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THE

JOURNAL

THE LINNEAN SOCIETY.

ZOOLOGY.

WoL, 24.

LONDON:
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON HOUSE,
AND BY
LONGMANS, GREEN, AND CO.,
AND
WILLIAMS AND NORGATE.
1890.

Dates of Publication of the several Numbers included in this Volume.

No. 116, pp. 1- 61, published October 27, 1886.
elds, a 55> Ol=107- 3 June 30, 1887.
oe lis. 55 LOT Ae ms October 29, 1887.
5 219, 4, LEH 188; i November 1, 1888.
» 120, ,, 189-274, * September 15, 1888.
5) Lal; 4,. 270-298; = January 31, 1889.
,, 122, ,, 299-394, - August 31, 1889.
» 123, ,, 3895-472, * December 31, 1889.
, |

, 125, 473-592, ss, July 31, 1890.

PRINTED BY TAYLOR AND FRANCIS,
RED LION COURT, FLEET STREET.

LIST OF PAPERS.

Page
Baty, Josupy §., F.L.S.

Descriptions of a new Genus and of some new Species of Gale-
rucine ; also Diagnostic Notes on some of the older described
MERRIE SWIMEODIIOTE oo circa 5 5.6 nie ence span cistnesnces 1

Descriptions of some Genera and Species of Galerucin@ ...... 156

BousrrExp, Epwarp, L.R.C.P. Lond. .
The Natural History of the Genus Dero. (Communicated by
eeeemiuiie tsi.) (riates TIT —-V.) 20... osc cece sence 91

Day, Dr. Francis, C.LE., F.L.S.
On the Loch-Leyen Trout (Salmo levenensis) .... 64. cece ees il

Duncan, Prof. P. Martin, F.R. 8. , F.LS., and W. Percy SuapEN,
F.G.S., Sec. Linn. Soc.
On fi Anatomy of the Perignathic Girdle and of other Parts
of the Test of Discoidea cylindrica, Lamarck, sp. (With 10
NI, 0 DE Gack hicks Shs ees eer sissence 48

GuLick, Rey. Joun Tuomas.

Divergent Evolution through Cumulative Segregation. (Com-
municated by Alfred Russel Wallace, F.L.S.) ............ 189

Hoye, W. E., M.A. (Oxon), F.R.S.E., Keeper of the Manchester
Museum.
On the Deep-water Fauna of the Clyde Sea-area. (Communi-
cated by John Murray, LL.D., F.L.S., &c.) (With Map:
ieee i.) ......... ten issn ne wan ase oe.» 449

iv
Page
Kirpy, W. F., Assistant in the Zoological Department, British
Museum.

A Synopsis of the Genera of Chalcidide, Subfamily Eucharine ;
with Descriptions of several new Genera and Species of Chal-
cidide and Tenthredinide. (Communicated by Dr. J. Murie,
F.Li3.) (Plate.L):3°) ie % 2d bed meal ee

Lownek, Bensamin Tuompson, F.R.CS., F.L.S., Hunterian Pro-
fessor of Comparative Anatomy, Royal College of Surgeons.
On the Structure of the Retina of the Blowfly (Calkiphora
erythrocephala). (Plate XXVIL.) ......:500:s59s einen 406
On the Structure and Development of the Ovaries and their
Appendages in the Blowfly (Calliphora erythrocephala).
(Plate XXVIEIP). oo oa aS ee 418

LUBBOCK, Sir JoHN, Bart., M.P., D.C.L., LL.D., F.RS.
Observations on Ants, Bees, and Wasps.—Part XI. (With 2
UastrAtIONS:) oc se ives es ee 0 0 30 0 6 apg ee Keke © eee

Micaak., A. D., F.L.S., F.Z.S., F.R.M.S.
Researches into the Life-histories of Glyciphagus domesticus and

G.-spinipes, (Plate XVI.) . ... ss 00+ +00 +0015 017iphe eee 285
On some unrecorded Parasitic Acari found in Great Britain,
(Plate XXVIE) wicks s eae vee) ee er ... 400

Rak, Jonny, M.D., LL.D., F.RS.
Notes on some of the Birds and Mammals of the Hudson’s Bay
Company’s Territory, and of the Arctic Coast of America.
(Communicated ‘by G. J: Romanes, F.L:8:). "1.22. ae 136

Rroiny, H. N., M.A., F.LS.
Notes on the Zoology of Fernando Noronha. (Plate XXX.) .. 473

Ripiey; Stuart O., M.A., F.LS.
On the Characters of the Genus Lophopus, with a Description

of a New Species from Australia, (Plate II.) ............ 61
Romangs, Groree J., LL.D., F.R.S., F.LS.
Experiments on the Sense of Smell in Dogs ............00, , 66

; Page
SuurEtpt, R. W., M.D., C.M.Z.S., Captain, Medical Corps, U.S.

Army.
Studies of the Maerochires, Morphological and otherwise, with
the view of indicating their Relationships and defining their
several Positions in the System. (Communicated by Prof.
W. K. Parker, F.R.S., F.L.S.) (Plates XVII-XXIV.) .... 299

Sim, GEORGE. .

Occurrence of Lumpenus lampetriformis on the North Coast of

Scotland; with Notes on its Habits, Food, and the Ground

it frequents. (Communicated by Dr. Francis Day, F.L.S.)
(With 5 woodcuts.)......... Sg ee OS ee 38

SLADEN, W. Percy, F.G.S., Sec. Linn. Soc., and Prof: P. Martin
Duncay, F.R.S., F.LS.
On the Anatomy of the Perignathic Girdle and of other parts
of the Test of Discoidea cylindrica, Lamarck, sp. (With 10
EE SE CET Te eee eee eee Eee 48

SowERsy, G. B., F.L.S., F.Z.S.
Descriptions of fourteen new Species of Shells, (Plate XXV.) 395

THompson, Isaac C., F.R.M.S.
Copepoda of Madeira and the Canary Islands, with Descriptions
of New Genera and Species. (Communicated by Prof.
ere lao.) (Plates X—XI1). 1... esc ccc erie 145

WALKER, ALFRED O., F.L.S.

Notes on a Collection of Crustacea from Singapore. (Plates
NG ST cinrtewibiisl gs a Licinrikle dad bis siviavedeccaces 107

WatTERS, ARTHUR Wo., F.L.S., F.G:S.
On some Ovicells of Cyclostomatous Bryozoa. (Plate XIV.) .. 275
On the Ovicells of some Lichenopore. (Plate XV.).......... 280

.. Rerra of the Blowfly, illustrating B. T. Lowne’s paper.

EXPLANATION OF THE PLATES.

Species of Chalcidide and Tenthredinide.
Lornopus Lenpenrexpt, Ridley.

)
|

Illustrating the
Natural History of
the Genus Dero, by
E. C. Bousfield.

DERO OBTUSA.
Dero Perrier, D. latissima, D. Miulleri.
Dero uimosa, D. furcata.

Mata Mrnrst, Doclea tetraptera. )
=) ; I
XANTHO SCABERRIMUS.

PoRCELLANELLA PIcTA, Onychomorpha | Illustrating the

lamelligera, Goniosoma inaquale, Petro- | Oyystacea from

listhes corallicola, var., Diogenes avarus. | Singapore, by
Potyonyx comeTEes, Caphyra Archeri, Alfred O. Walker.

Typhlocarcinus villosus, Sesarma |

Bocourti.

CANDACE NiGROCINCTA, C, brevicornis.
Mecynocera CLAvst.

Illustrating the
Copepoda of Ma-

Macuatropus sancra-crucis, Acontiopho- { deira and the
rus angulatus. Canary Islands, by
CyMBASOMA RIGIDUM, Acontiophorus angu- Isaac C. Thompson.

latus.
Ovicetts of Bryozoa.
Ovicattis of Lichenopore.
Lire-Hisrories of Glyciphagus domesticus and G. spintpes.

Srupres of the Macrocurrss, illustrating Dr. R. W. Shufeldt’s
paper.
New Sprecizs of Sue ts, illustrating G. B. Sowerby’s paper.

Myocorres tenax, Symbiotes tripilis, Goniomerus musculinus,
illustrating some unrecorded Parasitic Acari, by A. D. Michael.

Ovarizs of the Blowfly and their appendages, illustrating B. T.
Lowne’s paper.

BarnyerapuicaL Cuart of the Clyde Sea-area, illustrating W.
E. Hoyle’s paper on the Deep-water Fauna of the Clyde
Sea-area.

Motuvsca of Fernando Noronha.

ERRATA.

Rage 446, line 19 from en, for Centronotus gunellus read Centronotus gun-
nellus,

a 474, line 12 from bottom, fr Pompilus nesophila read Pompilus nesophilus,
ae. 503, line 9 from bottom, for Chiton (Ischnochiton) carribeorum read Chiton
7 (Ischnochiton) caribheorum.

me line 7 from top, for A. eae read A. jaspidea,

e
|
3

THE JOURNAL

THE LINNEAN SOCIETY.

Descriptions of a new Genus and of some new Species of
Galerucine, also Diagnostic Notes on some of the older
described Species of Auwlacophora. By Joseru 8S. Baty,
F.L.S.

[Read 6th May, 1886. ]

Many of the insects characterized in the following paper were

collected in the Eastern Archipelago by Mr. A. R. Wallace.

Types of all the species mentioned are contained in my col-

lection.

Genus AULACOPHORA.

Many of the species of this genus, more especially those
described by the older authors, are in a state of great confusion,
arising to some extent from the great similarity in general
appearance and coloration of many of them; but principally
from the brief descriptions given by the various authors and from
the total absence of any reference to structural characters,

In the present paper I have attempted to work out some of
these species, pointing out the structural characters by which
they may be separated. I have retained, in most cases, the old
names, although I cannot be sure in every instance that the
species to which they are assigned were those from which the
descriptions were originally drawn.

I have not attempted to work out the whole genus, many
species sufficiently characterized by their respective describers, as

LINN. JOURN.—ZOOLOGY, VOL. XxX. 1

2 MR. J. 8. BALY ON SOME SPECIES OF GALERUCIN A.

well as others which, from want of sufficient material, I have not
had the opportunity of studying, having been omitted.

The Synoptical Table given below will, I trust, be of use to the
future student of the genus.

Table of Species.

MALEs.
I. Antenne filiform,

A. Elytra transversely depressed or excavated below
the basilar space.

A. Apical segment of abdomen trilobate, the middle
lobe oblong or oblong-quadrate, its surface
deeply concave or sulcate longitudinally.

a, Antenne (the base sometimes excepted) black or
fuscous.
a}, Elytra entirely black.
61, Breast black, the abdomen fulvous or

HA VOUS 0 a\sisdearedess sh0xce hen seh oe Cae 1. nigripennis.
6?. Breast and abdomen rufo-fulvous............ 2. melanopus.
a*, Elytra black, with flavous bands or markings.
CIP yeidiim ful VOUS. ..ccescne enue moos seeeeen eee 3. albofasciata.
eu Pywiditim ‘black’ siioe.c.ccnsersc 520000 eee 4. pygidialis.
b, Antenne fulvous or fiavous, rarely stained
towards the apex with fuscous, basal joints
sometimes fuscous or black.
d', Elytra entirely black.
e) Body broadly ‘ovabe:.....' J.2.3<t ce ee eee 5. melanoptera.
é*. Body oblong-Oyate: o...2....0%2<s..neeeeombeoamem 6. atripennis.

d?, Hilytra black, the extreme basal margin rufous. 7. lata.
d?. Hlytra black, their apices to a greater or less
extent TULOUS, cscs. -24. recone eaeenwacinae dagen 8. rosea.
d‘, Hlytra black, with flavous markings............ 9. instabilis.
s, Apical segment of abdomen trilobate, middle lobe
quadrate or subquadrate, its disk plane or only
slightly concave.
a. Thorax rufous or flavous.

j*. Tlytra entirely black’ <.350002..ccteneepe ae see 10. Boisduvaili.
f?. Elytra black with flavous markings.
g'. Body not exceeding 4 lines in length ...... 1l. propinqua.
r aaah 5 lines oe re ae eee eee : 4 rubrozonata.
. Elytra flavous, with black patches or markings. 13. approximata.
f*. Elytra flavous, the posterior half black ...... 14. postica.
y’, Milytra, entirely flayous | ...2:..24.<-.ecese esteem 15. testacea.
by Thorax blacks. )s.5.5040. egal seeks cane eee 16. flaviventris.

B. Elytra not transversely depressed or excavated
below the basilar space.
A. Trilobate apex of abdomen with its middle lobe
deeply concave or longitudinally sulcate.
hi, Elytra flavous, unicolorous.

z', Antenne, at the base excepted, nigro-fuscous.
gh, dese entirely MaVOus os i-.st)dacmsticninsemnte 17. abdominalis.
ya, Hour mind des black ;.....sc0sssssrsansereee 18. Fabricii.

¢. Antenne flavous or fulvous.
k1. Scutellum black or nigro-fuscous.

2, Thoracic sulcation deeply excavated
on the middle disk, obsolete or in-
terrupted towards the lateral margin. 19. nigroscutata.

2, Thoracic suleation entire, more deeply
impressed on the middle disk

k?, Scutellum flavous or fulvous.
m', Hace armed on either side below the

eeveeeeea

antennz witha compressed tubercle. 21.

m, Face unarmed.

n'. Thoracic groove deeply excavated and
abruptly sinuate on the middle disk,
the hinder margin of the groove
ete ee Peat ooen ponds is vessovdsans

n*. Thoracic groove straight or only
moderately sinuate on the middle
disk, its hinder margin not torulose.

o'. Legs entirely flavous
o*, Anterior pair of legs fulvous, the two

MR. J. 8. BALY ON SOME SPECIES OF GALERUCINA. 8

20. Wallacii.

cornuta.

. foveicollis.

23. similis, var.

hinder pairs black or nigro-piceous ... 23. similis.
2, Elytra flavous, with black markings.
p'. Hach elytron with two black spots ......... 24, tetraspilota.
p®. Hach elytron with four black spots ......... 25.. octomaculata.
h’. Elytra nigro-ceruleous, thoracic groove deeply
excavated on the middle disk .................. 26. excavata.

h*, Hlytra nigro-piceous, narrowly edged with

flavous; face with two compressed tubercles. 21.

B. Intermediate lobe of apical segment of abdomen
quadrate or subquadrate, plane or slightly
concave.

g'. Elytra flavous.
u', Legs black or nigro-piceous; the labrum

and the lower portion of the clypeus black. 27.

u*, Legs, labrum, and lower surface of clypeus

cornuta, var.

coffeze.

MM ass oy aye nar ovncsddatdedss+aa% 28. bicolor,
var. F & G.
gq’. Hlytra flavous, with black markings’............ 28. bicolor,
var. D & B.
q’®. Hlytra black, the apex rufo-fulvous ............ 28. bicolor, type.
SB mrtetyermentiroly DIACK .....cccecese.ccescessseensens 29. Nigrivestis.
g°. Elytra black, fulvous at the base ............... 30. Downesi.
II. Antennz with the apical joint alone incrassate.
‘4 vt, Hlytra transversely depressed below the basilar
ND Nahas shia fen detqyheviccsesssecsscascaden. Jl. antennata.
v’. Elytra not depressed below the basilar space. 32. luteicornis.
III. Antennz with three or four intermediate joints
dilated.
w', Elytra flavous, witha submarginal black line. 33. Stevensi.
ete MOEA VIFICI-GANCOUS. 6) .ccedeecesseressscerssveses 34. Duboulayi.
FEMALES.
I. Surface of elytra plane or obsoletely sulcate.
A. Elytra transversely depressed or excavated below
the basilar space.
A. Thorax nigro-fulvous or flavous.
a. Apex of last abdominal segment entire.
@. Body ontively flavous ..-...s.0...sessesosastes 35. unicolor.
a, Breast and abdomen black ; legs and antennzx
flavous *.
6*. Body above flavous ..........csssescscestecces 15. testacea.

* 4. palustris, Perroud, belongs to this section, but differs from A. ‘estacea

| in having the legs and antennz black.

MR. J. 8. BALY ON SOME SPECIES OF GALERUCINA.

62. Posterior half of elytra black............... 14. postica.
a°, Breast, legs, and antennz black; elytra
black with yellowish markings ............ 11. propinqua., var.

a*, Breast and abdomen (the apical segment
sometimes excepted) rufy - fulvous or
fulvous ; elytra more or less black.
c', Elytra entirely black.
ad‘, Antenzie black... foie bee 2. melanopus.
d?. Antenne flavous or fulvous, the basal
joint sometimes piceous or black.

el, Gabrumy black | ifsc. teeh-ck eee 10. Boisduvaili.
é*, Labrum rufo-fulyous ..:.f./¢:h. ees 3l. antennata.
c*, Elytra black, the basal margin more or
less distinctly edged with rufous ......... 7. lata.
e°, Elytra black, their apices to a greater or
less extent rufous: -25..5:2 05 teers commen 8. rosea.
ce*, Elytra black, with flavous bands or mark-
ings.

