period of reproductive activity is, in all probability, in July. 
A ripe male was found at Herm in August, and an advanced female, in the same month, 
at Lochmaddy, North Uist. The changes in the form of the body and bases of the feet 
at maturity are marked. In one the seventh segment was more dilated than the rest, but 
this may have been accidental. The ova are of an orange or brownish hue. 

. De St. Joseph l describes the eggs as green or brown, while the sperms are white. 
The natatory bristles commenced on the seventh segment. The eight segments filled with 
the sexual elements are largely dilated, and their colouration is reduced to a simple brown 
band. The eyes are largely increased in size, and cover almost the entire head. After 
the development of the natatory bristles the animal swims with great activity. The 
reproduction is direct, without alternation of generations. 

Malaquin 2 observed a young Amblyosyllis of eight bristled segments. Its proboscis 
is long and coiled, but the larval pharynx is still visible anteriorly. The proventriculus 
is present though small. The occipital wings and minute palpi are present. The stage 
appears to agree with that of an Autolytus of twenty-two segments. 

Habits. — It is a most interesting and beautiful annelid in confinement. In repose the 
cirri are gracefully coiled on the dorsum, but on touching the animal with a needle they 
are stretched out, the opposite of what occurred in Mr. Hancock's specimen, though it 
agrees in part with Mr. Dyster's experience as mentioned by Dr. Johnston. When moving 
about a large trough (of a dissecting-lens) it occasionally coiled up the long cirri, and 
swam through the water with a rapid wriggling motion like JEvame or Harmothoe. In 
its native sites it cannot have much scope for locomotion, and this may explain its habit of 
remaining quiescent for a long time in a glass vessel, its cirri being closely coiled. 

Specimens very readily fracture themselves in confinement, as, for instance, in a 
bottle amongst other marine animals, or when put in spirit. Yet they may be kept alive 
for a considerable time. Thus one obtained near the foot of the Lee hills at Lochmaddy 
was carried in a small bottle (sij) therefrom to the summit of the mountain and across to 
the shore on the southern side, where the sea-water was renewed. Moreover, ten or 
twelve days thereafter it was transported to Murthly in Perthshire, where the coloured 
figure was made and where it lived for six weeks. 

In a specimen from Herm the bases of the cirri were covered with minute whitish 
parasites, probably the forms alluded to by De St. Joseph, 3 viz. Infusorians. He mentions 
Trichodina Auerbachii, Cohn, Licrophora Auerbachii, Claparede, and an Acineta (Ophryo- 

1 Op. cit., p. 189 (1886). 

2 Op. cit., p. 423, pi. xiv, f. 30. 

3 < Ann. Sc. Nat./ 7 e ser., I, p. 190 (1886). 



228 MYEIANIDA. 

dendron annulatorum) on the tentacles and cirri. He also found a number of Cercarige 
and Distomes in the intestine. 

"What the Amblyosyllis rhombeata (1857) of Grube may be is doubtful ('Annul./ (Erst., 
1857, p. 29). It has fourteen sub-rhomboid segments, and came from St. Croix. 
De Quatrefages thinks it is near Ioida, but this can hardly be. Grube's A. lineata, 1 which 
is roseate and has sixteen segments, from Lussin Picolo, is nearer the British species, 
and may, indeed, be that form. 

Malmgren 3 describes the falcate bristles of his Gattiola finmarchica as simple, yet in 
other respects the form does not appear to differ from the British. It is possible that the 
minute secondary process may have been overlooked. 

De St. Joseph in 1886 points out that without any anatomical differences four species 
of this form have been made. 

It is doubtful if the species described by Marenzeller (1874), viz., P. plectorhyncha, 
differs in any essential degree from the present form. A further examination of P. lineata 
is also necessary. 

A similar species (I. cincinnata, Verrill, 1879) occurs on the American shores; indeed, 
it is probable that the same form is found there. The Stephanosyllis picta 3 and ornata 4 " of 
Verrill do not appear to differ materially from the British annelid. Two names are thus 
given to the same species, and two figures (PI. II, fig. 1, and PL IV, fig. 1) seem to refer 
to the same form. 

Malaquin 5 points out the structure of the proventriculus of this annelid, the muscular 
columns having the form of a pyramid, which in transverse section presents two fissures. 
They also show from fifteen to twenty striations. 



Genus LVII. — Mteianida, Audouin and Milne Edwards (1845). 

Head devoid of palpi, three tentacles — a median and two lateral. A pair of tentacular 
cirri. The foregoing and the cirri thick and clavate, or almost foliaceous. No ventral 
cirrus. Reproduction by schizogamy. 

The genus was established by Milne Edwards, who termed the phenomena of its 
reproduction, multiplication by buds. Ehlers placed it near Autolytus and Procerxa. 
The second example was procured by Claparede at Naples, and the author considered that 
it was not mature, being only 3'5 mm. 6 in length, whereas the former specimen (Milne 
Edwards') was 64 mm. He termed it M. maculata. Viguier found the female form 
(Sacconereis) of M. fasciata. De St. Joseph observed two examples of M. maculata. 
Giard showed that M. fasciata, M. Edwards, agreed with M. jpinnigera, Montagu. 
Malaquin again seems inclined to ignore Giard's diagnosis. 

1 ' Archiv f. Naturges./ 1863, p. 48, Taf. v, fig. 1. 

2 < Annul. Polych./ p. 38, Tab. vi, fig. 36 (1867). 

3 ' Americ. Journ. Sc./ iii, vol. vii, p. 132 (1874), pi. v. 

4 'Proc. Americ. Assoc. Sc./ 1873, p. 378. 

5 Op. cit., p. 230. 

6 This is probably a misprint for 35 mm. 



MYRIANIDA PINNIGBRA. 229 



1. Myeianida pinnigera, Montagu, 1808. Plate LXXI, figs. 5 and 6— feet; Plate LXXIX, 

figs. 24 and 24 a — bristles. 

Specific Characters. — Head comparatively small, bluntly conical or irregularly 
rounded. Eyes large, the two anterior being wide apart, while the pairs on each side are 
close together. Palpi apparently fused, forming two rounded bosses with a median 
fissure in front. Median tentacle long and somewhat clavate, lateral shorter and dilated 
at the tip. First pair of tentacular cirri curved and short; second clavate and rather 
longer. Body (of nurse-stock and buds) about 1^ in. long, the two sections being nearly 
equal. Nurse-stock dull whitish and brilliantly marked by thirteen rich orange spots, 
two irregular longitudinal patches in front, followed by a pair, and then nine well- 
defined spots, having a deep orange marginal belt and a paler orange centre. Two 
orange dots occur on the last segment of the nurse-stock. Behind the tentacular cirri 
are two or three cirri of similar form, though with a tendency to flattening, but becoming 
ringed in spirit, and thus differentiated from the succeeding. Foot with the dorsal 
cirrus in the form of a flattened oar, which is translucent, with a faint milky opacity, and 
of a tissue more lax than the ceratophore. Setigerous lobe has superiorly a prominent 
papilla above the spines, whilst the outline of the ventral border is nearly semi- circular. 
Bristles rather feeble, elongated, the distal end of the shaft dilated, and the bevelled 
region spinous distally. Terminal piece short and diminished towards the bifid tip. 

Synonyms. 

1808. Nereis pinnigera, Montagu. Trans. Linn. Soc, ix, p. Ill, Tab. 6, f. 3 (nurse-stock and bud). 

1812. „ „ Pennant. Brit. Zool., iv, p. 95. 

1815. „ maculosa, Montagu. Trans. Linn. Soc, xi, p. 21, pi. iii, f . 4 ( $> bud). 

1845. Myrianida fasciata, M. Edwards. Ann. Sc. Nat., 3 e ser., t. hi, p. 170, pi. xi, f. 65 — 68. 

1851. Nereis pinnigera, Grrube. Fam. Annel., p. 63. 

1864. „ „ Kolliker. Kurz. Ber. Scliotl., p. 9. 

1865. Nereis pinnig era, Johnston. Cat. Buit. Mus., p. 281. 
„ „ maculosa, idem. Ibid, pp. 198, 280 (bud). 

„ Myrianida fasciata, De Quatrefages. Annel., II, p. 63. 

„ „ pinnigera, Johnston. Cat. Brit. Mus., p. 196. 

1867. „ „ Parfitt. Trans. Devon. Assoc., ii, pt. i, p. 236. 

„ Amytidea maculosa, idem. Ibid., p. 29. 

1869. Myrianida fasciata, Langerhans. Zeitsch. f. w. Zool., xxxii, p. 581. 

1885. „ „ et maculata, Carus. Fauna Medit., p. 237. 

1886. „ ,. Viguier. Arch. Zool. Exper., 2 e ser., iv, p. 432, pi. xxvii, f. 21 and 22. 
„ „ „ Harvey Gibson. Proc. Lit. Phil. Soc. Liverp., vol. xi, p. 154. 

1890. „ pinnigera, Griard. Bull. Sc. Fr. Belg\, t. xxii, p. 79. 

1893. ,, fasciata, Malaquin. Recherch. Syllidiens, ubique in text and figs. 

1895. „ maculata, De St. Joseph. Ann. Sc. Nat., 7 e ser., torn, xx, p. 195. 

1904. „ pinnigera, Allen. Journ. M. B. A., n.s., vol. vii, p. 220. 

Habitat. — Dredged in 10 fathoms at the northern end of Bressay Sound, amongst 
horse-mussels, July, 1871 ; and, again, in the Sound of Harris, in 3 to 4 fathoms, in 



230 MYRIANIDA PINNIGERA. 

August, 1872, amongst the roots of tangles (nurse-stock with female buds). Amongst 
ascidians and sponges, Plymouth (Allen). 

Shores of France (Audouin and Edwards). Madeira (Langerhans) . 

Head comparatively small, bluntly conical or irregularly rounded, the transverse 
exceeding the antero-posterior diameter in spirit-preparations, whilst the posterior border 
is rounded and the anterior truncate. The eyes are large, the two anterior being the 
wider apart and with lenses, though the pairs on each side are close together. The palpi 
appear to be fused, forming two rounded bosses with a median furrow in front. The 
median tentacle is long and somewhat clavate, the lateral on each side shorter, more 
cylindrical, and somewhat dilated at the tip. A short, curved pair of tentacular cirri 
follow, then another clavate pair, rather longer than the first. All have vibratile cilia. 
De St. Joseph describes the median and lateral tentacles in his M. maculata as foliaceous 
like the cirri in the nurse-stock of sixty segments. There would thus seem to be variation 
in this respect. 

Body (in nurse-stock and buds from Bressay Sound) about 1^- in. long, 1 the two 
sections being about equal in length. The dull whitish nurse-stock in front is brilliantly 
marked by thirteen rich orange spots, two irregular patches in front, the first commencing 
behind the eyes, and median in position, with the posterior end somewhat bifid, and the 
second occasionally is also chiefly longitudinal, followed by a pair (the third), and then 
by nine well-defined spots having a deep orange marginal belt and a paler orange centre. 
Two orange dots occur on the last segment of the nurse-stock. The posterior region or 
section is formed of buds, the largest (and oldest) being posterior. Eight or nine had 
distinct eyes, in the rest these were obscure. 

Behind the long clavate cirrus mentioned in connection with the head are two or 
three shorter cirri of similar form, the last, however, having a tendency to flattening. 
All the foregoing cirri and tentacles are more or less ringed in spirit, thus being at once 
differentiated from the succeeding cirri, which assume the form of flattened oars, and 
continue of this shape to the posterior end. In those in process of regeneration the 
organs are narrower and more pointed. The cirri throughout are translucent, with a 
faint, milky opacity. The buds have minute cirri of the same form as the nurse-stock. 
The terminal (and oldest) bud had two longer caudal cirri. On the dorsum of the buds 
the orange specks are chiefly arranged in pairs. 

Claparede (1868) describes the proboscis as similar to that in Procerdea aurantiaca, 
the anterior edge without papillae or denticles, and the proventriculus with about thirty- 
four rows of points. 

De St. Joseph describes the nurse-stock as having in his example sixty-six segments. 
Then followed a bud of ten segments, and behind it fifteen others. The first had a 
rudiment of a head without eyes, with two rudimentary palps. It had three segments. 
The second and third were similar, but with four segments. The fourth agreed with the 
foregoing, but had eight segments. The fifth had four minute eyes and twelve segments, 
and a simple pigment-touch. The sixth and seventh were similar, and the palps were 

1 Malaquin gives 32 mm. for nurse-stock and 38 mm. for stolons in one, sixty-six segments 
(twenty-nine stolons) . 



MYRIAMDA PINNIGEKA. 231 

differentiated ; they had twenty segments. The eighth and ninth corresponded, and the 
bifurcation of the palps had commenced. In the eleventh and twelfth the bifurcation of 
the palps was marked, and the tentacles were present. They had twenty-three segments. 
From the sixth to the twelfth bud there were four reddish-orange touches. In the 
thirteenth and fourteenth the palps were completely bifurcate, the tentacles well 
developed, and the eyes large. The fifteenth and last ripe stolon became detached under 
observation. 

Foot. — In the nurse-stock (Plate LXXI, fig. 5) dorsally is a long ceratophore 
rounded at the end. It bears the large flattened cirrus, the tissue of which is much 
more lax than that of the basal process, and has pigment and glands. The cirrus forms 
a long flattened blade, somewhat pointed at the tip, and having a central nerve of con- 
siderable size, which gives off branches on its way to the tip. The inferior lobe has 
superiorly a prominent papilla above the tips of the spines, which form a group. The 
outline of the ventral border is nearly a semicircle, the swollen process inferiorly having 
pigment and glandular tissue. The ventral bristles (Plate LXXIX, fig. 24) are rather 
feeble, with elongated, slightly curved shafts dilated at the tip, the edges of the bevelled 
region distally being minutely spinous, whilst the terminal piece is short and diminished 
towards the bifid tip. As usual the points of the hooks are directed dorsally. 