‘fi. Legs entirely black.
g', Body not exceeding 4 lines in
Teng thy cscs comesans cde 52 004 cee ee ll. propinqua.
g°. Body upwards of 4 lines in length. 12. rubrozonata.
f?. Legs with the thighs and the basal
portion of the tibiz rufo-fulvous or
flavous, the rest black.

fe, “Pyeidnam black. -.......0c.-s0s see 4. pygidialis.
h?, Pygidium rufo-flavous .........+0006 3. albofasciata.
sgt ®, Legs entirely flavous .............2-.0+0+ 9. instabilis.

*, Elytra nigro-piceous, the lateral margin
and the suture from below the base to

the apex.fulyous-i.0.vcscust eae eee 38. semilimbata.
_ ¢8, Elytra flavous, with black markings.
2, Surface of elytra obsoletely sulcate...... 36. Nigrosignata.
¢. Surface of elytra plane ...5:.2---. sane 13. approximata.
b. Apex of anal segment of abdomen emarginate.
Kh. Antennigs bak. s/c. «ne aoweace eee 1, nigripennis.
k*, Antenne and legs flavous,
it, Body broadly ovate .:....001--seesserus 5. melanoptera.
, Body oblong-ovate .....,....s0s0.0+sees 6. atripennis.
B:. ‘Thorax entirely black. occ. anncomnuoseeme eee 16. flaviventris.

B. Elytra not transversely depressed or excavated
below the basilar space.
a. Apex of anal segment of abdomen entire.
m', Body, not exceeding 3 lines in length.
n', Elytra nigro-piceous, the base flavous ... 30. Downesi.
n”, Elytra flavous, with black markings.
o', Elytra each with four black spots...... 25. octomaculata.
o”, Elytra each with two black spots’...... 24, tetraspilota.
m. Body 3 lines in length or upwards.
p'. Budy subelongate or oblong, slightly
dilated posteriorly.
g'. Elytra flavous.
wu, Lower portion of clypeus and the
labrum black ’...,..:..isseeesssereers> 22s 27. coffeze.
u*, Lower portion of clypeus and the
inbrum flavous.- ......esseeeee et: cncsaet 28. bicolor, var.
qg°. Elytra variable in tint and markings,
HOt metallic. ..... sss ueeaes end oaiae 28. bicolor.
q*. Elytra metallic green ............sessccees 34. Duboulayi.

MR. J. 8S. BALY ON SOME SPECIES OF GALERUCINA. 5

p?. Body ovate or oblong - ovate, dilated

posteriorly.
v'. Elytra flavous, the suture and a sub-
Pierevonl vitta black ....:........00+0 33. Stevensi.
vy’, Elytra and abdomen black............... 37. Lewisii.

b. Apex of anal segment of abdomen rounded, its
middle portion produced into an oblong -
nl 19. nigroscutata.
e. Apex of anal segment of abdomen slightly
sinuate, its surface deeply concave on either

ee eee 22. foveicollis.
d. Apex of anal segment of abdomen broadly
subangulate-emarginate ..........esceesseeeeees 18. Fabricii.
e, Apex of anal segment of abdomen abruptly ;
CONCAVE-CMATLINALE ....00....00..esceseaceceenees . Wallacii.
f. Apex of anal segment of abdomen broadly
COMGAVE-CMALPINALG ...........2.sesccccccscecneee 21. cornuta.
g. Apex of anal segment of abdomen bilobate.
NTIS CONIA. 50.200. --ascnsnpsasnnccecsccsseces 23. similis.
w*. Lobes plane.
BSE ERYQUG) «0 5.ncsecnde. ocosncaeccesecscones 40. excisa.
Se AMIV EOS, MIGLO-CRFULCOUS ........-+00. 0000005 26. excavata.

h. Apex of anal segment of abdomen bisinuate ... 39. Duvivieri.
t, Apex of anal segment of a3domen trilobate.

y'. Lobes nearly equal in length ................ 41. Mouwhoti.
y*. Lobes with the intermediate much shorter
Pai tno laberal ONES .......3......cseseseoseeee 32. luteicornis.
PS PMPETRGIEOOZIY COSLALG ......2.0..0c0esereresevsscroesccooe 42. costatipennis.

1. AULACOPHORA NIGRIPENNIS, Motsch. Etud. Entom. 1857,
p. 38.

A. atripennis, Hope, Proc. Ent. Soc. 1841, p. 64.

Anguste ovata, postice ampliata, convexa, fulva aut flava, antennis,
pedibus, pectore elytrisque nigris, his infra basin transversim depressis,
distincte punctatis; thorace vix pone medium sat profunde trans-
versim sulcato, sulco fere recto. Long. 3-33 lin.

Mas. Abdominis segmento anali trilobato, lobo intermedio oblongo,
longitudinaliter concavo.

Fem. Abdominis segmento anali apice concavo-emarginato.

Hab. Mantchuria; Japan (Lewis); China (Bowring).

Labrum and antenne black, the latter with the basal joints,

and sometimes with the four or five outer ones, nigro-piceous.
Thorax nearly twice as broad as long; sides sinuate and slightly
diverging from the base to beyond the middle, thence obliquely
converging towards the apex; disk rather strongly sulcate just
behind the middle. Elytra distinctly punctured.

2. AULACOPHORA MELANOPUS, Blanch.Voy. au Péle Sud, p. 346,
pl. 19. fig. 20.

Anguste oblongo-ovata, postice ampliata, convexa, rufo-fulva, nitida,

6 MR. J. 8. BALY ON SOME SPECIES OF GALERUCINEA.

antennis, pedibus totis, pygidio elytrisque nigris; thorace levi, sat
profunde transversim sulcato; elytris infra basin transversim de-
pressis, tenuissime punctatis. Long. 33-65 lin.

Var. A. Femoribus pygidioque rufo-fulvis.

Mas. Abdominis segmento anali trilobato, lobo intermedio transverso-
quadrato, angulis anticis rectangulis, apice truncato, disco longi-
tudinaliter sulcato. .

Fem, Abdominis segmento anali late sed leviter angulato-emarginato.

Hab. New Guinea, Dorey ; Amboina (Wallace).

Antenne slender, filiform, nearly equal to the body in length
in either sex, black, the two to four lower joints usually rufo-
piceous ; labrum and mouth nigro-piceous. Thorax transverse ;
sides nearly straight and parallel from the base to beyond the
middle, thence rounded and converging towards the apex,
anterior angle obtuse; disk smooth, impunctate, discoidal
groove nearly straight, more deeply impressed on either side
the median line. Elytra dilated posteriorly, convex, transversely
excavated below the basilar space, very minutely punctured.

The dark antenne, equally long in both sexes, will separate
this species from others with similarly coloured elytra, It is
with some doubt that I have placed var. A under the same
specific head; it agrees, however, entirely in structural cha-
racters.

3. AULACOPHORA ALBOFASCIATA., Anguste ovata, postice paullo
ampliata, convexa, fulvo-nitida, antennis, ore, tiblis tarsisque nigro-
piceis aut fuscis ; thorace transverso, disco transversim sulcato; elytris
convexis, infra basin prope suturam leviter transversim excavatis,
nigris, utrisque fascia obliqua prope medium communi, ad marginem
abbreviata, albida. Long. 4-5 lin.

Var. A, o. Elytrorum fascia albida fere obsoleta.

Mas. Abdominis segmenti analis lobo intermedio oblongo, profunde

~ concavo.

Fem. Abdominis segmento ultimo late obtuso.

Hab. New Guinea, Dorey (Wallace).

Head scarcely longer than broad, trigonate. Antenne
slender, equal to the body in length in both sexes; the four or
five lower joints sometimes obscure fulvous; eyes in the male
large, prominent. Thorax nearly twice as broad as long; sides
slightly diverging or nearly parallel from the base to beyond the
middle, thence obliquely converging to the anterior angle; disk
transversely sulcate, the sulcation rather more deeply impressed

ey ‘

MR. J. 8. BALY ON SOME SPECIES OF GALERUCINE. 7
on either side. Elytra broader than the thorax, rather broadly
dilated behind the middle, convex, rather strongly excavated
below the basilar space, minutely punctured, the puncturing on
the hinder disk nearly obsolete.

4. AULACOPHORA PYGIDIALIS. Oblongo-ovata, postice ampliata, con-
vexa, flava, nitida; abdominis segmento ultimo plus minusve,
pygidio, pedibus posticis fere totis, tarsis anticis quatuor, tibiis in-
termediis (basi exceptis), labro, antennis (basi exceptis) elytrisque
nigris; his infra basin transversim excavatis, minute punctatis, fascia
lata prope medium, interdum interrupta aut abbreviata, flavo-fulva ;
thorace vix pone medium transversim sulcato. Long. 3-33 lin.

Var. A. Abdominis disco plus minusve nigro-piceo tincto.

Mas. Abdominis segmento anali trilobato, lobo intermedio oblongo,
profunde concavo.

Fem. Abdominis segmento anali apice obtuso, integro.

Hab. Aru Islands; New Guinea, Dorey ; Ceram; Ké Islands
(Wallace).

Antenne slender, filiform, black, more or less piceo-fulvous at
the base. Thorax more than half as broad again as long; sides
obliquely diverging from the base to beyond the middle, sub-
angulate anteriorly ; disk transversely sulcate just behind the
middle. Elytra very finely punctured.

Closely allied to A. albofasciata, smaller and the male broader
than that species, its eyes less prominent; the coloration of the
legs varies greatly in degree. .

5. AULACOPHORA MELANOPTERA, Boisduval, Voy. de l Astrol.

“p. 549.

Late ovata, postice ampliata, convexa, flavo-rufa, nitida ; elytris nigris,
labro piceo; thorace lateribus fere parallelis, obtusis, ad apicem
convergentibus, disco minute punctato, transversim sulcato; elytris
convexis, infra basin prope suturam transversim excavatis, minute
punctatis. Long. 4 ln.

Mas. Abdominis segmento anali trilobato, lobo intermedio oblongo, pro-

funde concavo.
Fem. Abdominis segmento anali apice subangulato-emarginato.

Var. A, ¢. Tarsis piceis.
Galeruca melanoptera, Boisduval, Voy. de l’Astrol. p. 549, t. 8.

fig. 11.
Hab. Celebes (Wallace).
Antenne slender, filiform, nearly three fourths the length of
the body; labrum piceous. Thorax nearly twice as broad as

8 MR. J. S. BALY ON SOME SPECIES OF GALERUCINA.

long; sides nearly parallel, obliquely converging towards the
apex; disk smooth, minutely punctured, transverse groove rather
more deeply impressed on either side. Elytra broadly dilated
posteriorly, convex, transversely excavated below the basilar
space ; minutely punctured, lateral limb reflexed, more strongly
punctured than the disk.

Broader in both sexes than A. atripennis, Fabr., the lateral
margin of the elytra broader and more strongly punctured.

6. AULACOPHORA ATRIPENNIS, Fabr. Syst. El. i. p. 482.

Oblongo-ovata, postice ampliata, convexa, rufo-fulva aut rufa, nitida,
.~ antennis flavis, articulo basali interdum rufo-piceo tincto; thorace

transversim sulcato; elytris nigris, tenuissime punctatis, labro (an
mare) nigro-piceo. Long. 3-4 lin.

Var. A. Pedibus rufo-piceis.

Mas. Abdominis segmento anali trilobato, lobo mtermedio oblongo,

profunde concavo-excavato.

Fem. Abdominis segmento anali apice leviter angulato-emarginato.

Hab. Sumatra; Amboina; Gilolo; Tondano; Ké Islands
(Wallace).

Antenne slender, filiform. Thorax transverse ; sides obliquely
diverging from the base to beyond the middle, thence converging
towards the apex ; disk smooth, impressed on either side with a
few minute punctures; discoidal sulcation nearly straight. Elytra
dilated posteriorly, convex, transversely excavated below the
basilar space, very finely punctured.

This species is smaller in size than either of the following two;
the deeply concave intermediate lobe of the anal segment of the ab-
domen in the male, together with the obtusely emarginate apex of
the same segment in the female, will separate it from either. Many
of the specimens that I have seen differ from the diagnosis given
by Fabricius in not having the basal joint of the antenne darker
than the following ones; in other respects they agree with the
description given. I possess three specimens of the male; in
this sex the labrum is piceous, in the other one it is concolorous
with the; upper face; im some individuals the legs are stained
with p.ceous.

7. AULACOPHORA LATA. Late ovata, postice ampliata, convexa, rufo-
fulva, nitida, tarsis, tibiis plus minusve elytrisque nigris, his basi
extrema plus minusve rufo-marginatis ; antennis pallide flavis ; tho-
race transverso, lateribus obtuse rotundatis, reflexis; disco trans-

MR. J. S. BALY ON SOME SPECIES OF GALERUCINA. 9

versim sulcato, levi, ad latera minute punctato ; elytris late oblongo-
ovatis, postice ampliatis, convexis, infra basin transversim excavatis,
minute punctatis, limbo externo reflexo. Long. 4-55 lin.

Mas. Abdominis segmento anali trilobato, lobo intermedio quadrato,
longitudinaliter suleato.

Fem. Abdominis segmento anali apice obtuso; pygidii apice leviter
emarginato.

Hab. Ceram ; Macassar; Celebes; Gilolo; Batchian (Wallace).

Antenne filiform, nearly equal to the body in length in both
sexes ; Jabrum concolorous with the upper face. Thorax nearly
twice as broad as long; sides obtusely rounded, the hinder angle
obsolete, the anterior one very minute; lateral margin more
broadly reflexed than in most species of the genus ; disk smooth,
very minutely punctured on the sides, transverse sulcation nearly
straight. Elytra broadly oblong-ovate, dilated posteriorly, con-
vex, transversely excavated below the basilar space, minutely
punctured ; outer limb rather broadly dilated and reflexed for
nearly its whole extent.

This species is closely allied to A. rosea, Fabr.: both insects
closely agree in the form of the thorax and in all other struc-
tural characters ; but the present one is distinctly broader in form
and its elytra are (the extreme basal margin excepted) unicolorous.

8. AuLacoPpHoRA ROSEA, Fubr. Syst. El. i. p. 479.

Ovata, postice ampliata, convexa, rufo-fulva aut rufa, nitida, tibiis tar-
sisque nigris, antennis flavis, articulo primo sepe rufo; thorace sat
profunde transversim suleato; lateribus obtuse rotundatis, reflexis ;
elytris infra basin transversim depressis, nigris, apice plus minusve
rufis. Long. 33-6 lin.

Mas. Abdominis segmenti analis lobo intermedio late oblongo, longitu-
dinaliter sulcato.

Fem, Abdominis segmento anali integro.

Hab. Sumatra; Java; Philippine Islands; Malacca.

This species so closely resembles A. lata in the form of its
thorax and in other structural characters, that I do not give a
detailed description; it is separated by its less broadly ovate
form and by the rufous apex of its elytra. The rufous colora-
tion of the apex of the elytra extends upwards to a greater or
smaller extent over the disk. A. albicornis, Chapuis, is apparently
identical with the present insect; at any rate, 1 cannot separate
it on the slight diagnosis given by the author.

10 MR J. 8. BALY ON SOME SPECIES OF GALERUCINA.

9, AULACOPHORA INSTABILIS. Oblongo-ovata, postice ampliata,
flava, nitida, antennis extrorsum fuscis; thorace transversim sulcato,
suleo fere recto; elytris convexis, infra basin leviter transversim de-
pressis, nigris, fascia interrupta lata prope medium, interdum inter-
rupta aut extrorsum abbreviata, flava. Long. 4 lin.

Mas. Abdominis segmento anali trilobato, lobo intermedio oblongo,
concavo, longitudinaliter sulcato.

Fem. Abdominis segmento anali apice obtuso.

Hab. Batchian; Mysol (Wallace).

Antenne slender, filiform, entirely, or with the apices of the
six or seven outer joints alone, fuscous. Thorax about one half
broader than long; sides parallel and slightly sinuate from the
base to beyond the middle, thence slightly converging towards
the apex; disk deeply impressed with a transverse sulcation.
Elytra dilated posteriorly, convex, slightly flattened along the
suture, and faintly excavated behind the basilar space ; nitidous,
very minutely punctured. }

10, AuLAcopHORA BoIsDUVALI. Ovata, postice amplhiata, convexa,
rufo-fulva aut fulva, labro nigro-piceo, nitido, antennis flavis ; tibiis
plus minusve tarsisque piceis, elytris nigris ; thorace profunde trans-
versim suleato. Long. 4-5 lin.

Var. A. Labro rufo-fulvo.