In the bud a distinct papilla occurs between the ceratophore of the dorsal cirrus and 
the base of the ventral lobe (Plate LXXI, fig. 6), from which a long tuft of swimming- 
bristles (Plate LXXIX, fig. 24 a) projects. 

Reproduction. — The specimen procured on August 8th, in the Sound of Harris, carried 
female buds, all with ova except the minute ones next the nurse-stock. The latter had 
fully sixty segments, and there were ten buds. On the other hand, that procured in 
Bressay Sound, Shetland, in July, carried male buds. 

In life the species is one of the most beautiful and graceful in the family. Its 
brilliant orange spots, the translucency of its tissues, and its active movements as it trails 
the long series of finely-coloured buds behind it, make it as striking as it is interesting. 

Montagu's Nereis pinnigera (1808) is a nurse-stock with a developing bud 
posteriorly. He thought that at the posterior end it (the nurse-stock) " suddenly 
decreases and becomes very small, as if that part had been newly formed ; a circum- 
stance of no unreasonable conjecture, as it is well known that many of the Mollusca tribe 
are capable of reproduction." The Nereis maculosa of the same author appears to be a 
pelagic female bud of this species, 1 with swimming-bristles. Dr. Johnston made a 
separate family for it (Amytiacese). In his manuscript volume of drawings (1808) in 
the Linnsean Society, Montagu shows in Plate XVIII, fig. 2, under the name of Nereis 
doridea, a female bud, and in fig. 4 the species itself (nurse-stock) with buds. 

The Myrianida longissima of Audouin and Edwards 2 is probably the same form. 

De St. Joseph describes the proventriculus in M. maculata, Claparede, as barrel- 
shaped, with thirty-two rows of "oleaginous" dots, and occupying segments 13, 14, 

1 ' Trans. Linn. Soc./ xi, p. 21, Tab. hi, fig. 4, and also MS. vol. Drawings in the Linn. Soc., 
Plate XXXV, fig. 4. 



2 ' 



Ann. Sc. Nat./ t. xxix, p. 240. 



232 MYRIANIDA PINNIGERA. 

and 15. The proboscis is 4 mm. long, and has fifty to sixty pointed papillas at the edge. 
There are no lateral ponches or stomach. Little difference exists between this and the 
foregoing species. 

Malaquin a considers from his observations on three examples that the budding 
occurs after the sixty-sixth segment of the nurse-stock. The segment concerned is the 
penultimate one, and he terms it " Zoonite formateur" The pygidium of the nurse-stock, 
therefore, he found at the extremity of the twenty-ninth bud. With this segment 
{zoonite formateur) the new buds are connected. On the other hand, he has observed 
zoonites formateur s at a distance from the pygidium and independent of it. This author 
has studied the growth of the buds, the early segments of which present no appendages ; 
then minute processes appear in the succeeding segments, in the first or youngest bud on 
the pygidium (Malaquin). In the next series of segments forming a bud, these appen- 
dages are better developed, and so until in the third bud {e. g.) bristles occur, and the 
alimentary canal of the nurse-stock goes through each to the terminal bud. Each stolon 
possesses a zoonite formateur in front of the pygidium, the former having no appendages, 
the latter having them. Malaquin thinks the formative segment in the buds gives rise to 
additional segments in front. The anterior segment, on the other hand, is thickened 
dorsally and gives rise to head and post-cephalic segment. The author minutely describes 
each successive bud, and the appearance of the various appendages in it, including the 
eyes for dorsal and ventral vision, and the swimming-bristles. The terminal stolons — 
male or female — thus developed show great activity, and by-and-by break from the 
nurse-stock and swim freely away. The male bud (Polybostrichus) had thirty segments 
besides head, peristomium, pygidium, and the formative ring in front of it. Their 
sexual elements were ripe. The first four segments have no swimming-bristles, and 
contain the genital glands (four pairs). They have spots of brilliant orange-red 
pigment. The head bears dorsally a long median and two shorter lateral tentacles, and 
beneath are the split palpi, or, as he terms them, the anterior lateral antenna and the 
palpus. The peristomial segment has a long dorsal and a shorter ventral tentacular 
cirrus. 

The female bud (Sacconereis) has about thirty-five segments, and twenty-three bear 
swimming-bristles. The head carries a median and two lateral foliate tentacles. The 
post-cephalic (peristomial) has a single tentacular cirrus. The eyes are, as in the male, 
a larger anterior and a smaller posterior. The ovigerous sac occupies the fifteen or 
sixteen median segments, and has a median constriction. It is proportionally smaller 
than in Autolytus, and has exactly forty-three large orange eggs (Malaquin). The sac is 
a transparent secretion of the mucous glands of the ventral division of the foot. The 
colouration is similar to that of the male. The natatory bristles begin on the first foot, 
though occasionally they vary. These stolons, ■$ and ? , are devoid of a proboscis and 
proventriculus. 

• l ' Recherches sur les Syllidians/ pp. 287 — 305. 



ERRATA ET CORRIGENDA. 

PAGE 

2. Explanation of Fig. 35, for nephridispore read nephridiopore. 

6. „ „ 37, for below read above. 

8, line 14 from bottom, for Plate LXXVII read Plate LXXYI. 

17, „ 15, Plate XL111, for fig. 6 read fig. 3. 

17, „ 16, for Plate LXXVII read Plate LXXYI. 

20, „ Al, ,, „ „ ,, 

91 "1 & 

22, „ 24, (Me^e the before priority. 

23, „ 4 from bottom, for PI. LXXYII read PI. LXXYI. 

24, „ 21 from bottom, 1867, for Aimel. read Annul. 
26, „ 6, /o?- dilation read dilatation. 

29, „ 12 from bottom, for Plate LXXYII read Plate LXXYI, 

30, „ 7, 1867, for Annel. read Annul. 

31, lines 21 and 24, for Plate LXXYII read Plate LXXYI. 
49, line 6 from bottom, for Annel. read Annul. 

61, Synonyms, line 4, 1868, for Neap, read Nap. 

62, line 22, for bacilliporous read bacilliparous. 

75, „ 10 from bottom, for rev. read etc., and for Mgrm. read Mgrn. 
99. Explanation of Fig. 43, for spetzbergensis read spetsbergensis. 
104. Head-line and line 15 from bottom, ditto. 

104, line 18, for Mgm. read Mgrn. 

105. Head-line, for sjpetzbergensis read spetsbergensis. 
108, „ 7, for preparation read preparations. 

Ill, „ 13, for umbricate read imbricate. 

114, „ 11 from bottom, for organs read organ. 

130. Foot-note 1,/or Annel. Neap, read Annel. Nap. 

13^ 2 

j- <-»*-'• j) ^i •)■> )) )) >) 

142, „ 4, „ 

143, line 23, for moniliconis read monilicomis . 

144, ,, 19, for cliitine read chitin. 
154, „ 18, for Long. 55 read Log 55. 
161, „ 3, for epithelium read coat. 
170, „ 11, insert a comma after bilobed. 
178, ,, 4, delete and in rear of. 

197, and in succeeding entries under Synonyms, for Annel. Nap. read Annel. Nap. 

214, line 18, insert a comma after front. 

228, „ 11 from bottom, insert apparently before devoid. 

228, „ 7 „ delete the comma after reproduction. 

232. Foot-note, for Syllidians read Syllidiens. 



PLATE XLIIL 

Fig. 

1. Nephthys caeca, 0. Fabr., from the dorsum. About natural size. The example is a 

little more deeply coloured than usual. 

2. N. caeca, var. ciliata (epitokous form?), the K longisetosa of Johnston, from the 

dorsum. Slightly enlarged. 

3. Nephthys Hombergii, Lamk., from the dorsum. The filament at the tip of the tail is 

unusually long. Slightly enlarged. 

4. Anaitis rosea, Mcintosh. St. Andrews. From a drawing by Eliza Lorimer. It is 

incomplete posteriorly. Enlarged. 

5. Eulalia bilineata, Johnst., var., from the tidal region, St. Andrews, July 1st, 1904. 

Enlarged. 

6. Eulalia nebulosa, Mont. Drawing of a specimen from Plymouth by Mrs. Sexton. 

X 2. 

7. Eulalia viridis, 0. F. Mull., var. = E. aurea, Gravier, from Plymouth. From 

a drawing by Mrs. Sexton. X 3. 

8. Eulalia? brick red, Dr. Allen, Plymouth (six specimens found July 26th, 1904). 

Enlarged. 

9. Eumida sanguinea, (Erst. The body shows various constrictions and dilatations. 

Enlarged. 
10. Eumida sanguinea. A few segments, behind the middle, of a variety from Plymouth 
having a dull orange hue. Enlarged. 



Pl.XLII; 




-*•- 
fe 






\N 



■/, 




10. 










PLATE XLIV. 

Fig. 

1. Eulalia viridis, 0. F. Mull., from the East Kocks, St. Andrews. Enlarged. 

2. Gelatinous cocoon with ova of the foregoing (?) appended to a fragment of Fucus 

serratus by a cord of mucus, from St. Andrews (in March). Enlarged. 

3. Eulalia tripunctata, Mcintosh, from St. Andrews. Considerably enlarged. 

4. Eumida sanguinea, (Erst., with white bar at collar and white touches, from Plymouth; 

drawn by Mrs. Sexton. Enlarged. 

5. Eulalia viridis, 0. E. Mull., var. omata, De St. Joseph, from Plymouth; drawn by 

Mrs. Sexton. Enlarged. 

6. Phyllodoce Paretti, De Blainv. Head and anterior segments, from the dorsum. From 

Plymouth. Enlarged. 
6a. Middle segments of the foregoing, viewed laterally. 
6b. Middle segments from the dorsum. 
6c. Tail of the foregoing, recently reproduced. 



Pl.XLIV. 







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LithWernerk Winter, Frankfort 



PLATE XLV. 

Fig. 

1. JSTotophyllum foliosum, Sars, showing the dorsum, and, posteriorly, the ventral aspect 

of the tail (which is incomplete), from St. Andrews. Enlarged. 

2. Phyllodoce maculata, Linn., in full activity, from St. Andrews. Enlarged. 

3. Eteone picta, De Quatref., from St. Andrews. Enlarged. The red tints are less 

bright than originally. 



PL XLV. 










M *<** 









W *> v ZS 





# <* -fir 1 






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R.M ad nat. fecit. 



Litri.Weirier&¥iii[er,'Fr-;i!ii-:[ ... 



PLATE XLVI. 

Fig. 

1. Pionosyllis prolifera, Krohn (?), from St. Andrews. Enlarged. 1 

2. Gastalia punctata, 0. F. Mull., from the deeper water off St. Andrews Bay. Enlarged. 

3. Gastalia fusca, Johnst., from the tidal water, St. Andrews. The tail is incomplete 

and the bristles are absent from a few feet on the left anteriorly. Enlarged. 

1 There is doubt in connection with this figure, the preparation of which has been lost. The 
dorsal cirri showed no articulations in life. 



P1.XLV1 







I 



Li th .Werner & Winter Frai ikfort°/M . 



PLATE XL VII. 

Fig. 

1. Autolytus pictus, Ehlers, male bud (Polybostrichus), from St. Andrews. The bifid palpi 

have the limbs approximated, so that the separation is scarcely visible. Enlarged. 

2. Phyllodoce lamelligera, Linn., from a specimen at St. Andrews. X about 2. The 

majority are considerably darker. 

3. Phyllodoce maculata, Linn., portion of the dorsum of an example of a deep greenish 

hue, in March, from St. Andrews. Enlarged. 

4. Phyllodoce rubiginosa, De St. Joseph, from the dorsum, from Plymouth. A few of the 

dorsal cirri behind the head have been injured. Enlarged. 

5. Middle segments of the above, from Plymouth. Enlarged. 



Pl.XLVII. 



\ Hi I 




Fig.lR.M'. Fig-.2R.M.& A.H.W. 
Castera A.H.W. 



PLATE XLVIII. 

Fig. 

1. Eumida sanguinea, (Erst., from St. Andrews. Enlarged. 

2. Variety of the foregoing, with a white touch behind the head, and with transverse 

white bars, from Plymouth. Enlarged. 

3. Eteone spetsbergensis (?), from the mouth of the Bay of St. Andrews. Enlarged. 

4. Amblyosyllis lineata, Grrube, from Lochmaddy, North Uist. Lived at Murthly for a 

month or two. Enlarged. 
b. Sylline rubropunctata, Grrube, from a Plymouth example. Enlarged. 
6. Autolytus prolifer, 0. E. Mull., animal surculare. Enlarged. 




Fig.4.R.M.del. 
Castera A.H.W. 






Lith Werner kWinterTrarikfort °/M . 



PLATE XLIX. 

Fig. 

1. Phyllodoce lamelligera, Linn., a variety, from Plymouth Sound, September 30th, 1905. 

2. Dorsal aspect of a few segments from the middle of the foregoing. Enlarged. 

3. Odontosyllis gibba, Clap., dorsal view of an example from Plymouth. 

4. Middle segments of the foregoing, from the dorsum. Enlarged. 

5. Odontosyllis fulgurans, Aud. & Ed., from Plymouth. The tail is incomplete. Enlarged. 

6. Syllis Krohnii, Ehlers, from the tidal rocks, Lochmaddy, North Uist. It lived for 

months at Murthly. Enlarged. 

7. Autolytus, deeply tinted like pictus, but probably prolifer, pelagic female, with eggs 

in ventral sac. Enlarged. 

8. Nurse-stock of Autolytus pictus, Ehlers, from the West Pocks, St. Andrews. Enlarged. 



PI. XILIX 




.M. fecit 
Castera A.H.W. 