Mas. Abdominis segmento anali trilobato, lobo intermedio transverso-
subquadrato, plano.

Fem. Abdominis segmento anali apice obtuso.

Hab. New Guinea (Boisduval); Borneo, Sarawak; Celebes;
Bouru (Wallace).

Antenne slender and filiform in either sex; nearly equal to
the length of the body in the male, shorter in the other sex;
labrum nigro-piceous, rarely concolorous with the upper face.
Thorax with its sides sinuate and slightly diverging from the
base to beyond the middle, thence obliquely rounded and con-
verging towards the apex; upper surface smooth, finely but not
closely punctured; discoidal sulcation nearly straight, deeply
impressed, more deeply so on the sides. Hlytra much broader
than the thorax, their sides nearly parallel anteriorly, abruptly
dilated posteriorly, convex, slightly excavated transversely below
the basilar space, minutely punctured.

Nearly allied in coloration to A. lata, differing in the form
of the thorax, the sides of which in the present species are

MR. J. S. BALY ON SOME SPECIES OF GALERUCINZ. ll

obtusely angulate and not rounded as in the before-named
insect; there is also a distinct difference, both in the anal
segment of the abdomen in the male and in the male genital
organ.

11. AuLAcopHORA prRopinavaA. Anguste oblongo-ovata, postice
ampliata, convexa, flavo-fulva, nitida, labro, antennis (his basi inter-
dum exceptis), pedibus elytrisque nigris; his infra basin transversim
excavatis, tenuissime punctatis, utrisque fascia lata prope medium,
plerumque ad marginem abbreviata, flavo-fulva. Long. 33-4 lin.

Var. A. Pectore nigro, ceteris ut in typo.

Var. B. Pectore, abdomine pygidioque nigris, ceeteris ut in typo.

Mas. Abdominis segmento anali trilobato, lobo intermedio fiavo, obtuso.

Fem. Abdominis segmento anali obtuso.

Hab. New Guinea, Dorey; Ké Islands; var. A, Batchian ;
var. B, Sulu Islands (Wallace).

Antenne slender, filiform. Thorax about one half as broad
again as long; sides nearly straight and slightly diverging from
the base to beyond the middle ; upper surface transversely sul-
cate behind the middle. Elytra dilated posteriorly, distinctly
excavated on the suture below the basilar space, very minutely
punctured.

Var. B is at first sight very similar to A. dorsalis, Boisduvyal ;
but that species is smaller, and its elytra are more strongly punc-
tured and not depressed below the base.

12. AuLacopHoRA RUBROZONATA, Blanch. Voy. au Péle Sud,
p. 845, pl. 19. fig. 19.

Ovata, postice ampliata, convexa, rufa aut fulva, nitida, antennis (his
basi plerumque exceptis) pedibusque nigris; thorace sat profunde
transversim sulcato; elytris infra basin leviter transversim depressis,
fere impunctatis, nigris, fascia lata mediali rufa aut fulva. Long. 4-
5 ln.

Var. A. Elytrorum margine apicali rufo aut fulvo, ceteris ut in
typo.

Var. B. Elytris fulvis, utrisque basi lata, maculaque subapicali,
nigra,

Var. C. Elytris macula subapicali nigra obsoleta.

Mas. Abdominis segmento anali trilobato, lobo intermedio quam late-
ralia paullo longiore, plano, apice obtuse rotundato.

Fem. Abdominis segmento anali apice obtuso.

Hab. New Guinea, Dorey; Celebes (Wallace).

12 MR. J. 8S. BALY ON SOME SPECIES OF GALERUCINE.

Head longer than broad, trigonate ; mouth nigro-piceous or
piceous. Antenne slender, rather shorter than the body.
Thorax nearly twice as broad as long ; sides slightly sinuate and
very slightly diverging from the base to beyond the middle; disk
deeply impressed with a transverse sulcation. Elytra nitidous,
nearly impunctate.

13, AULACOPHORA APPROXIMATA. Anguste oblongo-ovata, postice
ampliata, convexa, fulvo-flava, nitida, tibiis (basi exceptis) tarsisque
nigro-fuscis ; thorace transversim sulcato, sulci medio leviter sinuato ;
elytris obsolete sulcatis, levibus, minute punctatis, utrisque fascia
basali plagaque pone medium, nigris. Long. 33-9 lin.

Mas. Abdominis segmento anali trilobato, lobo intermedio quadrato,
leviter concavo.

Fem. Abdominis segmento anali apice obtuso.

Hab. Celebes (Wallace).

Antenne slender, filiform, nearly equal to the body in length,
their outer joints more or less stained with fuscous. Thorax
rather more than one half broader than long; sides nearly
straight and parallel from the base to beyond the middle; disk
smooth, deeply impressed transversely, the suleation faintly
sinuate in the median line. LElytra oblong-ovate, dilated poste-
riorly, very finely punctured; a common transverse band at the
base, and a subrotundate patch halfway between the middle and
apex of each elytron, black.

The pale underside, together with the absence of any longitu-
dinal sulcations on the elytra, will separate the female of this
species from that of A. nigrosignata (sp. 36); the male of the
latter insect is unknown to me.

14, AuLAcoPHora PosTiIca, Chapuis, Ann. Soc. Ent. Belg. xix.
1876, p. xcix.

Elongato-ovata, postice ampliata, sordide fulva, nitida, pedibus posteri-
oribus quatuor, pectore, abdomine elytrorumque dimidio postico
nigris ; thorace sat profunde transversim sulcato; elytris infra basin
leviter transversim depressis, distincte punctatis. Long. 44-5 lin.

Mas. Abdominis segmenti analis lobo intermedio oblongo, profunde lon-
gitudinaliter sulcato.

Fem. Abdominis segmento anali apice leviter sinuato.

Hab. Philippine Islands; Java; Malacca.
Thorax nearly twice as broad as long; sides nearly straight
and slightly diverging from the base to beyond the middle ; disk

Se ee eo

MR. J. 8. BALY ON SOME SPECIES OF GALERUCIN A. 13

transversely sulcate, the sulcation nearly straight. Elytra obso-
letely depressed below the base, very minutely punctured.

15. AutacopHora TesTacra, Fabr. Mant. Ins. i. 1787, p. 87.

Oblonga, postice paullo ampliata, convexa, flava, nitida; labro, pectore
abdomineque nigris; thorace vix pone medium transversim sulcato ;
elytris infra basin leviter depressis, tenuissime punctatis. Long. 3 lin.

Mas. Abdominis segmenti analis lobo intermedio transverso-quadrato,

plano.

‘em. Abdominis segmento anali rotundato, integro.

Hab. India (Fabr.); Assam (Chennell: coll. Baly).

Labrum shining black; antenne entirely flavous. Thorax
nearly twice as broad as long; sides nearly straight and slightly
diverging from the base to beyond the middle; disk transversely
sulcate immediately behind the middle, the sulcation straight.
Elytra slightly but distinctly depressed below the basilar space,
very minutely punctured. Hinder femora in the male slightly
stained on their outer surface with piceous.

The entirely black abdomen, together with the entirely dif-
ferent structure of the anal segment of the abdomen, will at
once separate this species from A. foveicollis and A. abdominalis,
with which insects it is usually confounded; it also differs from
both in the depression below the basilar space of the elytra.

16. AULACOPHORA FLAVIVENTRIS, Late ovata, postice ampliata,
convexa, nigra, nitida, pectore piceo, abdomine antennisque (articulo
basali excepto) pallide flavis ; thorace levi, ad latera minute punctato,
sulco transverso, fere recto, sat fortiter impresso; elytris convexis,
infra basin transversim excavatis, minute punctatis. Long. 4 lin.

Mas. Abdominis segmento anali trilobato, lobo intermedio transverso-
quadrato, plano, angulis anticis obtusis.

Fem. Abdominis segmento anali obtuso, leviter sinuato.

Hab. Malay Peninsula, Penang, Tringano, Malacca.

Antenne from the second to the tenth joint slender, filiform
(the terminal joint broken off). Thorax nearly twice as broad
as long; sides nearly straight and slightly diverging from the
base to beyond the middle, thence obliquely converging towards
the apex ; uppersurface smooth, very finely punctured on the sides;
discoidal sulcation nearly straight, deeply impressed. Elytra much
broader than the thorax, dilated posteriorly ; convex, transversely
excavated below the basilar space, very minutely punctured.

Similar in form to A. lata, differing in the colour of the thorax
and lower surface of the body, which is uniform in the three spe-

14 MR. J. 8. BALY ON SOME SPECIES OF GALERUCINE.

cimens (two males and one female) before me; the sides of the
thorax are also straight posteriorly, instead of rounded as in the
former species.

17. AULACOPHORA ABDOMINALIS, Fabr. Spec. Ins. i. p. 151.

Oblonga, postice paullo ampliata, convexa, flava, nitida, antennis ex-
trorsum, pectore abdomineque (ano excepto ) nigris; thorace transversim
suleato, sulci medio magis profunde impresso. Long. 3 lin.

Mas. Abdominis segmenti analis lobo intermedio oblongo, profunde
longitudinaliter concavo.

Hab. Islands of the Pacitic Ocean (Fabr.) ; Western Australia
(my collection). \

Very similar in general appearance to A. foveicollis, Kiist., the
thorax less deeply excavated on the middle disk, the outer half of
the antenne black. I only know the male of this species.

18. AuLAcopHoRA Fasriciit. Anguste subelongato-ovata, postice
paullo ampliata, convexa, flava, nitida, antennis (basi exceptis), pedibus
posticis quatuor, pectore abdomineque (hujus apice excepto) nigris ;
thorace transversim sulcato, sulco in mare disci medio magis pro-
funde excavato ; elytris tenuiter punctatis. Long. 3+33 lin.

Mas. Antennarum articulo basali incrassato, abdominis segmento anali
trilobato, lobo intermedio alteris longiore, sat profunde concavo-
excavato.

fem. Abdominis segmento anali apice sinuato.

Hab. Tonga Islands.

Antenne filiform, nearly three fourths the length of the body,
the basal joint in the male strongly thickened. Thorax nearly
twice as broad as long; sides nearly straight and slightly diverg-
ing from the base to beyond the middle, thence rounded and con-
verging towards the apex; disk transversely excavated, the
sulcation in the male more broadly and deeply excavated on the
middle disk. Elytra narrowly oblong, slightly dilated posteriorly,
very finely punctured.

Separated from A. nigroscutata by the flavous scutellum and by
the thoracic sulcation extending entirely across the disk; the
apical abdominal segment in the female is also differently formed.

19. AULACOPHORA NIGROSCUTATA. Anguste oblonga, postice paullo
ampliata, convexa, flava, nitida, pectore, abdomine scutelloque nigris ;
thorace transverso, disci medio transversim excavato, suleo utrinque
abbreviato; elytris tenuiter punctatis. Long. 3 lin.

Mas. Thoracis sulco profunde impresso ; abdominis segmento anali tri-
lobato, lobo intermedio profunde excavato.

MR. J.'8. BALY ON SOME SPECIES OF GALERUCINA. 15

Fem. Abdominis segmento anali apice rotundato, medio processu ob-

longo retrorsum producto armato ; pygidio apice emarginato.

Hab. Gilolo; Amboyna (Wallace).

Antenne filiform. Thorax nearly twice as broad as long; sides
parallel from the base to beyond the middle; disk convex, deeply
impressed with a large transverse excavation, which terminates on
either side some distance before reaching the lateral margin ; this
excavation is less deeply impressed in the female; in this sex on
either side, between the excavation and the lateral border, is a
small fovea. LElytra oblong, dilated posteriorly, convex, finely
and rather closely punctured.

20. AULAcoPHORA Wa.uacil. Subelongato-ovata, postice ampliata,
convexa, flava, nitida, pectore abdomineque nigris, scutello femoribus-
que posticis quatuor nigro-piceis ; thorace transversim sulcato; ely-
tris subnitidis, minute punctatis. Long. 4 lin.

Mas, Thoracis suleco medio profunde excavato; antennis filiformibus,
articulo primo sat valde incrassato; abdominis segmento anali tri-
lobato, lobo intermedio oblongo, profunde excavato.

Fem. Abdominis segmento avali apice emarginato, incisura transverso-
quadrata, basi angulato-producta.

Hab. Timor (Wallace).

Antenne filiform, about four fifths the length of the body,
basal joint strongly thickened in the male; anterior half of
labrum nigro-piceous. Thorax nearly twice as broad as long;
sides more or less distinctly sinuate and slightly diverging from
the base to beyond the middle, thence obliquely rounded towards
the apex ; upper surface finely but distinctly punctured on the
sides, impressed across the middle with a nearly straight trans-
verse sulcation, which is of nearly equal depth for its whole course
in the female, but which is much broader and more deeply exca-
vated on the middle disk in the male. Elytra subnitidous, not
distinctly excavated below the basilar space, minutely punctured.

This insect in coloration closely resembles A. similis, but

the four posterior thighs alone are nigro-piceous, the middle disk
of the thorax is more deeply excavated in the male, and the
emargination at the apex of the abdomen in the female is of an
entirely different form. I only know three specimens, two males
and one female, all from Timor.

21. AunLAcoPHoRA cornuTaA, Baly, Cist. Ent. ii. p. 445.
Oblonga, postice paullo ampliata, convexa, flava, labro, peetore abdomi-

16 MR. J. S. BALY ON SOME SPECIES OF GALERUCINE.

neque nigris; thorace transversim sulcato; elytris tenuiter punctatis.
Long. 33-4 lin.

Var. A. Elytris nigro-piceis, anguste flavo-limbatis.

A. robusta, Duviv. Notes Leyd. Mus. vi. p. 124.

Mas. Antennarum articulo basali incrassato, intus compresso, carinato ;
fascie utrinque infra antennas tuberculo compresso, subconico, medio-
que lamina brevi, pube nigro circumdata, instructa ; telo apice hastato ;
abdominis segmento anali trilobato, lobo intermedio profunde longi-
tudinaliter sulcato.

Fem. Abdominis segmento anali late concavo-emarginato, emargina-
tionis medio dente brevi instructo.

Hab. Assam ; Siam (Mouhot) ; Celebes; Ceram; Timor ; Sulu
Islands ; Flores (Wallace); var. A, Wagaiou (Wallace); New
Guinea (v. Rosenberg).

Antenne filiform. Thorax twice as broad as long; sides
rounded, nearly straight at the base; upper surface deeply
grooved just behind the middle, the sulcation nearly straight.
Elytra oblong, slightly dilated posteriorly, convex, not depressed
below the basilar space, finely punctured.

Var. A at first sight is very dissimilar to the type, and has
been described by M. Duvivier as a separate species ; it agrees,
however, so closely in structural characters that I cannot but
place it under the same specific head.

22. AULACOPHORA FOVEICOLLIS, Kister, Kaf. Eur. xxviii. p. 100.

Oblonga, postice paullo ampliata, convexa, flava, nitida, pectore abdomi-
neque (ano excepto) nigris ; thorace transversim sulcato, suleco medio
angulato, magis fortiter excavato; elytris tenuissime punctatis.
Long. 3-33 lin.

Mas. Antennarum articulo basaliincrassato; thoracis sulcomedio ampliato
et profunde excavato et margine postico utrinque tuberculo conico in-
structo; abdominis segmento anali trilobato, lobo intermedio oblongo,
profunde longitudinaliter concavo.

Fem. Abdominis segmento anali apice breviter sinuato ; disco utrinque
concavo-excavato, pygidio apice emarginato.

Hab. Southern Europe; Northern Africa ; India.

The peculiar sculpture of the thorax, combiee with the form
of the anal segment of the abdomen in either sex, will distinguish
this species at once from its congeners.

23. AULACOPHORA SIMILIS, Oliv. Ent. vi. p. 624, pl. 2. f. 28.

A. femoralis, Motsch. Etud. Ent. 1857, p. 37.

Elongato-ovata, postice ampliata, flava, nitida, pectore abdomineque
(ano flavo plerumque excepto) nigris; pedibus posticis quatuor plus

MR. J. S. BALY ON SOME SPECIES OF GALERUCINA. 17

minusve nigris; thorace transversim sulcato, sulco medio sinuato ;
elytris tenuiter sat crebre punctatis. Long. 3-33 lin.
Var. A. Pedibus flavis.
Rhaphidopalpa flavipes, Jac. Notes Leyden Mus. v. p. 202.
Mas. Thoracis suleco transverso medio profunde excavato; abdominis
segmento anali trilobato, lobo intermedio oblongo, profunde longitu-
dinaliter excavato.
Fem. Abdominis segmento anali bilobato, lobis concavis.
Hab. Mantchuria; Japan; Northern India, China; Canton;
Cochin China; Malay Archipelago. |

The male of the present species is very similar in form and in
the structure of the anal abdominal segment to the same sex
of A. foveicollis and of A. abdominalis; the four hinder legs are
usually stained more or less with nigro-piceous, although in some
specimens from Canton they are entirely flavous; it may be
known, however, by the thorax being less deeply excavated. The
female is at once distinguished by the deeply bilobate anal segment
of the abdomen.