Iith .Werne r <S Win ter. Frankfor t °/M . 



PLATE L. 

Fig. 

1. Early stage in the development of Nephthys, October 23rd, 1897. X 100. 

2. An older stage of the same date. X 100. 

3. Specimen captnred a few days later (October 28th) showing certain alterations of the 

head and body. X 100. 

4. Head of Eulalia bilineata, Johnst. Enlarged. 

5. Outline of the head and anterior region of Eumida sanguinea, from St. Andrews. 

Enlarged. 

6. Young Phyllodoce, from St. Andrews, June 15th, 1898. Enlarged. 

7. Dorsal cirri of a variety of Phyllodoce lamelligera, Linn., from Plymouth. Enlarged. 

8. Head of Phyllodoce rubiginosa, De St. Joseph, from a Plymouth specimen. Enlarged. 

9. Head of Trypanosyllis zebra, Grube, from Plymouth. Enlarged. 

10. Tail of the foregoing. Enlarged. 

11. Pelagic male Autolytus (= Polybostrichus) . Enlarged. 

12. Larval Nereis (Alitta) virens, from St. Andrews Bay, October 26th. X 50. 

13. Older larval stage apparently of the same form, from St. Andrews. X 45. 
14 and 15. Dorsal and front views of larval Nereis, from St. Andrews. X 100. 

16. Post-larval Nereis (Alitta ?), September 26th, St. Andrews. X 40. 

17. Head of Nereis irorata, Grube, from Plymouth. Enlarged. 



PL L. 



io 



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thm ; 






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17. 



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15 





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13. 



















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PLATE LVIL 



Fig. 

1. Head of Nephthys eseca, 0. Fabr., from the dorsum. Enlarged. 

2. Under surface of head and anterior region of the same. Enlarged. 

3. Extruded proboscis of the foregoing — from the ventral surface. Enlarged. 

4. Extruded proboscis of the same form viewed from the free end. Enlarged. 

5. Proboscis slit longitudinally and exposing the teeth. Enlarged. 

6. Head of Nephthys Hombergil, Lamk., from the dorsum. Enlarged. 
6a. Extruded proboscis of the same species from the dorsum. Enlarged. 

7. Under surface of the foregoing, showing the mouth and the parts behind it. Enlarged. 
7 a. Head of the same with eyes from St. Magnus Bay (Dr. Gwyn Jeffreys). Enlarged. 

8. Head and exserted proboscis of Nephthys hystricis, McL, from the dorsum. Enlarged. 

9. Lateral view of the same proboscis. Enlarged. 

10. Dorsal view of the head and proboscis of Nephthys longisetosa, (Erst. Enlarged. 

11. Proboscis of the same partially extruded. Enlarged. 

12. Proboscis of the foregoing wholly withdrawn. Enlarged. 

13. Head of Nephthys Grubei, McL, from the dorsum. Enlarged. 

14. Ventral surface of the foregoing. Enlarged. 

15. Head of Nephthys Johnstoni, Ehlers, viewed from the dorsum. Enlarged. 

16. Ventral surface of the same. Enlarged. 

17. Head of Nephthys cirrosa, Ehlers, with extruded proboscis, from the dorsum. 

Enlarged. 

18. The same from the ventral surface. Enlarged. 

19. Head and anterior region of Nephthys incisa, Mgrn., from Graiway. Enlarged. 

20. Ventral surface of the foregoing showing the mouth. Enlarged. 



PL LVII. 



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PLATE LVIII. 

Fig. 

1. Head of Eidalia nebidosa, Mont., from the dorsum. Enlarged. 

2. Head of the foregoing with the proboscis extruded. Enlarged. 

3. Head of Uulalia viridis, 0. E. M., with the proboscis extruded. Enlarged. 

4. Head of Eulalia macroceros, Grube. Enlarged. 

5. Head of Phyllodoce groenlandica, (Erst., from St. Andrews. Enlarged. 

6. Head of Anaitis Kosteriensis, Mgrn. Enlarged. 

7. Head of Anaitis Jeffreysii, McL, from Valencia Harbour. Enlarged. 

8. Head of Genetyllis hibernica, McL Enlarged. 

9. Head of Eteone arctica, Mgrn., from H.M.S. ' Valorous.' Enlarged. 

10. Head of Mysta barbata, Mgrn., dredged off the Isle of Man. Enlarged. 

11. Head of Mystifies Lizzise, n.s., from Cape Sagres, 'Porcupine,' 1870. Enlarged. 

12. Head of Gphiodromus flexuosus, D. Ch. Enlarged. 

13. Anterior end of the foregoing with the proboscis extruded. Viewed from the ventral 

surface. Enlarged. 

14. Head of Gastalia arctica, Mgrn., from west coast of Ireland. The frontal tentacles 

have been lost. Enlarged. 

15. Dorsal surface of the head of Gastalia fusca, Johnst. Enlarged. 

16. Ventral view of the same species with the proboscis exserted. Enlarged. 

17. Head of Leocrates atlantica, Poule. The specimens unfortunately were injured and 

imperfectly preserved. Enlarged. 

18. Head of Dalhousiella Garpenteri, McL, from an indifferent specimen. Enlarged. 

19. Head (imperfect) of " Sph&rosyllis ovigera, Lghns.," from Dr. Allen. X about 35. 

20. Head and extruded proboscis of Phyllodoce lamelligera, L. (drawn by Mrs. Sexton). 

Enlarged. 

21. Head of Phyllodoce maculata, L., with proboscis extruded (drawn by Mrs. Sexton). 

Enlarged. 

22. Head of Eumida sanguined, (Erst., with proboscis exserted (drawn by Mrs. Sexton). 

Enlarged. 

23. Head of Eulalia tripunctata, McL, with proboscis thrust out (drawn by Mrs. Sexton). 

x 17. 

24. Abnormal tail of Phyllodoce maculata; L., with one large and four smaller anal cirri 

in pairs, from St. Andrews. Enlarged. 

25. Tip of tail of Phyllodoce rubiginosa, De St. Joseph, from Plymouth. Enlarged. 



PLLVIII. 




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PLATE LIX. 

Fig. 

1. Head and anterior region of Mag alia per armata, De St. Joseph. Enlarged. 

2. Candal region of the same. Enlarged. 

3. Female of Splixrosyllis hystrix, Clap., with ova attached to the ventral surface. 

Ventral view. Enlarged. 

4. Male example of the foregoing with long (swimming-) bristles, from the coast of 

Co. Down, Ireland. Enlarged. 
4a. Head of the foregoing species. X Zeiss oc. A, obj. 4, with draw-tube. 

5. Exogone gemmifera, Pagenstecher, from St. Andrews, with flask-shaped embryos 

attached to feet on the ventral surface ; c natural size, a and b enlarged. 

6. Proboscis of the same and the proventriculus. Enlarged. 

7. Xenosyllis Kinbergi, n.s., from the dorsum. Enlarged. 

8. Head of Sphserosyllis hystrix, Clap., with an additional eye-speck on each side in 

front. The tentacles are incomplete. Enlarged. 

9. Anterior region of Mlcrosyllis Marenzelleri, n.s. (fragmentary), from the Sound of 

Harris, x about 40. 

10. Head of Pionosyllis prolifera, Krohn, var. Malmgreni. Enlarged. 

11. Anterior region of Eusyllis tubifex, Gosse, in life, showing the arrangement of the 

proboscis, proventriculus, and stomach. X about 45. 

12. Head of Pionosyllis divaricata, Kef erst. Enlarged. 

13. Head of Eusyllis tubifex, Gosse. Enlarged. 

14. Head and anterior region of Odontosyllis gibba, var. Robertianse, in life, from Guernsey. 

Enlarged. 
14a. The same in spirit. Enlarged. 

15. Head and anterior region of Odontosyllis fulgurans, Aud. and Ed. Enlarged. 
15a. Tail of the same. Enlarged. 

156. Tail (incomplete) of example of the foregoing from Plymouth. Enlarged. 

16. Head of Ioida from Plymouth, September 14th, 1905. Enlarged. 
16a. Tail of the foregoing. Enlarged. 



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PLATE LX. 

Fig. 

1. Head of Syllisarmillaris, 0. F. M. Enlarged. 

2. Head of Ioida macrophthalma, Johnst. Enlarged. 

3. Head of Syllis cucullata, Mel. Enlarged. 

4. Early larva of Autolytus prolifer, 0. P. M., from the sac. X 100. 

5 and ha. Anterior and posterior ends of free bud of Autolytus, St. Andrews, May 25th, 

1885, the former from the ventral surface. Magnified. 

6 and 6a. Dorsal and ventral aspects of the head and proboscis of Nereis pelagica, L. 

Enlarged. 

7 and 7a. Dorsal and ventral views of the head and proboscis of Nereis cultrifera, Grube. 

Enlarged. 

8. Head of Nereis irorata, Mgrn., from the dorsum. Enlarged. 

9. Head and proboscis of Nereis Marionii, Aud. and Ed., from the dorsum. Enlarged. 

10. Proboscis of Nereis Dtimerilii, Aud. and Ed., from the ventral surface (partly 

extruded). Enlarged. 
10a. Proboscis of the same from the ventral surface (from St. Andrews). Enlarged. 
106. Head and proboscis in an epitokous form, in which the eyes are much larger than 

usual. Enlarged. 
10c. Epitokous male from the surface of the sea, Castle Bay, Barra. Enlarged. 

11. Head and proboscis of Nereis diversicolor, 0. F. M. Enlarged. 
11a. Ventral aspect of the foregoing. Enlarged. 

12. Head of Marphysa Belli, Aud. and Ed. Enlarged. 



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PLATE LXI. 

Fig. 

1. Head of Eunereis longissirna, Johnst., from St. Magnus Bay, Shetland. Enlarged. 
la. Head and proboscis of the same from Montrose Bay. Enlarged. 

lb. Ventral patches of teeth (paragnathi) on the proboscis of la. Enlarged. 

2. Dorsal view of head and proboscis of Nereis (Alitta) virens, Sars. Enlarged under a 

lens. 
2a. Ventral aspect of the foregoing. Enlarged. 

3. Coelomic corpuscles of this species. X 700. 

4. Contracted muscular fibril showing peculiar cross-striations. X 700. 

5. Another contracted fibril with a granular band in centre, and transparent sarco- 

lemma (b) externally. X 350. 

6. Dorsal view of head and proboscis of Nereilepas fucata, Savigny. Enlarged. 
6a. Ventral view of the same. Enlarged. 

7. Upper dental apparatus of Staurocephalus rubrovittatus, Grube, from Herm. Seen 

from below. X about 40. 
7a. Shorter teeth of the same, x Zeiss oc. 2, obj. D. 
7b. Longer (outer) upper teeth. X Zeiss oc. 2, obj. D. 
7c. Mandible of the foregoing, x 55. 

8. Mandible of Staurocephalus Kefersteini, Mel. X Zeiss oc. 2, obj. D. 

9. Dental (upper) apparatus of Staurocephalus ciliatus, Keferst., from St. Peter Port, 

Guernsey. The specimen is incomplete. X Zeiss oc. 2, obj. D. 
9a. Mandible of the foregoing, x Zeiss oc. 2, obj. D. 
10. Young example of Ojohryotrocha puerilis, Clap, and Mecz. Enlarged. 
10a. Dental apparatus of the same. X Zeiss oc. 2, obj. D. Eeduced from original 

drawing. 



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PLATE LXIL 



Fig. 

1. Dental apparatus of Lumbriconereis fragilis, 0. F. M., viewed from above. X about 14. 
la. Mandibles. X about 14. 

2. Mandibles of Lumbriconereis impatiens, Clap. X 35. 

3. Head of Lumbriconereis hibemica, n.s., twenty-five miles off the Blasquet. Enlarged. 
3a. Proboscis extruded. Enlarged. 

4. Head of Lumbriconereis gracilis, Grube, from the dorsum. From Loch Slyne, Co. 

Cork. Enlarged. 
4a. Armature of the proboscis as extruded. X 41 . 

5. Head of Lumbriconereis G. Enlarged. 

6. Head of Zy globus laurentianus, Grube. Enlarged. 
6a. Dental apparatus. x about 50. 

66. Mandibles, x about 50. 

7. Head of Drilonereis JElisabethse, n.s., with the proboscis partly protruded. From the 

stomach of a haddock, St. Andrews. Enlarged. 
7a. Dental apparatus. X 20. 
7b. Mandibles of the same. X Zeiss oc. 2, obj. A. 

8. Ventral surface of the head of Arabella tricolor , Mont., from Gruernsey. Enlarged. 
8a. Dental apparatus. X 20. 

Sb. One side of the same in another example. X about 40. 
8c. Mandibles. X about 25. 

9. Notocirrus scotieus, McL, from the dorsum. From Lochmaddy. Enlarged. 

9a. Dental apparatus drawn from an example from the Isle of Man, off Peel. The 
anterior fang of the maxillary plates (second from rear) was broken off. X 50. 

9b, Mandibles of the same. X 50. 
10. Dental apparatus of Eunice Harassii, Aud. and Ed. x 30. 
10a. Mandibles of the same. X 30. 






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PLATE LXIIL 

Pig. 

1. Dental apparatus of Marphysa sanguined, Mont. X 15. 
la. Mandibles. X 15. 

2. Dental apparatus of Marphysa Belli, Aud. and Ed. X 15. 

3. Dental apparatus of Eunice vittata, D. Ch. (=limosa, Ehlers), from G-alway. X Zeiss 

oc. 2, obj. A. 
3a. Mandibles. X Zeiss oc. 2, obj. A. 