This species is very widely spread, and is often confounded in

collections with A. coffee, Hornst.

24. AULACOPHORA TETRASPILOTA. Elongato-ovata, postice ampliata,
pallide flava, nitida; pectore abdomineque nigris; thorace lateribus
rotundatis, basi fere rectis, disco transversim suleato; elytris tenuiter
punctatis, utrisque puncto subbasali maculaque transversa pone
medium, nigris. Long. 23 lin.

Mas. Abdominis segmento anali trilobato, lobo intermedio oblongo,
profunde concavo.

Fem. Abdominis segmento anali apice obtuse truncato.

Hab. Batchian ; Amboina; Gilolo (Wallace).

Antenne filiform, equal to the body in length in the male.
Thorax about one half broader than long; sides rounded, nearly
straight from the base to beyond the middle; disk rather deeply
transversely sulcate. Elytra finely but distinctly punctured;
each elytron with a small round spot immediately below the
middle of the basal margin, together with a second, transverse,
placed obliquely a short distance below the middle, black.

25. AULACOPHORA OCTOMACULATA. Late ovata, postice ampliata, con-
vexa, pallide flava, nitida; pectore abdominisque segmentorum maculis
(his triseriatim dispositis) nigris; thorace lateribus a basi fere ad apicem
subrectis, angulis acutis, disco utrinque transversim excavato; elytris
tenuiter sed distincte punctatis ; utrisque maculis quatuor (2, 2 trans-
versim dispositis) nigris. Long. 3 lin,

LINN. JOURN.— ZOOLOGY, VOL. XX. 2

, MR. J. 8. BALY ON SOME SPECIES OF GALERUCINA.

. Mas. Abdominis segmento anali trilobato, lobo intermedio oblongo-
quadrato, concavo.

Fem. Abdowminis segmento anali apice obtuso.

Hab. Northern India.

Eyes black, vertex with two small piceous spots, sometimes
obsolete; antenne filiform. Thorax about one half broader than
long; sides slightly diverging, and almost straight from the base
nearly to the apex; the anterior and posterior angles acute ;
upper surface broadly and deeply excavated transversely on either
side, the excavation extending nearly across the entire disk.
Elytra subquadrate-ovate, dilated posteriorly ; moderately convex,
distinctly but not very closely punctured ; each elytron with four
large black patches, placed obliquely in pairs, two at the base and
two below the middle; of the upper pair the first, near the
suture, is attached to the basal margin, the other is placed just
below the humeral callus; the outer one of the lower pair is
attached to the lateral margin.

26. AULACOPHORA EXCAVATA. Oblongo-ovata, postice paullo am-
pliata, flava, nitida; thoracis disco sat profunde excavato ; elytris
subnitidis, tenuiter subcrebre punctatis, nigro-cyaneis; tibiis epee
tarsisque infuscatis. Long. 43-5 lin.

Mas. Abdominis segmento anali profunde trilobato, lobo intermedio
alteris paullo longiore, oblongo, apice truncato, concavo.

Fem. Abdominis segmento anali bilobato, lobis apice incurvatis, pygidio
apice angulato-emarginato.

| Hab. India.

Antenne filiform. Thorax about one half broader than long;
sides nearly parallel and slightly sinuate from the base nearly to
the apex ; middle disk deeply excavated transversely, the hinder
margin of the excavation obsoletely trisinuate. Elytra finely
punctured.

Nearly allied to A. impressa, Fabr., its thorax less deeply ex-
cavated, the excavation different in fina. the elytra unicolorous,
and the under surface of the body entirely flavous.

27. AULACOPHORA COFFE®, Hornst. Schrift. Berl. Ges. viii.
1788; pb. tia. a: 7.

Aulacophora varians, var. B?, Chapuis, Ann. Soc. Ent. Belg. t. xix. p. ¢.

Subelongata, postice paullo ampliata, convexa, sordide flava, nitida ;
oculis magnis; pectore, abdomine pedibusque nigris, vertice plus
minusve, clypei margine antice labroque nigro-piceis; scutello piceo ;
thorace levi, sat profunde transversim sulcato ; elytris minute punc-
tatis, subnitidis. Long. 33-4 lin.

MR. J. 8S. BALY ON SOME SPECIES OF GALERUCINZ. 19

Var. A. Pedibus anticis flavis.

Mas. Abdominis segmento anali trilobato, lobo intermedio quadrato,

plano.

Fem. Abdominis segmento anali integro, obtuso.

Hab. Java ; Sumatra; Philippine Islands ; Tondano ; ; Tringano;
Cambodia; India.

jee filiform, equal to the body in length in the male,
rather shorter in the female ; anterior margin of clypeus, together
with the entire labrum, nigro-piceous; vertex also stained
more or less with the same colour. Thorax about one. half
broader than long; sides very slightly diverging and obsoletely
sinuate from the base to beyond the middle, thence converging
and slightly rounded towards the apex; disk smooth, deeply im-
pressed transversely immediately behind the middle, the sulcation
nearly straight. Elytra convex, not excavated below the basilar
space, minutely punctured.

This species closely resembles A. bicolor in general form and in
the structure of the anal segment of the abdomen in both sexes ;
in addition, however, to the uniform coloration of the elytra, the
nigro-piceous labrum and anterior margin of the clypeus will
separate it from all the varieties of that species.

28. AULACOPHORA BICOLOR, Weber, Obs. Ent. 1801, p. 56;
Fabr. Syst. El. i. p. 482.

Subelongata, postice paullo ampliata, convexa, nigra, nitida; capite,
thorace scutelloque flavis aut fulvis; pedibus anticis femoribusque
intermediis sordide flavis, plus minusve piceo tinctis; elytris opacis,
nigris, macula parva basali apiceque fulvis. Long. 4-5 lin.

Var. A. Corpore subtus toto fulvo.

Var. B. Elytris nigris, apice marginibus laterali et suturali et interdum
fascia interrupta prope medium fulvis.

Var. C. Elytris rufo-fulvis, dimidio apicali et utrisque maculis duabus
infra basin, nigris; pedibus anticis quatuor fere totis sordide fulvis.
Var. D. Elytris fulvis, utrisque maculis transversis irregularibus duabus,

una ad basin, altera vix pone medium positis, nigris; pedibus ut in typo.

Var. E. Elytris flavis aut fulvis, utrisque maculis duabus, infra basin
transversim positis, fasciaque transversa vix pone medium, utrinque
abbreviata, nigris.

Aulacophora sexnotata, Chapuis, Ann. Soc. Ent. Belg. t. xix. p. e.

Var. F. Corpore flavo, oculis nigris.

Var. G. Minor; corpore flavo, elytris nitidis.

Mas. Abdominis segmento anali trilobato, lobo intermedio quadrato,
leviter concavo.

Fem. Abdominis segmento anali integro, obtuso.

20 MR. J. S. BALY ON SOME SPECIES OF GALERUCINEA.

Hab. Type, Sumatra, Java; var. A, Celebes, Gilolo, Flores ;
var. B, C, and D, Java; var. E, Java, Philippines, Lombock ;
var. F, Ceylon; var. G, Celebes.

The above species is very nearly allied to A. coffee, Hornst.,
agreeing closely in form and structural characters. The head in
the male of the present insect is narrow and the eyes are larger and
more prominent, and the apical surface of the intermediate lobe
of the anal segment of the abdomen is slightly concave instead
of plane. A. bicolor, however, although extremely variable in
coloration, always has the lower edge of the clypeus and the
entire labrum concolorous with the upper face; in A. coffee, on
the contrary, these parts are nigro-piceous or black.

29. AULACOPHORA NIGRIVESTIS, Boisd. Voy. del Astrol. p. 548.

Subelongata, postice paullo ampliata, convexa, fulva, nitida; tibiis apice,
tarsis antennarumque articulis intermediis piceis ; abdomine elytrisque
nigris, his zneo tinctis; thorace lateribus a basi fere ad apicem
rectis, disco sat fortiter transversim sulcato, suleo utrinque magis
distincte impresso. Long. 3} lin.

Mas. Abdominis segmento anali trilobato, lobo mtermedio alteris lon-

giore, transverso-quadrato, plano, apice truncato.

Hab. Fiji Islands, a single specimen.

Antenne nearly equal to the body in length, slender, filiform.
Thorax nearly twice as broad as long; sides straight and parallel,
obliquely converging near the apex. LElytra narrowly oblong,
slightly dilated posteriorly, convex, distinctly punctured. Ex-
treme apex of the abdomen piceous.

30. AULACOPHORA DownEsI. Ovata, postice ampliata, nigra, nitida,
capite, thorace, scutello pedibusque flavis; thorace lateribus ante
medium obtuse angulatis, disco modice transversim sulcato; elytris
tenuiter punctatis, plaga magna basali, trigonata, communi, flava.
Long. 23-3 lin.

Mas. Abdominis segmento anali trilobato, lobo intermedio late oblongo-
quadrato, plano.

Fem. Abdominis segmento anali apice sinuato.

Hab. India, Bombay (Dr. Hzra Downes).

Antenne filiform. Thorax about one half broader than long;
sides obtusely angulate before the middle, nearly straight pos-
teriorly ; upper surface transversely sulcate, the sulcation less
deeply impressed than in most other species of the genus.
Elytra dilated posteriorly, moderately convex, very finely punc-
tured.

MR. J. 8S. BALY ON SOME SPECIES OF GALERUCINA. 21

In the two specimens before me the flavous patch on the elytra
: varies in extent.

31. AULACOPHORA ANTENNATA. Anguste ovata, postice ampliata,
convexa, rufo-fulva, nitida ; antennis flavis, pectore rarius pedibusque
piceis; elytris convexis, infra basin prope suturam transversim excavatis,
nigris, tenuissime punctatis; thorace levi, sat profunde transversim
suleato. Long. 4-45 lin.

Mas. Antennarum articulo ultimo incrassato, apice oblique sinuato-
emarginato, dente acuto armato; abdominis articulo anali trilobato,
lobo intermedio transverso-quadrato, disco concavo.

Fem. Antennarum articulo ultimo non incrassato, abdominis segmento
anali late truncato, obsolete sinuato.

Var. A. Pedibus fulvis.

Hab. Java; Singapore; Borneo, Sarawak ( Wallace).

Antenne filiform, the apical joint in the male thickened, its

apex obliquely emarginate and armed with a short acute tooth;
labrum and mouth concolorous with the upper face. Thorax
nearly twice as broad as long; sides nearly straight and very
slightly diverging from the base to just beyond the middle, thence
obliquely converging towards the apex, anterior angle obliquely
truncate, armed laterally with a small obtuse tooth; upper
surface smooth, with a few scattered punctures on either side
behind the anterior angle; transverse groove nearly straight,
deeply impressed. Elytra dilated posteriorly, convex, transversely
excavated below the basilar space, minutely but not closely

punctured.

32. AULACOPHORA LUTEICORNIS, Habr.

Late ovata, postice ampliata, thorace sat profunde transversim sulcato,
lateribus obtuse rotundatis ; elytris tenuissime punctatis. Long. 4 lin,

Var. A. Nigra, nitida; thoracis lateribus angustis antennisque pallide
flavis; elytris rufis, apice nigris ; tarsis interdum rufo-piceis.

Var. B. Thoracis lateribus disco concoloribus, ceteris ut in var. A.

Galeruca luteicornis, Fabr. Syst. El. i. p. 482.

Var. C. Elytris totis rufis, caeteris ut in varr. A et B.

Aulacophora simplicipennis, Clark, Ann. §& Mag. Nat. Hist. ser. 3, xv.

1865, p. 145.

Var. D. Nigra, antennis (basi exceptis) pallide flavis.

Var. E. Tota flava.

Mas. Antennarum articulo ultimo incrassato, apice emarginato ; abdo-
minis segmenti analis lobo imtermedio plano, transverso-quadrato

angulis anticis obtusis.

22 MR. J. 8S. BALY ON SOME SPECIES OF GALERUCINA.

Fem. Abdominis segmento anali transversim emarginato, trilobato, lobo

intermedio brevi, truncato, lobis lateralibus subacutis, apice incurvatis.

Hab. Malay Peninsula, Singapore, Penang, Tringano, Malacca ;
Borneo, Sarawak ; Sumatra (Wallace).

This variable insect in point of coloration is, owing to the
dilated apical joint of its antenne in the male and to the peculiar
emargination of the anal segment of the abdomen in the female,
one of the best defined species of the genus; the formation of
the anal segment of the abdomen will at once distinguish the latter
sex from all others known to me of the genus. Aulacophora
antennata, described in the present paper, is the only insect with
which the male can be confounded ; in the male of A. antennata
the body is larger and narrower, the dilated apical joint of its
antenne is differently notched at the apex, and the intermediate
lobe of the anal segment is concave, not plane as in A. luteicornis.
The apical segment of the abdomen in the female of A. anten-
nata is not emarginate.

33. AULACOPHORA STEVENSI. Anguste ovata, convexa, pallide flava,
nitida, scutello nigro-piceo, oculis nigris; thorace transversim sulcato;
elytris tenuissime punctatis, basi extrema, sutura, apice, nec non linea
submarginali, a basi fere ad apicem extensa, nigris. Long. 33-4 lin.

Mas. Antennarum articulis 3°-5™ incrassatis ; abdominis segmento anali
trilobato, lobo intermedio oblongo-quadrato, profunde concavo.

Fem. Antennis filiformibus; abdominis segmento anali apice sinuato.

Hab. Ceylon; India, Dinapore.

Front impressed on either side with an oval fovea in the male.
Antenne in the same sex with the second joint very short, tur-
binate, the following three thickened; the third compressed, tri-
gonate, its antero-external angle very acute; the fourth more
strongly thickened, oblong-ovate; the fifth slightly compressed,
subtrigonate, its outer edge armed near the apex with a short
acute tooth, its upper surface excavated.

34. AULACOPHORA DuBouLaAyti. Anguste oblonga, postice paullo
ampliata, convexa, flava aut fulva, nitida, collo utrinque nigro, tibiis ad
apicem tarsisque nigro-piceis; thorace transversim sulcato; elytris
tenuissime, subcrebre punctatis, viridi-eeneis. Long. 3-33 lin,

Mas. Antennarum articulis tertio ad sextum dilatatis, fronte utrinque
intra oculum foveolata; abdominis segmento anali trilobato, lobo in-
termedio oblongo-quadrato.

Fem. Antennis filiformibus nigris, basi fulvis; abdominis segmento
anali apice leviter sinuato.

Hab. Western Australia (Duboulay).

NE EES a ee eee

MR. J. S. BALY ON SOME SPECIES OF GALERUCIN A. 23

Upper portion of front in the male impressed just within the
eye with a large round fovea ; antenne in the same sex entirely
flavous, the second joint very short, moniliform, the third thickened,
subtrigonate, its antero-external angle armed with a long acute
tooth, the fourth joint strongly thickened, compressed externally,
equal in length to the third, the fifth shorter than either of the
preceding, compressed and dilated externally at the base; an-
tenne in the female simple, filiform. Thorax transverse, trans-
versely sulcate, more deeply excavated on either side the median
line ; its surface smooth, impunctate ; sides nearly straight and
parallel, slightly converging anteriorly. Elytra narrowly oblong,
parallei, very finely punctured.

This species is separated from A. smaragdipennis, Duv. (viridi-
pennis, Chap.), by the dilated joints of the antennz in the male
Sex.

Females of which the males are unknown to me.

35. AULACOPHORA UNICOLOR, Jac. Notes Leyd. Mus. v. p. 201.

Ovata, postice ampliata, convexa, flava, nitida; thorace sat profunde
transversim sulcato ; elytris infra basin transversim depressis, tenuiter
punctatis. Long. 5 lin.

Fem. Abdominis segmento anali integro.

Hab. Saleyer.

This insect, of which Mr. Jacoby has kindly sent mea specimen,
exactly resembles A. Bozsduvalz in size, form, and sculpture, only
differing in the uniform flavous coloration of the whole body ;
whether it is a distinct species or only a local form of A. Boisdu-
vali it is impossible, in the absence of the male (which is at
present unknown), to determine.

36. AULACOPHORA NIGROSIGNATA. Anguste ovata, postice ampliata,
flava, nitida; pectore abdomineque nigris ; thorace transversim sulcato,
sulco medio subinterrupto; elytris subnitidis, tenuiter punctatis,
utrisque plaga basali transversa, utrinque abbreviata, fasciaque pone
medium, interdum ad suturam abbreviata, nigris. Long. 4-43 lin.

Fem. Abdominis segmento anali apice integro, obtuso.

Hab. Flores (Wallace).

Antenne nearly three fourths the length of the body, filiform.
Thorax nearly twice as broad as long; sides nearly straight and
slightly diverging from the base to beyond the middle, thence
converging towards the apex; upper surface finely but distinctly
punctured on either side in front; disk deeply sulcate transversely,

24 MR. J. 8. BALY ON SOME SPECIES OF GALERUCIN A.

the sulcation less deeply impressed in its middle. Elytra sub-
nitidous, minutely punctured; on each elytron are several faint
longitudinal sulcations.

37. AULAcoPHORA LeEwisiI. Anguste ovata, postice ampliata, con-
vexa, rufo-fulva, nitida; abdomine nigro-piceo; thorace vix pone
medium transversim sulcato; elytris nigris, infra basin non excavatis,
minute punctatis. Long. 3 lin.

Fem. Abdominis segmento anali integro.

Hab. China, Hongkong (Lewis).

Thorax nearly twice as broad as long; sides nearly straight
and parallel from the base to beyond the middle, thence converging
towards the apex ; disk smooth, impressed just behind the middle
with a nearly straight transverse sulcation. Elytra not depressed
transversely below the basilar space, minutely punctured.

At first sight very close to A. nigripennis, but at once to be
known by the different coloration of the under surface of the
body, by the pale legs, and by the absence of the subbasilar de-
pression on the elytra. Although from so widely different a
locality, it is just possible that this insect may be the female of
A. nigrivestis, Boisd.

38. AULACOPHORA SEMILIMBATA. Anguste oblongo-ovata, convexa,
flava, nitida; antennis (basi exceptis),tibiis apice tarsisque nigro-piceis ;
abdomine piceo tincto ; thorace profunde transversim sulcato; elytris
convexis, infra basin excavatis, nigris, nitidis, limbo suturali, nee non
margine externo, his ante medium obsoletis, flavis. Long. 3 lin.

Fem. Abdominis segmento anali obtuso, obsolete sinuato.

Hab. New Guinea, Dorey ( Wallace).

Antenne slender, filiform, the basal joint flavous. Thorax
nearly twice as broad as long; sides distinctly angulate; disk
transversely impressed with a deep sulcation, which is rather
more strongly excavated on either side the median line. LElytra
oblong, slightly dilated posteriorly ; convex, distinctly excavated
on the suture below the basilar space, shining, impunctate.

39. AULACOPHORA DvuvivieRI. Anguste ovata, postice paullo am-
pliata, convexa, flava, nitida; pectore, abdomine pedibusque poste-
rioribus nigris, anticis piceis ; thorace quam longo fere duplo latiore,
transversim sulcato, sulco prope marginum fere obsoleto; elytris
minute punctatis. Long. 4 lin.

Var. A. Tibiis tarsisque intermediis piceis.

Fem. Abdominis segmento anali apice late emarginato, leviter bisinuato,

Hab. Java; Malacca.

MR. J. 8S. BALY ON SOME SPECIES OF GALERUCINEA. 25

Antenne filiform; labrum concolorous with the upper face.
Thorax nearly twice as broad as long; sides nearly straight and
parallel from the base to beyond the middle, thence obliquely
converging towards the apex; disk transversely excavated across
the middle, the sulcation nearly obsolete near the lateral margin.
Elytra narrowly oblong, slightly dilated posteriorly, convex, not
excavated below the basilar space, minutely punctured.

40. AULACOPHORA EXCISA. Ovata, postice ampliata, convexa, flava,
nitida; pectore abdomineque (hoc apice excepto) nigris; femoribus pos-
ticis quatuor piceo-nigris ; thorace sat profunde transversim sulcato ;
elytris minute punctatis. Long. 33 lin.

Var. A. Antennis extrorsum pedibusque posticis quatuor nigris, pedibus
anticis piceo tinctis.

Fem. Abdominis segmento anali apice bilobato, lobis planis, ad apicem
incurvatis.

Hab. Celebes; Flores; var. A, New Guinea (Wallace).

- Antenne slender, filiform, nearly two thirds the length of the
body, the outer half slightly stamed with piceous; labrum
piceous. Thorax transverse ; sides nearly straight and parallel
from the base to beyond the middle, thence converging towards
the apex, the posterior angle obtuse; disk deeply impressed across
the middle with a transverse groove. LElytra minutely, but not
closely punctured. Apical lobes of abdomen plane, separated by
a broad deep incision.

The peculiar form of the apical segment of the abdomen in the

female (the only sex known to me) will at once separate this
species from its allies.

41. AutacopHoRA MovunorTi. Anguste ovata, postice ampliata,
convexa, flava, nitida ; pectore abdomineque nigris, scutello nigro-
piceo ; thorace sat profunde transversim sulcato ; elytris minute sub-
crebre punctatis. Long. 3 lin.

Fem. Abdominis segmento anali trilobato, lobis zquilongis, intermedio
apice emarginato.

Hab. Cochin China; Malacea.

Antenne slender, filiform, more than half the length of the
body, stained with piceous. Thorax transverse; sides nearly
straight and parallel from the base to just beyond the middle;
disk deeply transversely sulcate. Elytra minutely punctured.

The above insect, of which I know only the female, agrees with
A. luteicornis in the anal segment of its abdomen being trilobate,

26 MR. J. 8S. BALY ON SOME SPECIES OF GALERUCINA.

but differs in the narrower form and in the intermediate anal
lobe being nearly equal in length to the lateral ones.

42. AULACOPHORA COSTATIPENNIS. Ovata, postice ampliata, convexa,
rufo-fulva, nitida; thorace sat profunde transversim suleato, lateribus
angulatis; elytris infra basin leviter transversim depressis, nigris,
elevato-vittatis, interspatiis punctatis. Long. 33 lin.

Fem, Abdominis segmento anali apice obtuso.

Hab. A single specimen, my collection.

Antenne slender, filiform. Thorax nearly twice as broad as
long; sides straight and diverging from the base to beyond the
middle, thence obliquely converging towards the apex; disk mi-
nutely punctured on either side, deeply transversely sulcate.
Elytra broadly ovate, dilated posteriorly, convex, slightly depressed
transversely below the basilar space; each elytron with nine
distinctly elevated longitudinal coste, their interspaces minutely
punctate.

Nearly allied to A. Batesi, Jac., and A. anchora, Redt. ;
elytra nitidous, entirely black, and distbiniate (cpa hielo: ae
basilar space.

Genus PsEuDocoPHORA, Jacoby.

PsEUDOCOPHORA BRUNNEA. Ovata, postice ampliata, valde convexa,

pallide fulvo-picea, nitida, pectore, abdomine pedibusque posterioribus
quatuor nigris; thorace leviter transversim sulcato, lateribus angulatis;
elytris distincte punctatis. Long. 3 lin.

Hab. Celebes (Wallace).

Antenne filiform. Thorax rather more than one half broader
than long; sides distinctly produced and angulate before the
middle; disk impressed transversely with a shallow sulcation.
Elytra broadly oblong-ovate, dilated posteriorly, convex, distinctly
punctured, the puncturing nearly obsolete on the hinder disk.

In the typical species of the genus, as well as in the present
one, the prosternum is distinct and nearly equal in height to the
coxe ; this character is not mentioned by Mr. Jacoby.

Genus PARIDEA.

Corpus ovatum, postice ampliatum. Caput exsertum; oculis rotundatis,
integris ; antennis filiformibus. Thorax transverso-quadratus, dorso
transversim sulcatus. Elytrorum epipleura a basi ad medium extensa.
Pedes simplices; tibiis apice spina brevi armatis ; tarsis posticis articulo

MR. J. 8. BALY ON SOME SPECIES OF GALERUCIN”|. 97

basali ad duos sequentes equilongo ; wnguiculis appendiculatis. Pro-
sternum coxis anticis occultum ; acetabulis anticis apertis.
Type Paridea thoracica.
Closely resembling Awlacophora in general form, separated by
the prolonged elytral epipleura and by the appendiculated claws.

PARIDEA THORACICA. Ovata, postice ampliata, convexa, flava,
nitida ; pectore abdominisque maculis nigris; capite thoraceque rufo-
fulvis; antennis (basi exceptis) nigro-piceis; thorace quam longo latiore,
lateribus fere parallelis, sinuatis, disco profunde transversim excavato ;
elytris tenuiter sed distincte punctatis, utrisque plaga basali subrotun-
data, alteraque pone medium, transversa, nigris. Long. 23 lin.

Fem. Abdominis segmento anali integro, obtuso.

Hab. India.

Antenne rather slender, filiform. Thorax about one fourth
broader than long; sides parallel and rather deeply sinuate from
the base to beyond the middle, very slightly dilated before the
latter ; disk smooth, broadly and deeply excavated transversely,
more deeply excavated on either side. Elytra dilated posteriorly,
finely but distinctly punctured. Abdominal segments stained
with black on either side.

Genus AtnrprA, Baly.

ZENIDEA FACIALIS. Elongata, flavo-fulva, nitida; thorace levi, disco
leviter transversim excavato; lateribus a basi ad ultra medium diver-
gentibus, hine ad apicem rotundato-convergentibus ; elytris parallelis,
puncto humerali suturaque basi, nigris. Long. 4 lin.

Mas. Facie infra antennas lamina transversa trigonata, antrorsum pro-
ducta armata; clypeo transverso, tumido, apice superiori bispinoso ;
antennarum articulo tertio subtus emarginato.

Hab, Ceylon.

Clypeus in the male transversely swollen, its anterior border
emarginate, its upper edge free, bispinose ; face below the antennz
armed with a large, transverse, horizontal, triangular plate, its
apex acute ; antenne in the same sex robust, the third joint sinuate
beneath. Thorax about one half broader than long ; sides straight
and diverging from the base to beyond the middle, thence con-
verging towards the apex. Elytra finely punctured.

28 MR. W. F. KIRBY ON GENERA AND SPECIES

A Synopsis of the Genera of the Chalcidide, Subfamily Eucha-
rine ; with Descriptions of several new Genera and Species
of Chalcidide and Tenthredinide. By W. F. Kirsy,
Assistant in the Zoological Department, British Museum.
(Communicated by Dr. J. Muriz, F.L.S.)

[Read 17th June, 1886. ]
(Puate I.)

Synopsis of the Genera of Eucharine.

I have here attempted to give a generic revision of the sub-
family Eucharine, which includes some of the most beautiful
and remarkable species of the Chalcidide. With few exceptions,
the species are extra-Huropean, and appear to be most numerous
in Tropical America and Australia. The species in the British
Museum represent at least fifteen well-marked genera, of which
only ten have been previously characterized. I have selected
only the most salient characters of each genus ; for minute and
detailed characters are unnecessary in the present imperfect |
state of our knowledge of the subfamily.

The Eucharine are large, strongly-sculptured, metallic-coloured
Chalcidide ; the abdomen is always more or less petiolated, and
is frequently raised and compressed, giving the insects some
resemblance to the Cynipide. From the Perilampine, to which
they have some resemblance, they may be distinguished by the
longer petiole, the absence of the stigmatic nervule, &c.

List of Genera of Eucharine, with indication of Types, and
characters of new Genera.

1. Evcnaris, Latr.,
Hist. Nat. Crust. Ins. xiii. p. 210 (1805).

Type, CYNIPS ADSCENDENS, Fabr.
Cynips adscendens, Fabr. Mant. Ins. i. p. 251. n.1 (1787).
Eucharis Kollari, Forst. Verh. Ver. ERheml. xvi. p. 91. n. 25

(1859).
Details: Westw. Thes. Ent. Oxon. pl. xxviii. f. 15, a-d (1874).
Hab. Kurope.

Panzer (Faun. Germ. Heft 88, pl. 10) represents the legs as
yellow, saying only “antenne tarsorumque ungule nigre;” and

OF CHALCIDIDE AND TENTHREDINIDA. 29

his insect thus appears to differ from that of Fabricius, who says
“ pedes pallide femoribus basi nigris.” Forster regarded them as
distinct ; and as Fabricius merely gave a brief diagnosis in his
Syst. Piez. p. 157, and this apparently founded on Panzer’s
figure, Forster was not unnaturally misled into redescribing and
renaming the Fabrician instead of the Panzerian insect.

2. OrnasEMA, Cameron,
Biol. Centr.-Amer., Hym. p. 104 (1884).

Type, ORASEMA FLAVIPES, Cam.

Orasema stramineipes, Cam. 1. c. p. 105 (1884).

Orasema flavipes, Cam. op. cit. pl. v. figs. 20,20 a-c, pl. vi. figs.
18, 18 a—e (1884).

Hab. Panama.

There seems to be some error about this species, as the descrip-
tion does not well agree with the figure. Judging by the figure,
I would refer Lucharis festiva, Fabr., E. rapo, Walk., and
Schizaspidia maculata, Westw., to this genus. All these are
South-American insects.

3. TRICORYNA, gen. nov.

Male. Antenne 11-jointed, thick, moniliform, striated and
finely pubescent; scape short; second joint short and trans-
verse ; third joint four times as long as broad, much thickened
beyond the middle; fourth narrower, longer than broad; the
remainder gradually decreasing in length, but not in width, to
the ninth ; the tenth shorter and narrower than the ninth, and
the eleventh minute. Head and thorax coarsely punctured,
moderately pubescent; scutellum gibbous, but not produced;
petiole longitudinally striated, moderately broad, about two thirds
as long as the abdomen. Abdomen about four times as long as
broad, tapering at both ends, and rather finely punctured; first
joint of tarsi very thick, and much longer than ail the rest put
together.

Type, Evcuaris Ienio, Walk.
Eucharis Iello, Walk. Mon. Chale. ii. p. 12 (1839).

Details: Haliday, Entomol. i. pl. P. figs. 3, 3a, 6 (1842).
Hab. Tasmania.

30 MR. W. F. KIRBY ON GENERA AND SPECIES OF

4, METAGHA, gen. nov.

Male. Antenne nearly naked; the third joint four times as
long as broad, and gradually thickened to the extremity ; the
remainder thickened at the extremity, and gradually decreasing
in length to the eighth, whichis only slightly longer than broad ;
the rest broken off; petiole short. First joint of the tarsi as
long or longer than all the rest together, but not thickened.
Abdomen obtuse at tip.

Type, Evcnaris Zauatres, Walk.
Eucharis Zalates, Walk. Mon. Chale. ui. p.13 (1839).
Details: Haliday, Entomol. i. pl. P. figs. 4, 4 a—-c (1842).
Hab. Australia.

5. PsrnogastEer, Blanch.,
Hist. Anim. Art. iii. p. 260 (1840); ? Brullé, Ins. Hym. iv.
p. 573 (1846).

Type, PstLoGasTER cuPREUS, Blanch.

Psilogaster cupreus, Blanch. l.c. (1842).

Hab. Egypt.

Brullé has figured a Tasmanian insect to represent this genus
(P. pallipes, Brulié, 1. c. pl. xxxix. figs. 1 & 2), and the British
Museum possesses many species congeneric with this from
Australia and Tasmania. It is, however, very probable that the
Egyptian insect may prove to be generically distinct from these,
in which case the Australian group will require a new name.

6. CHALCURA, gen. nor.

Male. Antenne pubescent, 12-jointed; joints 8-11 throwing
off a long pubescent ramus from the extremity on the outside.
Scutellum gibbous, unarmed. Petiole moderately stout, about
two thirds as long as the conical abdomen, and longitudinally
striated.

Female with scarcely the rudiments of the antennal rami.
Petiole short and broad. Abdomen oval. J irst joint of tarsi
nearly as long as all the remainder, but not expanded.

Type, HucHaris DEPRIVATA, Walk.
Eucharis deprivata, Walk. Ann. §& Mag. N. H. (8) vi. p. 359
(1860) ; Westw. Thes. Ent. Oxon. p. 155, pl. xxviii. fig. 6 (1874).
Hab, Ceylon.

CHALCIDIDE AND TENTHREDINIDS. 81

7. RHIPIPALLUS, gen. nov.

Male. Antenne pubescent, 13-jointed, with short joints ;

‘joints 4-12 throwing off long pubescent rami (thickened at the

Vere,

tips) from the outside, and shorter ones from the inside. Scu-
tellum gibbous, unarmed. Petiole moderately slender, as long
as the oval and moderately raised abdomen.

Type, Evcnaris votusus, Walk.
~ Hucharis volusus, Walk. Mon. Chale. ii. p. 9 (1839).

Details: Haliday, Entomol. i. pl. P. figs. 1, 1 a-d (1842).
Hab. Australia.

8. SrrinpuLa, Spin.,
Ann. Mus. xvii. p. 150 (1811).

Type, IcHNEUMON CYNIPIFORMIS, /oss?.

Ichneumon cyniformis (sic), Rossi, Mant. Ins. p. 125, pl. vi.
fig. G (1792).

Hab. 8. Europe.