4. Dental apparatus of Eunice norvegica, L. Prom Norway. X about 30. 
4a. Mandibles of ditto. X 55. 

5. Dental apparatus of Nematonereis unicornis, Grrube. Incomplete. X 21. 
ha. Mandibles (unfortunately imperfect). X 21. 

6. Dental apparatus of Lysidice Ninetta, Aud. and Ed. X 20. 
6a. Mandibles. X 20. 

7. Head of Onuphis britannica, Mel. Enlarged. 
la. Dental apparatus. X 50. 

7b. Mandibles. X 50. 

7c and 7cl. Tubes of this species composed of shells and secretion. Enlarged. 

8. Dental apparatus of Onuphis brevibrachiata, Ehlers. X 20. 
8a. Mandibles. X 20. 

9. Dorsal view of Onuphis conchylega, Sars, in its tube. Enlarged. 

10. Head aud anterior region of Onuphis brevibrachiata, Ehlers, after figure by Ehlers 

(Taf. Ill, fig. 2). Enlarged. 
10(2. Ventral surface of the same, after Ehlers. Enlarged. 



Pl.LXIIL 



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PLATE LXIV. 

Fig. 

1. Dental apparatus of Onuphis conchylega, Sars. Prom the ' Knight Errant.' X 30. 
la. Mandibles. X 30. 

2. Dental system of Onuphis quadricuspis, 0. Pabr. x 30. 
2a. Mandibles. X 30. 

3. Head of Onuphis (Diopatra) fragosa, Ehlers. Enlarged. 
3a. Dental apparatus. X Zeiss oc. 2, obj. A. 

3b. Portion of tube (of mud) from Norway. Enlarged. 

4. Ventral surface of the head of Onuphis sicula, De Quatref. Enlarged. 
4a. Dental apparatus of the foregoing. X as before. 

4b. Mandibles of the same. X as before. 

4c. Tube of the same from Plymouth. Enlarged. 

5. Head of Hyalinoecia tubicola, 0. P. M., from the mouth of Bantry Bay. Enlarged. 
ha. Dental apparatus. X 21. 

hb. Hyaline tube containing annelid. Enlarged. 

6. Anterior end of Goniada maculata, (Erst. Enlarged. 
6a. The same with proboscis extruded. Enlarged. 

6b. B,ow of V-shaped denticles of the proboscis. X 100. 

6c and 6c' . Intermediate small denticles, x Zeiss oc. 2, obj. D. 

6d. Larger denticle somewhat palmate in outline. X Zeiss oc. 2, obj. D. 

7. First and part of second region of the body of Ghjcinde Nordmanni, Mgrn. Enlarged, 
7a, Tail. Enlarged. 

7b. Smaller denticles of Glycinde Nordmanni. x Zeiss oc. 2, obj. D. 

7V and 7b" . Larger denticles. X Zeiss oc. 2, obj. D., with 1 in. draw-tube. 

8. Proboscis (exserted) of Glycera siphonostoma, D. Oh., from Balta. Enlarged. 
8a. Tip of a regenerated tail in an example from Herm. Slightly enlarged. 

9. Tooth of Glycera capitata, (Erst. X 380. 

9a. Tooth of Glycera lapidum, De Quatref. X 350. 



Pl.LXIV. 




A.H.VA del 



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PLATE LXV. 



Fig. 

1. Terminal region of the extruded proboscis of Glycera alba, H. Rathke. Enlarged. 

2. Anterior end of Glycera Goesi, Mgrn. (somewhat contracted). Enlarged. 
2a. Proboscis extruded. Enlarged. 

3. Head of Aricia norvegica, Sars. Enlarged. 

4. Head of Scoloplos armiger, 0. F. M. Enlarged. 

4a. Tip of tail, from Symbister Harbour, Shetland. Enlarged. 

5. Dorsal view of the head of Naidonereis quadricuspida, 0. Fabr. Enlarged. 
5a. Ventral view. Enlarged. 

6. Head of Eteone pusilla, (Erst. Enlarged. 

7. Head of Eunice floridana, Pourtales. X about 15. 
7a. Upper dental apparatus of the foregoing. X 12. 
7b. Mandibles of the same. X 15. 

8. Upper dental apparatus of Lumbriconereis impatiens, Clap, x about 21. 
Sa. Mandibles of the foregoing. X 15. 

9. Foot of Nematonereis unicornis, Grrube. X 90. 

10. Proboscis of Magalia perarmata, Mar. and Bobr., from the figure given by these 

authors. Enlarged. 

11. Upper dental apparatus of Marphysa Belli, Aud. and Ed. X 50. 
11a. Mandibles of the foregoing. X 50. 

12. Upper dental apparatus of Lumbriconereis Nardonis, Aud. and Ed. x 50. 
12a. Anterior region of the mandibles of the same. X 50. 

13. Upper dental apparatus of Onuphis britannica, McL, in which reproduction of the 

maxilla3 has occurred. X Zeiss oc. 2, obj. A. 

14. Ninetieth foot of Marphysa sanguinea, Mont. X 55. 

15. View of the mouth and adjacent parts of Hyalinoecia tubicola, 0. F. M., from life. 

Enlarged. 

16. Foot of Exogone gemmifera, Pag., after Viguier. 

17. Mandible of Staurocephalus rubrovittatus, Grrube. X Zeiss oc. 2, obj. D. 

18 and 18a. Isolated teeth of Staurocephalus Kefersteinii, McL X Zeiss oc. 4, obj. 0. 







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PLATE LXVI. 

Fig. 

1. Anterior foot of Nephthys cdeca, 0. Fabr., from the front. X 50. 

2. Posterior foot of the same from the front. X 50. 

3. Anterior foot of the variety ciliata (N longisetosa of Johnst.). May 17th. X 

about 50. 

4. Anterior foot of Nephthys Hombergii, Lamk. Similarly magnified. 

5. Anterior foot of Nephthys Hombergii var. Kersivalensis. X about 50. 

6. Posterior foot of the same. Similarly magnified. 

7. Anterior foot of N Hombergii var. vasculosa, from twenty-five fathoms, Buono Bay, 

6 Porcupine.' 

8. Posterior foot of the same. X 90. 

9. Anterior foot of Nephthys ciliata, 0. F. M. X 50. 

10. Anterior foot of Nephthys hystricis, Mel. 

10a. The same viewed from the edge. X about 50. 

11. Anterior foot of Nephthys longisetosa, (Erst. Enlarged. 



Pl.LXVI. 




















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PLATE LXVIL 



Fig. 

1. Anterior foot of Nephthys Grubei, n.s. X 40. 

2. Typical foot of Nephthys Johnstoni, Ehlers, from Balta. X 50. 

2a. Foot from the same region of an example from the West Sands, St. Andrews. X 50. 