9. Scurzasprp1a, Westw.,
P. Z. 8. 1835, p. 69.
Type, SCHIZASPIDIA FURCIFERA, Westw.
Schizaspidia furcifera, Westw. 1. c. (1835); Thes. Ent. Oxon.
p- 151, pl. xxviii. fig. 2 (1874).
Hab. India.

10, TETRAMELIA, gen. nov.

Male. Antenne 12-jointed; joints 3-11 each throwing off a
long pubescent ramus. Scutellum bidentate. Metathorax
throwing off a stout horn-like process on each side, curving out-
wards horizontally nearly as far as the level of the petiole, which
is slender and about as long as the height of the abdomen. The
latter is of an irregular shape, elevated, compressed, and rather

pointed in front, the first segment covering about half the
abdomen.

Type, Scuizasprp1a PLaciatTa, Walk.

Schizaspidia plagiata, Walk. Trans. Ent. Soc. (8) i. p. 885
(1862); Westw. Thes. Ent. Oxon. p. 152, pl. xxviii. fig. 11
(1874).

Hab. Para.

32 MR. W. F. KIRBY ON GENERA AND SPECIES OF

11. Lopnyrocrera, Cameron,
Biol. Centr.-Amer., Hym. p. 103 (1884).

Types, LopHyROCERA STRAMINEIPES and L. NIGROMACULATA,
Cam.
(1) Lophyrocera stramineipes, Cam. 1. c. pl. v. figs. 18, 18 a
(1884).
Hab. Panama.

(2) Lophyrocera nigromaculata, Cam. 1. c. p.104, pl. v. figs.
19, 19 a, b (1884).

Hab. Nicaragua.

12. Kapana, Cameron,
Biol. Centr.-Amer., Hym. p. 102 (1884).

Chirocerus, Brullé (nec Latr.), Ins. Hym. iv. p. 571 (1846).
Type, Evcnaris rurcata, fbr.

Eucharis furcata and E. flabellata, Fubr. Syst. Piez. p. 158
(1804).

Thoracantha fureata, Haliday, Entomol. 1. pl. P. figs. 2, 2.a-c
(1842).

Kapala furcata, Cam. 1. c. p. 103, pl. v. figs. 17, 17 a-d (1884).

Hab. South and Central America.

There are several species of this genus, from different parts of
South America, in the British Museum. Chirocerus furcatus,
Bruilé (Ins. Hym. pl. xxxviu. figs. 5, 5 a, 6, 1846), seems to me
to agree better with Thoracantha atrata, Walk. (Trans. Ent. Soc.
(3) i. p. 883, 1862), than with the Fabrician species.

13. THoracantHa, Latr.,
Cuv. Régne Anim. ed. 2, v. p. 297 (1829).

Galearia, Brullé, Spec. Hym. iv. p. 572 (1846).
Type, THoracantHa Larreruui, Gueér.

Thoracantha Latreillil, Guér. Icon. Réegne Anim., Ins. p. 415,
pl. lxvii. fig. 8 (1829-44).

Thoracantha Latreillii vel coleopteroides, Waterh. Trans. Ent.
Soe. ii. p. 196, pl. xvii. fig. 3 (1839).

Galearia violacea, Brullé, l.c. pl. xxviii. figs. 6, 6 a, 6 (1846).

Hab. Brazil.

Although Latreille specified no type, yet his description clearly
indicates this genus; moreover, 7. Latreillit was figured and de=
scribed as a Lhoracantha before any other species to which the
generic name could apply.

— eS

ee Be ae

CHALCIDIDHE AND TENTHREDINIDZ. 33

14. Larara, Cameron,
Biol. Centr.-Amer., Hym. p. 102 (1884).

Type, LiraTa LUTEOGASTER, Cam.
Lirata luteogaster, Cam. l.c. pl. v. figs. 16, 16 a (1884).
Hab. Panama.

15. UROMELTA, gen. nov.
‘Thoracantha, Brullé (nee Latr.), Ins. Hym. iv. p. 572 (1846).

Type, THoRACANTHA SsTRIATA, Perty.
Thoracantha striata, Perty, Del. Anim. Art. p. 185, pl. xxvii.
figs. 15, 16 (1834).
Hab. Amazons.

Table of Genera of Eucharine.

A. Scutellum not bidentate.
B. Antennz simple in male.
C. Antennz moniliform.
D. Abdomen compressed, ascending ... 1. Hucharis.
DD. Abdomen not compressed, nor as-
cending.
E. First joint of tarsi much
PRGEROMED civccsssavecececae--s 3. Tricoryna.
EE. First joint of tarsi very long,
but notthickerthan the others 4. Metagea.
CC. Antenne not moniliform.

F’. Joints of antenne long .................. 5. Psilogaster.
FF’. Joints of antenne short ..............2.65 2. Orasema.
BB. Antenne ramose in male.
G. Antenne ramose in male .................0065 6. Chalcura.
GG. Antenne biramose in male .................. 7. Rhipipallus.

AA. Scutellum bidentate.
H. Scutellum of moderate size.
I. Antenne simple in male ..............0.ssseeeee 8. Stilbula.
II. Antennzx ramose in male.
K. Metathorax unarmed ...............0.000 9. Schizaspidia.
KK. Metathorax with a strong lateral pro-
jection.
L. Metathoracic processes curving
eo a: a 11. Lophyrocera.
LL. Meiathoracic processes consisting
of two diverging horizontal teeth 10. Tetramelia.
HA. Scutellum often as long as the abdomen.
M. Scutellar processes covering the whole abdo-

men.
N. Scutellar processes very broad ............ 13. Thoracantha.
NN. Scutellar processes long, contiguous, and
tapering to the extremity ............... 15, Uromelia.

MM. Scutellar processes long and slender, gene-
rally curving inwards towards the tips.
O. Third joint of antenne as long as all

the rest together ...............000008 14. Lirata.
OO. Third joint of antenne not much
fonger than fourth, .........scsecee 12. Kapala

LINN. JOURN.— ZOOLOGY, VOL. Xx. 3

34 MR, W. F. KIRBY ON GENERA AND SPECIES OF

TENTHREDINID 2.

HyYLoTroMIn».

1. Hemripraneura CameEroni. (PI. I. fig. 10.)

Long. corp. 8 millim., exp. al. 20 millim.

Female. Head, antenne, knees, tibie, tarsi, and abdomen (ex-
cept base) black ; thorax, base of abdomen, coxe, trochanters, and
femora (except tips) luteous ; on the abdomen the five first ventral
segments are luteous, but the first two dorsal segments only are
wholly luteous, the sides of the third and fourth being clouded
and shading into the deeper black of the following segments ;
wings purplish hyaline, with a strong smoky tinge, especially at
the base.

Hab. Rio Grande do Sul (Lhering).

Not closely allied to any known species.

SELANDRIIN #&.

2. SELANDRIA MERIDIONALIS. (PI. I. fig. 12.)

Long. corp. 6 millim., exp. al. 17 millim.

Female. Inky black ; mesothorax above and front of scutellum
dullred, prothorax yellower; wings smoky hyaline, with blackish-
brown nervures, and a black dot in the second submarginal cell.

Hab. Rio Grande do Sul. .

TENTHREDININA. ~

3. MacropHya (?)Saunpersr. (PI. I. fig. 11.)

Long. corp. 11 millim., exp. al. 28 millim.

Female. Black; labrum,scutellum, a spot at the extremity of the
hind coxe, an oval spot on the side of the fifth and a round one on
the side of the sixth segment yellow ; cox otherwise black, four
front femora and tibie yellowish red, the middle tibiz just tipped
with black ; hind femora entirely red, except a small black spot
at the extreme base; all the tarsi and the hind tibie black:
wings iridescent hyaline, rather more clouded beyond the middle ;
nervures and stigma castaneous, the latter paler towards the base ;
lanceolate cell contracted.

Two specimens from Albania, from the late SirS. E. Saunders’s
collection.

Allied to IL. femoralis and corallipes, Eversm., in both which
species the cell is subcontracted.

CHALCIDIDE AND TENTHREDINIDS. 35
CHALCIDID A.
CHALCIDIN &.

4, TRICHOXENIA CINERARIA, Walk. (PI. I. figs. 4, 5.)

Long. corp. 11 millim., exp. al. 18 millim.

Male. Black; antenne very thick, with the second and third
joints very short; scape red at the tip above; face clothed with
silvery pubescence, shading into golden yellow above; thorax
black, variolose; tegule black, a bright golden spot on each
side at the base of the scutellum, and the sides of the metathorax
fringed with golden hairs above ; abdomen with the first segment
black, variolose, its sides and extremity above, and the sides and
upper surface of the remaining segments thickly clothed with
bright golden hairs; abdomen black beneath; femora black,
clothed with silvery hairs; tibie and tarsi with a reddish shade,
and hind tibiz clothed with yellow hairs; spines of the hind
tarsi golden beneath; wings and general structure as in the
female.

Hab. Australia (Du Boulay).

This beautiful insect, which strongly contrasts with the black
female, has a striking general resemblance to some species of
Mutilla; but it is worthy of note, first, that this resemblance
is confined to the male, and, secondly, that the Mutille which it
most resembles are not Australian species.

ee ee eee ee

I am indebted to Signor P. Magretti for permitting me to de-
scribe the three following species, taken by him in the Soudan.
All the other species in this paper are in the collection of the
British Museum.

5. Stomatoceras Macrerri. (PI. I. fig. 7.)

Long. corp. 9 millim.

Female. Black, variolose, with silvery hairs and down; antenne
—-scape red, black at the tip ; four following joints red, with short
white pubescence, the rest black; third joint with avery distinct
patch of white pubescence on the outside: tegule, stylus, and
sides of first segment and under surface of abdomen shading into
dullred ; hind coxe and last segment of abdomen before the stylus
clothed with silvery hairs ; legs black, shading into reddish, espe-
cially on the inner surface; middle femora somewhat thickened, all
the tibiz with a strong projecting spine beneath, and middle tibiz
armed with strong, stout, short bristles or spines, which are much

36 MR. W. F. KIRBY ON GENERA AND SPECIES OF

more distinct upon the tarsi; hind femora armed with three large -
obtuse teeth; wings hyaline, with two brown bands.

Hab. Kassala (Magretti).

Allied to Halticella liberator, Walk., from Natal.

In describing the genus Stomatoceras, I stated that the meta-
thorax was unarmed. In the two species here described it is
armed with two strong teeth on each side; and I should expect
to find this to be the case in well-preserved specimens of the
previously described species of the genus.

6. STOMATOCERAS DIVERSICORNIS. (PI. I. fig. 8.)

Long. corp. 4 millim.

Female. Closely allied to S. Magrettit, of which at first [regarded
it as only a small variety. Black, variolose, silvery pubescence
very slight; scape of antenne and joints 2, 3, 4, and 11 wholly red ;
the joints longer and slenderer in proportion than in §. Magrettii ;
four front femora black; tibiz and tarsi red, the former with a
black streak on the inner side; hind femora black and shining, the
middle tooth distinct, the others merely undulations; a small
red spot at the base ; hind tibie black, red at the tip, hind tarsi
red; all the claws black ; tegule black ; wings hyaline, fore wings
with two brown bands; abdomen reddish beneath, along the
sheath of the stylus, which is comparatively short ; metathorax
with two strong teeth on each side.

Hab. Kassala (Magretti).

7. Cuatcrs Kassatensis. (Pl. I. fig. 9.)

Long. corp. 5 millim.

Male. Black ; head, thorax, and scutellum moderately closely
variolose, each pit with a setigerous depression in the centre; the
interspaces finely punctured; abdomen very finely reticulose ;
scutellum edged behind with silvery-white hairs ; lower orbits,
cheeks, pleure, terminal abdominal segments, and coxe more or
less clothed with silvery-white pubescence (antennz wanting);
tegule ivory-white ; coxe black; four front femora black, tipped
with white; four front tibis white, with a short black line on the
inner side; all the tarsi white, with the claws black; hind femora
thickened, denticulated beneath, red, with a large white spot at
the tip; hind tibiz white, with a reddish-brown ring at the base ;
wings hyaline.

Hab. Kassala (Magrettt).

Allied to C. amenocles and C. restituta, Waik.

IDIDA, AND

DINT DAE .

.
4

CHALC

SPECIES OF

t Limp

Hanhar

7
4

PEN cee

Hammond. delet ith

Ce ae
a*
ye
:

%
D
>

a
. ine
*s\

=
U

‘faq ote

a ty “ ie ae

|
|

CHALCIDIDA AND TENTHREDINIDZE, 37

EvcHARIssiIn&.
SACCHARISSA, nov. gen.

Antenne 18-jointed, scape rather long, second joint small,
transverse, third and fourth simple, third longest, fifth to fifteenth
short, with a series of long bristly fin-shaped projections on the
upper surface, that on the fifth segment half the length of that
on the sixth, the rest gradually decreasing until on joints 16-18
they almost disappear ; scutellum with a long spear-shaped pro-
jection behind ; head very narrow, transverse; ocelli arranged in
a straight line; petiole short; abdomen short, ascending. (The
abdomen is damaged in the type, and the figure may not there-
fore give the correct shape.)

Type, Evcnaris continaens, Walk., from Borneo. (Pl. I.
figs. 6, 6 a, 6 6.)

The species of Hucharissa are South-African insects, and their
scutellum is unarmed.

KUCHARIN.

8. Rurpreautus Cameron. (PI. I. figs. 2, 2a.)

Long. corp. 5 millim.

Male. Coppery, rugose ; head and metathorax greener, antennz
brown, the rami less thickened than in &. volusus, Walk.; legs flavo-
testaceous ; hind coxe globular, very dark green; petiole purplish
cupreous, longitudinally striated; abdomen flavo-testaceous,
brown on the back at the base; wings yellowish hyaline, with
yellowish and unusually distinct nervures; a large smoky cloud
below the cubitus.

Hab, Australia (?) or Celebes (?).

This very fine species much resembles Chalcura deprivata, Walk.,
in size and appearance.

DESCRIPTION OF PLATE I.

*Fig. 1. Schizaspidia Murrayi, Kirb., Fig. 6a. S. contingens. Antenna.
6b. Ditto. Scutellum.

2. Rhipipalius Cameroni,Kirb., 8 . 7. Stomatoceras Magrettii,Kirb., 2 .
2a. Ditto. Antenna. 8. S. diversicornis, Kirb., 2.

3. Stilbuia cynipiformis, Rossi, 3. 9. Chalcis Kassale, Kirb., 3.

8a. Ditto. Seutellum. 10. Hemidianeura Cameroni, Kirb.,
4, Trichoxeniacineraria,Walk., 3. ned

m. Ditto, 9’. 11. Macrophya Saundersi, Kirb., 2.

6. Saccharissa contingens,Walk., $. 12. Selandria meridionalis,Kirb., 2.

* §. Murrayi, Kirby, Ann. & Mag. Nat. Hist. (5) xiii. p. 403 (1884), from
Tongatabu.

38 MR. G. SIM ON THE OCCURRENCE ETC. OF

Occurrence of Lumpenus lampetriformis on the North Coast of
Scotland; with Notes on its Habits, Food, and the Ground
it frequents. By Grorcre Sim. (Communicated by Dr.
Francis Day, F.L.8.)

[Read 17th June, 1886.]

LUMPENUS LAMPETRIFORMIS, the subject of the follewing re-
marks, is anorthern form of the tribe Blenniide. It was described
by Collett, a naturalist who accompanied the Norwegian North-
Atlantic Expedition, 1876-78. According to this writer, Lum-
penus appears to be a rather common fish in most localities
along the Norwegian coast. As to its geographical distribution,
I cannot do better than quote from a paper on the species
under notice by Dr. F. Day. He says :—“ Up to the present
time (June 1884) this species has been recorded from the coast
of Greenland and Iceland. It is common off Spitzbergen and
on the shores of North-western Europe as far south as the
Cattegat. In the north its range extends certainly as high as
80°.” Collett says, “The southern limit of its range is probably
Bohuslan, in Sweden, one or two individuals having been obtained
off Gothenburg, 58° N.” This latter writer’s remarks on the
species are of a general description ; and nothing is said as to
the nature of the ground Lumpenus frequents, the other living
organisms that accompany it in its haunts, nor does he say a word
as to the food of this interesting species. To these points the
following notes more particularly refer.

The discovery of this species as British is of very recent date,
the first example having been brought to ight by Prof. M*Intosh
of St. Andrews in May 1884. This one he obtained fifteen miles
off St. Abbs Head, while engaged in a series of observations on
trawling, undertaken at the instance of the Scottish Fishery
Board. Prof. M*Intosh sent the specimen for examination to
Dr. Day, who has described and given an excellent figure of it in
the ‘ Proceedings’ of the Zoological Society of London.