3. Typical foot of Nephthys cirrosa, Ehlers. X 50. 

4. Typical foot of Nephthys incisa, Mgrn., from Connemara. X 55. 

5. Foot of Notophyllum folio sum, Sars. X 55. 

6. Foot of Eulalia bilineata, Johnst. X 100. 

7. Spermatozoa of foregoing. X 400. 

8. Tenth foot of Eulalia nebulosa, Mont. X 90. 

9. Fortieth foot of the same. X 90. 

10. Tenth foot of Eulalia viridis, 0. F. M., St. Andrews. X 210. 

11. Fortieth foot of the same. X 210. 

12. Anterior foot of Eulalia (Pterocirrus) macroceros, Grube. X 35. 

13. Anterior foot of Eulalia tripunctata, McL, from Herm. X 210. 

14. Tenth foot of Eumida sanguinea, (Erst., from Lochmaddy. X 90. 

15. Sixtieth foot of the foregoing. X 90. 

J 6. Twentieth foot of Marphysa Belli, Aud. and Ed., from Plymouth. X 15. 



PLLXVII. 




E -L.U.H.W.del 



f*P.WM.Trap impi 



PLATE LXVIII. 



Fig. 

1. Tenth foot of Pliyllodoce lamelligera, L., from the West Voe, Scalloway. X 55. 

2. Sixtieth foot of the same specimen. X 55. 

3. Foot (about sixtieth) of an example from Naples. X about 90. 

4. Tenth foot of Pliyllodoce groenlandica , (Erst., from St. Andrews. X about 60. 

5. Sixty-seventh foot of the same. X 60. 

6. Posterior foot of the foregoing example. X 60. 

7. Tenth foot of Pliyllodoce maculata, L., from St. Andrews. X about 102. 

8. Sixtieth foot of the foregoing. X about 102. 

9. Foot of Pliyllodoce rubiginosa, De St. Joseph. X about 55. 

10. Tenth foot of Pliyllodoce vittata, Ehlers, from St. Magnus Bay. X 210. 

11. Fiftieth foot of the foregoing. X 210. 

11a. Foot of the same form, middle of body. X about 180. 

13. Foot of Anaitis rosea, Mel. X about 150. 

14. Tenth foot of Anaitis Jcosteriensis, Mgrn. X 90. 

15. Thirty-fifth foot of the same. X 90. 

16. Tenth foot of Anaitis Jeffrey sii, Mel. X 210. 

17. Fortieth foot of the same. X 210. 

18. Tenth foot of Genetyllis lutea, Mgrn. The ventral cirrus is seen obliquely. X 

about 55. 

19. Fiftieth foot of the same. X about 55. 

20. Tenth foot of Genetyllis citrina, n.s. X 55. 

21. Sixtieth foot of the foregoing. X about 60. 



PI. IXVIIL 




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PLATE LXIX. 

Fig. 

1. Tenth foot of Genetyllis hibernica, Mel. X 90. 

2. Fortieth foot of the same. X 90. 

3. Twelfth foot of Eteone picta, Be Quatref. X 210. 

4. Thirtieth foot of the foregoing. x 210. 

5. Tenth foot of Eteone spetsbergensis, Mgrn. X 50. 

6. One hundredth foot of the same. X 50. 

7. Seventieth foot of Eteone pusilla, (Erst. X 50. 

8. Tenth foot of Eteone arctica, var. Bobertiana, from the West Rocks, St. Andrews. X 50. 

9. Seventieth foot of the foregoing. X 50. 

10. Foot of Mysta barbata, Mgrn. X about 90. 

11. Tenth foot of Mystides Lizzise, n.s. West Sands, St. Andrews. X 90. 

12. Fiftieth foot of the same. X 90. 

13. Typical foot of Ophiodromus flexuosus, D. Ch. : a, dorsal lobe ; b, ceratophore of 

dorsal cirrus ; c, long papilla of superior division ; d, long papilla of the ventral 
division ; e, ventral cirrus. X about 100. 

14. Anterior foot of G as talia punctata, 0. F. M. X 90. 

15. Anterior foot of Gastalia arctica, Mgrn. X about 100. 

16. Anterior foot of Gastalia fusca, Johnst. X about 100. 

17. Foot of Leocrates atlantica, Roule. X about 35. 

18. Foot of Mag alia per armata, De St. Joseph. X 50. 



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PLATE LXX. 

Fig. 

1. Foot of Sphxrosyllis hystrix, with swimming-bristles, from sea off Co. Down (Professor 

G. Brady), x 180. 

2. Foot of Xenosyllis Kinbergi, n.s. X about 50. 

8. Foot of Microsyllis Marenzelleri, n.s., from the Sound of Harris. X 250. 

4. Foot of Pionosyllis prolifera, Krohn, from Guernsey. X about 50. 

5. Foot of female bud of the foregoing, with swimming-bristles, from Scalloway. A 

papilla appears at the tip dorsally, and thus the example differs from the type. 
X about 50. 

6. Foot of variety (?) of the same form, from Herm. X about 50. 

7. Foot of Pionosyllis divaricata, Keferst. X about 50. 

8. Foot of Trypanosyllis zebra, Grube. X 50. 

9. Foot of Eusyllis tubifex, Gosse. X 50. 

10. Foot of bud of Autolytus prolifer, from St. Andrews. The swimming-bristles are 

developing, x 50. 

11. Foot of Odontosyllis fulgiirans, Clap. X about 50. 

12. Foot of Odontosyllis ctenostoma, Clap. X about 50. 

13. Foot of Odontosyllis gibba, var. Robertianse. X about 50. 

14. Foot of Syllis armillaris, O. F. M. X about 50. 

15. Foot of Ioida macrophthalma, Johnst., with long swimming-bristles. X about 50. 

16. Foot of Syllis comuta, var. Gollingsii, off Sark. x about 50. 

17. Foot of Syllis cucidlata, n.s. Luccomb Chine. X about 50. 

18. Foot of Syllis macroceras, Grube, from "Whitecliff Bay, Isle of Wight. X about 50. 

19. Foot of Syllis Ounninghami, n.s. X Zeiss oc. 2, obj. A, 1 in. draw-tube. 
19a. Foot of bud of the foregoing. X about 50. 

20. Foot of Syllis brevicirrata, n.s., from Herm. X about 50. 

21. Foot of pelagic form of Syllis spongicola, Grube. X about 50. 

22. Foot of Syllis comuta, H. Rathke, from St. Magnus Bay, Shetland. X about 50. 
22a. Foot of variety of the same form (off the Hebrides). X about 50. 

22b. Foot of another specimen from Donegal Bay. X about 50. 

23. Foot of Syllis Busldi, n.s. X about 55. 

24. Foot of Pionosyllis (Syllis) hyalina, Grube. X about 55. 

25. Anterior foot of Syllis gracilis, Grube. X about 55. 

2ba. Foot of the foregoing from the posterior part of the middle region of the body. 
X about 55. 

26. Foot of Sylline rubropunctata, Grube. X Zeiss oc. 4, obj. A. 



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BAY SOCIETY. 



(INSTITUTED 1844) 



OFFICERS AND COUNCIL, 



1907. 



president : 

The Right Hon. LORD AVEBURY, D.C.L., LL.D., F.R.S, Pres.S.A., For.Sec.R.A, F.L.S., F.G.S., 

ETC. 



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{Treasurer : 

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Secretary : 

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