The second specimen found in Britain I obtained on an Aberdeen
trawl-vessel on April 14th, 1885; and being quite unknown to
me, and not having at that time seen Dr. Day’s description of
Prof. M°Intosh’s specimen, I made a sketch of the one now

LUMPENUS LAMPETRIFORMIS ON THE COAST OF SCOTLAND. 39

under notice and sent it to Dr. Day, who at once recognized it
as L. lampetriformis, although differing very much in the form
of the caudal fin from Prof. M‘Intosh’s specimen, the dif-
ference being that M*Intosh’s specimen had the outer caudal
rays elongated beyond the central ones for nearly half the length
of the caudal fin; while in wy specimen the central ray was
longest, the others decreasing in length on each side of it, thus
giving the tail a lanceolate form, or, as Collett says, an “acumi-
nate form.” Strangely enough, all my specimens have the same
form of tail. On receipt of this information, the fish was for-
warded to Cheltenham for examination by Dr. Day; and in due
course I received a note from him stating that the specimen was
a female Lwmpenus, while that of Prof. M*Intosh was an old male ;
and this he considered would explain the difference in the form of
the tail.

The occurrence of this specimen Dr. Day recorded in ‘ Nature’
for July 9th, 1885. Thus stood the matter until March 25th,
1886, on which date a second example came to my hand, and on
March 31st eleven more; while from April 1st till May 31st fifty-
Seven specimens have made up my findings. This will indicate
pretty clearly that the species is not so rare upon the coast of
Britain as we were at first disposed to think.

In general appearance, Lwmpenus approaches very nearly
to that of our common Blennies, Centronotus gunnellus and
Zoarces viviparus, to which it is closely allied ; but on close exa-
mination it is seen to differ from them considerably, both in
external colouring, fin arrangement, and internal organization.
To give a minute description of the external appearance of Lwm-
penus is quite unnecessary, as this has been done by Dr. Day
in the article already mentioned, as well as by Collett. My
purpose, then, is merely to note the points where the three
species differ and where they agree, in so far as I have been able
to observe the same. Comparing Luwmpenus with the Spotted
Gunnel (Centronotus gunnellus), we find in both the spinous
dorsal fin, the number of rays in Centronotus being 76 to 78,
while in Lumpenus the number is 72 to 74; the latter number I
have only once observed. In Centronotus the ventral fins have
disappeared, and are represented by two short stout spines ; in
Lumpenus, however, these fins are comparatively well developed,
each haying one spinous ray and four soft ones. In the matter

40 MR. G. SIM ON THE OCCURRENCE ETC. OF

of dentition, Centronotus has a single row of conical teeth in both
jaws, becoming shorter as they approach the angle of the mouth ;
sometimes there exist a few irregularly set teeth in the upper jaw
in addition to the regular row; several strong teeth on the
vomer; four patches, two on each side, of irregularly set upper
pharyngeal teeth, and two long patches below, also set irregu-
larly. On the other hand, Lwmpenus is furnished in the upper
jaw with numerous irregularly set teeth, the outer ones being
conical, the others sharp-pointed, and all decreasing in length as
they approach the angle of the mouth. The lower jaw is set with
one, sometimes two rows of long incurved, rather sharp teeth,
not of uniform length, and all becoming shorter laterally ; none
on the vomer or tongue. ‘There are two irregularly set patches
of upper pharyngeal teeth, each point of which is finely tinged
with yellow; and four lower pharyngeals forming two sides of a
triangle set confusedly, and without the yellow colour of the
upper ones. In the number of vertebra, Centronotus outnumbers
Lumpenus by 3, the latter having 80 and the former 83.

But the greatest departure which Centronotus makes from
Lumpenus isin the form of the stomach and intestines. As will
be seen by the accompanying figures, the intestines of Centronotus
in some cases show little change in the stomach from the intestines
proper; that is, there is no very distinct line of demarcation
between them; presenting more or less the form of a simple
tube and are devoid of ceca. I have said in some cases. This
will be best understood by referring to figs. 1 and 2, both of
which represent the intestines of Centronotus, both drawn from
specimens caught by myself and while the intestines lay in situ.
This is a change in the internal arrangement of a species I have
never before observed carried to such a length, although the indi-
viduals of no species have the intestines formed or laid exactly
alike. The result of my investigation goes to show that fig. 1,
or something much like it, is the most usual arrangement to be
met with in Centronotus gunnellus. On the other hand, Lum-
penus has what may be called a properly developed stomach,
marked off from the intestines by its two cecal appendages, and
the intestinal track forms several convolutions in its course.

With reference to the ceca in Lumpenus, I find they differ
much in length in different individuals, even although these indi-
viduals are of themselves the same size. In some examples

LUMPENUS LAMPETRIFORMIS ON THE COAST OF SCOTLAND. 41

these appendages are about 3 inch long, while in others they are
barely half that length. Is this difference in the stomach and
intestines due to, or is this arrangement necessary to meet, the

Figs. 1 and 2. Stomach and intestines of Centronotus gunnellus.
Fig. 3. Stomach and intestines of Zoarces viviparus.
Fig. 4. Stomach and intestines of Lumpenus lampetriformis.

different conditions of food and habitat sustained by Luwmpenus
in comparison with its allies ?

Turning to the Viviparous Blenny (Zoarces viviparus), as com-
pared with Lwmpenus, the difference in some points is greater than
in the case of the latter and Centronotus ; while in other particulars
Lumpenus and Zoarces approach much more nearly to one
another. In external form, Lwmpenus is long and somewhat

42 MR. G. SIM ON THE OCCURRENCE ETC. OF

cylindrical, becoming compressed towards the caudal extremity,
while in Zoarces the body is compressed along its whole length.
Its dorsal fin has soft rays, 83 in number, after which are a few
spinous rays ; and the ventrals are well developed, having all the
rays soft. In dentition Zoarces differs from Lumpenus in having a
double row of teeth in both jaws, merging into one row, and
becoming less in size as they approach the angle of the mouth.
Six rows of upper pharyngeal teeth, viz. three on each side, the
first a single row, the other two double. Lower pharyngeal teeth
set ina double row which form a triangle; no teeth on the vomer or
tongue. The number of its vertebre is 116, whereas in Lumpenus,
as already pointed out, these bones only number 80. In the
case of the internal viscera, there is a great similarity in the
two species under notice as compared with that of Centronotus.
But the point wherein lies the greatest difference between these
closely allied species is the fact of Zoarces bringing forth its
young alive *.

Lumpenus is not viviparous. As to its time of spawning
I cannot speak with certainty; at the end of April some of
those I found had the roe pretty well advanced, much more
so than was the melt of the males caught at the same time.
Then, again, on May 20th I found one specimen measuring
only 12 inch. Assuming that the species breeds only once a
year, and taking into account the fact that those caught in April
had spawn fully halfway advanced towards maturity, we may
safely conclude that the spawning takes place in July or August,
and that my 12-inch specimen was one of last year’s young. On
being boiled, the vertebra of Zoarces become a fine deep-green
colour, those of Lwmpenus remain white.

It will thus be seen that although Lwmpenus has something in
its composition similar to both our common species; it is at
the same time so widely different that no doubt need be enter-
tained as to its being a distinct and well-marked species, nor any
difficulty experienced in recognizing it from its congeners.

* On April 22, 1886, I had two specimens of this fish—one of them with its
young, of which there were thirty, almost ready for expulsion, each measuring
21 inches; and in addition to these there were thirty-one more, evidently the
young of the former year, which had not been given birth to; they were all
firmly fixed together and in a hard wasted condition, yet capable of being

separated individually. Why they had not been born at their proper time is a
mystery difficult of solution.

LUMPENUS LAMPETRIFORMIS ON THE COAST OF SCOTLAND. 43

The only locality where Lumpenus has yet been found on
the north-east of Scotland is in a deep hollow in the Moray
Firth, which runs almost parallel to the land, and extends
from opposite the village of Roseharty on the Aberdeenshire
coast to near Tarbetness in Rosshire, where it bifurcates, the

DUNCANSBAY HEAD

Sxercu-Mar or Moray Firtn AND REGION AROUND, indicating
the position of the ‘‘ Witch Ground.”

other point running in the direction of the Cromarty Firth.
Reference to the sketch-map above will show the “lie” of
this ground. ‘This hollow slopes gradually from the shore,
but rises abruptly on the north-east or seaward side. It is
from six to ten miles off shore, and 35 to 110 fathoms deep,

44 MR. G. SIM ON THE OCCURRENCE ETC. OF

the greatest depth being at the Aberdeenshire end. This track
is known to the trawl-fishers as the “ Witch Ground.” The
reason for this name being given to it is because along a portion
of the area, principally off “ Covesea Scars,” enormous numbers of
Pleuronectes cynoglossus, the Pole or Craig Fluke, which these
trawlers know as the Witch-Sole, are taken: hence the “ Witch
Ground.” The bottom of this hollow is in some places composed
of grey muddy sand, while in other parts it is principally
covered with mossy matter. Concerning this latter material,
considerable difficulty was experienced by me in coming to any
definite conclusion regarding it. The question being, How came
peat to be in such a locality? Was it matter washed by some
river into this ground, or how came it to be there? It came first
under my notice whilst examining the stomachs of the Witch-Sole.
In many cases the stomach and intestines of this species contain
portions of this peat which had been taken in along with the
creatures upon which it feeds. However, during the storms
which occurred in the month of January last immense sheets of
peat were cast on our beach, which shows that at some long past
age a forest of birch, hazel, and other trees had flourished, as
also marshes filled with Sphagnum, Polytrichum, and other fresh-
water-loving plants, where now swells the mighty waves of the
North Sea. The remains of these sylvan beauties we now find
composing the large masses of peat to which reference has been
made. Itis amongst this soft peaty ooze that our fossorial little
friend Lumpenus loves to dwell, and amongst and on the surface
of the same material where it finds its varied food, and where,
in its turn, it becomes the food of its ever-watchful neighbours
Pleuronectes cynoglossus.

The other living organisms that have come under my notice
from the “ Witch Ground” are, first, a few of the common
Zoophyta, viz. Lubularca indivisa, Sertularia abietina, Thuiaria
thuia, known in the locality as the bottle-brush weed; the “ sea-
pen” (Pennatula phosphorea), Alcyonium digitatum, locally known
as “dead men’s fingers,’ and the common Llustra also occur, but
the last-named is perhaps only washed from the hard ground
on the east of this great hollow.

Threading their way through the above-mentioned forms are
those beautiful Ophiuroids Amphiura Chiajit, A. filiformis,
Ophiocnida brachata, Ophioglypha albida, O. lacertosa, and

LUMPENUS LAMPETRIFORMIS ON THE COAST OF SCOTLAND. 45

Ophiothrix pentaphyllum. Here and there very sparingly dis-
tributed is also to be found that link which joins the present to
the past—our elegant British Crinoid, Antedon rosacea.

From the depths of this vast aquarium we also sometimes
obtain a specimen of that fine, but most repulsively odoured
Starfish, Stichaster rosews, accompanied occasionally by Palmipes
membranaceus.

Here also is that form which the late Prof. Forbes calls one of
the rarest and most beautiful of our native Starfish, Hippasteria
plana; beautiful it unquestionably is; but in this long valley
it is by no means rare, dozens are sometimes brought up at one
drag of the net. Accompanying it, but in very much fewer
numbers, is its exquisitely coloured little brother, Porania pul-
villus. And yet again, along this great expanse of mud, sand,
and peat, live Astropecten irregularis and Luidia Savignyi, the
latter a form more troublesome and tantalizing in its habits
than any of our Starfish with which the lover of Nature may
come in contact. Hundreds of this fine form are brought up
both by traw] and line, but only to fall in pieces on coming to the
surface, or, as Forbes says, “in a minute he proceeds to dissolve
his corporation.”

Of the Echinoidea which frequent this ground, we have Echino-
cyamus pusillus, Spatangus purpureus, Brissopsis lyrifera, and
Echinocardium cordatum, while in each suitable resting-place are
specimens of Priapulus caudatus, a favourite food of the ‘* Witch-
Sole.” Aphrodita aculeata is also present in abundance.

The Crustacea to be met with along these grounds are numerous
and highly interesting. Heading the list comes Cancer pagurus,
LInthodes maia, Nephrops norvegicus, Corystes cassivelaunus, Calo-
caris Macandree (the latter is eaten in enormous numbers by the
Witch-Sole), Galathea squamifera, G. dispersa, and G. Andrewsi,
Pagurus Bernhardus, P. Thompsoni, and P. Prideauxi (always
accompanied by its humbler friend Adamsia palliata), Portunus
holsatus, P. pusillus, and P. depurator, Crangon vulgaris, C. bispi-
nosus,and O.trispinosus, Hippolyte spinus, Callianassa subterranea.
Pandalus annulicornis, various species of the genus Diastylus,
accompanied by their kindred form Eudora truncatula, and
several of the Mysidz literally swarm. These are all the higher
Stalk-eyed Crustacea; but in addition to the above list there is
another stalk-eyed form, of which I have taken several specimens

46 MR. G. SIM ON THE OCCURRENCE ETC. OF

from the stomach of the Witch-Sole; it is not, so far as I am
aware, described in any British work, and it therefore remains
unnamed.

In Sessile-eyed Crustacea of the orders Amphipoda and Isopoda
the ground is very rich. As to the identity of Anonyx denti-
culata, Hypera galba, and Arcturus longicornis there is no
doubt; but many others are in my possession which are not
yet identified. Pycnogonum littorale is abundant.

Turning to Molluscan life, the list is a fairly large one. First
of all, on account of its numbers, delicate structure, and fine sculp-
turing, comes that lovely little form Philine scabra, followed by
Nucula nitida, Solen pellucidus, Tellina fabula, Scrobicularia
prismatica, Psammobia Kerroensis, Donax anatinus, Mactra stul-
torum and MM. solida, Pleurotoma turricula, Tornatella fasciata,
Dentalium entale, Venus lincta, and Aporrhais pes-pelicani ; while
Pholas crispata finds a congenial home in the easily perforated peat.
Many other shells have been seen, but not living; so that in
all likelihood they have been merely washed by the waves to
the “ Witch Ground” after the death of their occupants.

Food-fishes other than the Witch-Sole are scarce and of bad
quality along this ground; and, according to the trawl-fishers,
many of the Cod and Haddock caught there have some wound
upon them, or are otherwise lean and out of condition. For this
reason some of the fishermen give this hollow the name of the “ Fish
Hospital,” because they think the sickly fish have come in so
that they might heal their sores and recruit their strength.

Such, then, is an outline, necessarily very imperfect, of the
living organisms which frequent the ground which Lumpenus
haunts and in which it burrows. Jor the latter statement
there is perhaps no absolute proof; but that it is fossorial
Tam strongly inclined to think, from the fact of its always being
caught by the trawl ground-rope. In the narrow openings
between the outer coils of this rope itis generally firmly jammed,
invariably accompanied by masses of sand, mud, and peat, of
which, as already stated, the bottom is composed.

It would seem that the ground-rope, in cutting through the
uneven surface, comes upon Lumpenus in its retreat, and fixes the
creature in its folds before there is any chance of escape. Were
the fish a free-swimmer only, the chances are very shght for such
a slender form to be caught at all.

LUMPENUS LAMPETRIFORMIS ON THE COAST OF SCOTLAND. A7

The next question to which attention had to be directed was,
upon what does Lwmpenus feed? Although many of the fish
obtained were so much injured as to be rendered useless as
specimens, fortunately the stomach of each had escaped muti-
lation, and therefore the contents were at my disposal; another
favourable circumstance was that each stomach was well filled.
Microscopical examination of the contents of their stomachs
reveals the fact that the food of Lwmpenus is almost a purely
crustacean one, confined in a very large measure to the Entomo-
straca and Copepoda; but in addition to these, I have found in
several numerous immature specimens of the two genera Dia-
stylus and Edora, minute bivalve mollusks, annelids, and several
very small fish-scales, minute starfish of the genus Amphiura,
a crustacean evidently parasitic (this I infer from the fact of its
being furnished on each foot with a strong long circular claw or
hook), very small forms of Priapulus caudatus, and a number of
brown pear-shaped objects quite unknown to me; also sessile-eyed
crustacea (Amphipoda).

As already stated, the Entomostraca and Copepoda largely
predominate, the species in greatest number being what seems
to me Dactylopus tisboides of Brady. Along with it is another
form of the same genus, and very like D. tisboides in every way
except that the last abdominal segment terminates in a long sharp
cylindrical telson, at each side of which, and somewhat under it,
are two short terminal segments from which issue several sete
of unequal length. Another creature presenting itself in some
abundance is in general form somewhat like Jdotea parallela of
Bate and Westwood; but differs from that species, first, in being
very small, and in having its body divided into thirteen segments
instead of nine as in Jdotea, and also in having the first pair of
feet very powerful, terminating in equally strong didactyle hands.
Its name remains unknown to me.

Then follow two species of the genus Oythere of Baird ; they
occur in about equal numbers; one seems to me to be Cythere
minna; but if Baird’s figure and description are correct, those I
have cannot belong to that species. OC. minna, according to Baird,
is “ obtusely rounded on the anterior extremity ;” my specimens
are equally acute at both ends. Can it be a species not described
by the author just quoted? ‘The second form answers to the
figure and description of Cythere pellucida of Baird.

48 PROF. P. M. DUNCAN AND MR. W. P. SLADEN ON THE

In naming these latter creatures specifically, I have done so
with considerable hesitation, as it must be confessed my know-
ledge of these groups is much too limited to warrant my speaking
with any great degree of certainty.

Thus ends my notes on Lumpenus lampetriformis ; and I trust
its habits and history have not been left shrouded altogether in
the darkness in which I found them.

Since writing the foregoing I got, on June 5th, five more
specimens of Lwmpenus, one of them, a female, carrying spawn,
which would have been deposited within a fortnight or three
weeks at latest. This brings my supposition relative to the
time of spawning to be pretty nearly correct, viz. the end of July
or beginning of August.

The longest specimen mentioned by Collett was 412 millim. ;
my longest one was 123 inches.

On the Anatomy of the Perignathic Girdle and of other Parts
of the Test of Discoidea cylindrica, Lamarck, sp. By Prof.
P. Martin Duncan, F.R.S., and W. Percy Stapen, F.G:S.,
Sec. L.S.

[Read 17th June, 1886. ]

DISCOIDEA OYLINDRICA, the Calerites cylindricus of Lamarck, is
one of the commonest of the Echinoidea from the Upper Creta-
ceous strata; and its shape and internal casts in flint are familiar
to all geologists. Desor, Wright, and Cotteau have described
the species; and the last-named paleontologist has enlarged the
generic diagnosis of Discotdea in consequence of some morpho-
logical details which had been elaborated by himself and some
previous observers, especially E. Forbes and Lovén.

Discoidea cylindrica has five basal plates in its apical system,
and the fifth or the posterior one is not perforated for a genital
duct. But the paleontologists just mentioned found a perforated
fifth basal in species which they felt bound to classify in the
genus Discoidea. Lovén, speculating on this association of im-
perforate and perforate basals in different species of the same
genus, considered it an instance of evolution during time.
Cotteau extended the generic diagnosis, and added to that of
Desor the following :—‘‘Apical system compact, subpentagonal,

—— a ee a ee

ANATOMY OF THE TEST OF DISCOIDEA CYLINDRICA. 49

presenting in some species five genital plates which are per-
forated, and in some other species a complementary imperforate
plate in the position of the odd generative plate”? (Heh. du Dépt.
de la Sarthe, 1869, Supp. p. 412). Desor had restricted his diagno-
sis to the structures with whicli he was acquainted ; but his speci-
mens do not appear to have been good ones, for he stated that the
apical system is small and rarely distinct in its details; but he
noticed that the odd genital plate is not perforated by the duct,
and that the ocular plates are very small (‘ Synopsis,’ p. 175).

The late Dr. Wright is at issue with all the other describers
of the species about the extent of the madreporite; and his de-
scriptions of the ambulacra and interradia of Discoidea cylindrica,
upon which he placed a specific value, are doubtful. There are
therefore many reasons why this familiar form should be studied,
and especially as in one of the many beautiful specimens in the
British Museum the perignathic girdle is exposed and can be
understood. We propose, therefore, offering the results of our
study of the most important details of the test in this and
some other species of the genus, and we trust that some of
the differences of opinion have now been settled, and that some
fresh anatomical details have been utilized.

Apical System.—The specimens examined in reference to this
and all the other anatomical structures are in the British Museum,
the Museum of Practical Geology, Jermyn Street,and one is in our
possession. ‘The full-grown specimens show how small the apical
system is in relation to the dimensions of the test, that there are
five basals, none being complementary, and that whilst four of
them (Nos. 1 to 4) are perforated by genital ducts, all of the five
are penetrated by the madreporite. ‘There are faint or decided
grooves between the basals. In the largest specimens the distri-
bution of the madreporite is well shown ; and in No. 4663, Brit.
Mus., the fifth basal is large and pentagonal, and is marked only
by a few of the pores of the madreporite ; and in a smaller spe-
eimen which is half-grown, belonging to us, the pores are absent
in the fifth basal.

Dr. Wright (Monogr. Cret. Echin., Pal. Soc. Lond. 1874, vol. i.
pt. vi. p. 207, pl. xlvi.) refers to the apical system. The type he
examined has a perfect apical system, and each of the basals is
perforated by the madreporite, and four basals are perforated by
the genital ducts. But the author, by an unfortunate oversight,
states that the madreporite is only seen in the right anterior

LINN. JOURN.—ZOOLOGY, VOL. XX. 4

50 PROF. P. M. DUNCAN AND MR. W. P. SLADEN ON THE

basal. This is erroneous, and so is the figure given in his plate.
The same specimen shows the very small size, comparatively, of
the radial plates (oculars).

It is evident then that the adult forms of Discoidea cylindrica
have all the five basals perforated for the madreporite, and all,
except the fifth, perforated for the genital duct.

The Ambulacra.—Wright was the first to point out that the
simple, straight condition of the pairs of pores was not univer-
sally found in the species, but that the pairs on the actinal area
might fall into a biserial arrangement.

Now it appears to us that allowance must be made for is age
and size of the tests in considering the particular distribution of
the pairs of pores ; for in the largest specimens the arrangement
of the pairs differs from that seen in the immature forms, espe-
cially near the peristome. Again, there is a point which must
be remembered, and it is that in full-grown specimens the am-
bulacra are flush with the test above the ambitus and raised
above its level actinally. The poriferous zones are on the slant
of the raised surface actinally; and it is on this slope that the
ereatest crowding of the pairs of pores happens. Above the
ambitus the simple arrangement of the pairs in one row is inva-
riable ; and it is seen in small forms even near to the peristome.
Asarule, the pairs are not crowded above the ambitus; and
they may be larger there than actinally, where the crowding may
be considerable.

The height of the ambulacral plates differs according to position
and age. They are high near the peristome in young forms, and
low close to the ambitus; and they are less unequal in adults.
The horizontal sutures between the plates are often slightly
furrowed.

The plates may be primaries, or compound with two or three
constituents. The plates near the apical system and extend-
ing towards the ambitus are primaries. Taking the specimens
in the British Museum, Nos. 38742, 728, and E180, and also the
half-grown specimen in our possession as examples, it is to be
observed that near the radial plates there are low, broad primary
plates, each having a pair of pores placed close to the interradial
suture*. The pores are large and separated by a ridge; and

* The late Dr. Wright (op. cit. p. 207) seemed to separate the poriferous
zones from the ambulacral plates; for he wrote, “The poriferous zones are

ANATOMY OF THE TEST OF DISCOIDEA CYLINDRICA. 51

they are very obliquely placed in reference to the transverse
sutures of the ambulacral plates. The outer pore is aboral, and
is very close to the edge of its plate and the interradium ; and the
inner pore is adoral, and either close above Fie. 1,

the line of suture or on it; and in the first uence
instance a narrow linear prolongation of the (% 429 O90
pore may occasionally be seen passing down to

the suture between the plate and the next in Fig. La.
adoral succession (figs. Land1la@). (B.M.38742.) ©) Or |
The number of these primaries 1s consider- \%O 0

able; and they almost reach the ambitus in
specimens which are not adult. They are followed by, or alter-
nate with, compound plates composed of a primary and a low,

short demi-plate (fig. 2), or a demi-plate Fig. 2.

may be intercalated between two primaries, EL OLOTO
the three not forming a compound plate O O
(fig. 2a). (B.M. 723.) The demi-plate in eoN O

this instance was a primary which has been

crowded-out by the growth-pressure of the
primary above and below; but in the other
instance (fig. 2) the demi-plate has been so
pressed upon that it has been fused, as it

were, with the primary.

Both in the specimen marked 723 in the British Museum and
in one in our possession the primaries are followed, at the ambitus
or just above it, by taller compound plates, each of which consists
of a large primary placed abactinally, and a low,
broad, triangular demi-plate situated actinaily.
The two plates are united by very delicate
sutures and form a geometrical compound plate
(fig. 3). (See also B.M. 180.)

Sometimes, at the ambitus, there is a third
plate in a compound plate; and the arrange-
ment seen is very unusual in the Echinide. For :—(1) The pairs
of pores are very slightly out of the straight vertical line. (2)
The upper plate is a large primary which occupies the whole of

very narrow, and have one pair of small round holes opposite each ambulacral
plate.” The pores are of course in ambulacral plates. On the same page it is
stated, “near the mouth-opening two rows [of pairs of pores] go to each plate.”
This is incorrect ; for there is but one pair to a plate.

52 PROF. P. M. DUNCAN AND MR. W. P. SLADEN ON THE

the compound plate at the median suture of the ambulacrum
(fig. 4). (B.M. 180.) (8) There are two Fig. 4.
demi-plates; the upper one has its aboral
suture passing actinally so as to reach the
adoral suture of the compound plate at a
little distance from the median suture ;
whilst the lower demi-plate has its aboral
suture also sloping actinally and reaching
the common adoral suture not far from the interradial edge.

The position of these demi-plates is exactly opposite to that
noticed in the compound plates of the Triplechinide*.

In nearly all the specimens the size of the pores diminishes
actinally, but this is not invariable; and in all cases the pairs
become closer, and alternate pairs are crowded out of the straight
vertical line.. There is no intercalation of pairs—that is, no new
growths amongst the original pairs. Actinally, and on the slope
of the raised ambulacra, which have already been noticed, the
pairs of pores are crowded and biserial in their arrangement,
and the pores are placed so obliquely that the aboral one is
nearly vertical to the adoral. Although the crowding of the
pairs is often excessive, still there is never more than a pair to
a plate; and the plate, often very small, forms part of a com-
pound plate. Solitary primary or demi-plates which do not enter
into the composition of compound plates are not found below the
ambitus. The compound plates there consist of a primary with
one or two demi-plates.

The following is a description of four compound plates placed
near the ambitus and actinally (fig. 5). (B.M. 38742.) Plate],
the abactinal one of the series, consists of
a large middle primary and a small aboral
demi-plate and a larger adoral demi-
plate. The aboral demi-plate is perfo-
rated by a pair of the outer set of pores
of the biserial zone, and the primary
by a pair of the inner set, whilst the
adoral demi exhibits a pair of pores
similar to those of the aboral demi-
plate. But the primary occupies the
whole of the median suture, and the
demi-plates do not come in contact ; for the adoral suture of the

* Lovén, ‘Etudes,’ pl. xvii.

ANATOMY OF THE TEST OF DISCOIDEA CYLINDRICA. 53

upper one curves aborally to reach the aboral suture of the com-
pound plate, and the adoral demi-plate has its aboral suture curved
so as to reach the adoral suture of the compound plate.

The next compound plate (2) is formed in the same manner as
that just described ; but the demi-plates are smaller, and are per-
forated by pairs of pores which belong to the inner set; on the
other hand, the primary placed between the demi-plates has a
pair of pores of the outer set, and this is the reverse of what
occurs in the plate above. Plate 3 has only two components,
and the abactinal plate is a low broad demi-plate perforated by a
pair of pores of the outer set; the actinal plate is a low primary,
and its pair of pores belongs to the inner set.

Plate 4 is formed like plate 2, but the demi-plates are larger ;
moreover, the pairs of pores of the upper demi-plate are of the
inner set, like the pair seen in the demi-plate placed immediately
abactinally in the compound plate above; but they are placed
nearer the interradium than the pair above, and the result is to
give a triple series of pairs of pores with the pair of the primary
of plate 4. This triple appearance is rare in Discoidea. It will
be observed that the composition of the four compound plates,
omitting the biserial condition, is suggestive of that seen in the
genus Calopleurus*.

At some little distance from the peristome the alternation of
outer and inner pairs of pores continues, and the compound
plates are rarely formed of more than two plates. The primary
of the compound plate is low and broad, and is placed abactinally,
whilst the very small demi-plate is found at the actinal and
outer part of the plate. When a tubercle
is present, the primary becomes high. The
primary has a pair of the outer series of pores,
and the demi-plate a pair of inner series
(fig. 6).

In half-grown specimens a decided change
occurs in the pairs of pores at a little distance from the peri-
stome, and they become larger, wider apart, and the arrangement
is nearly in simple series (fig. 7). The pores of each
pair are very oblique and almost vertical; and the
greater number belong to primaries which are higher
than those nearer the ambitus. A few are in demi-
plates. In older specimens the simple series close
to the peristome is much shorter, and indeed barely

Fig. 6.

Fig. 7.

* Duncan and Sladen, Journ. Linn. Soc., Zool. vol. xix. 1885, pl. i.

54 PROF. P. M. DUNCAN AND MR. W. P. SLADEN ON THE

exists, the pairs not being so large comparatively as in the
younger forms, and with their arrangement biserial.

In both instances the expansion of a tubercle-bearing plate
increases its height ; and it is followed by small low primaries or
by a demi-plate. The adult specimens conform very much to the
drawing given by Lovén of the peristome of Discordea conica
(‘ Etudes,’ pl. xiv. fig. 125).

The peristomial end of the poriferous zones is contracted, and
a series of tubercles separates the pairs from the position of the
interradial suture; so that there were spines in rows up the
narrow space of the peristomial funnel-shaped tube. The first
pairs of pores are not visible from without, for they are high up
in the peristomial tube, and are placed in the processes of a
structure which is termed the perignathic girdle (Journ. Linn.
Soc., Zool. vol. xix. p. 179.)

In a specimen attributed to Discoidea cylindrica in the
British Museum (No. E 180), which is balf-grown, and has not
become cylindrical in outline above the ambitus, but is simply
hemispherical, the arrangement of the plates above the ambitus
is somewhat exceptional. ‘The pores are large and oblique, being
in simple series and rather distant. The first four plates of a
certain set are low and broad; then comes a tubercle-bearing
primary with a considerable downward expansion towards the
median line. It is followed by a low primary which is almost a
demi-plate on account of the diminution of its vertical measure-
ment near the median line. This loss of size has been due to
the growth of the primary above, and the pressure has made
both plates to combine within a geometrical outline to form a
compound plate. The next plate is a decidedly very low primary,
and it is succeeded by a primary with a downward expansion
forming a compound plate with a low primary, which is almost
a demi-plate. This compound plate is followed by a low
primary. Just above the margin in this specimen, the growth-
pressure has altered the shape of the plates in a very instructive
manner. There is a triplet, and the first plate of it is a large
downwardly expanded primary; the next is a low and broad
demi-plate, for the pressure has blocked out part of a low
primary near the median line, and a demi has resulted. The
third plate is a small narrow and low demi-plate; and this
was once a very low primary resembling those further up.
Pressure made it assume the shape of a common small demi,

——~——

ANATOMY OF THE TEST OF DISCOIDEA CYLINDRICA. 55

and then the three plates were joined into a triple compound one.
It appears that in some specimens the outer set of pairs of
pores on the flank of the raised ambulacra are larger than the
inner series; and when this is the case, the alternations of the
pairs are very well seen.

The width of the ambulacra, compared with that of the inter-
radia, is from one fourth to one third; and above the ambitus
there are nine pairs of pores opposed to two interradial plates*.

The Interradial Plates.—These gradually diminish in size
towards the peristome, and still more so as they pass up the
funnel of the peristome to the perignathic girdle. At the girdle
the odd interradium (No. 5) has a single piate; and although
we are not quite satisfied on the point, it appears that there are
single plates at the peristomial margin of the interradia Nos. 1
and 8. Double plates are seen at the margin of the peristome
in interradia Nos. 2 and 4.

This is the arrangement of plates noticed in the funnel-shaped
peristomes of Hchinoneus and Amblypygus +. Loven has figured
a solitary plate in the peristome of Discovdea conica in inter-
radium 5 f.

The Peristome.—The actinal aperture is very small in relation
to the dimensions of the test; it leads up through a sort of funnel-
shaped tube to the peristomial margin, which is therefore well
within the test, and not flush with its actinal surface. The peri-
stomial tube becomes narrower upwards, and its upper margin
merges into a remarkable perignathic girdle. The openiug of the
peristome actinally is very nearly circular, and in no instance is
it worthy of the term decagonal ; there is the slightest departure
only from a continuous curvature. But when the upper margin
of the peristome is seen from below, the outline departs more
from that of a circle, and ten slight curves are seen—tive, broad
and decided, are interradial, and five, narrower and less formed,
are ambulacral. There are no notches actinally; but at the
margin of the peristome and at the edge of the perignathic
girdle there are faint groove-like notches between the interradia
and the ambulacra.

* Jn describing the ambulacra, the late Dr. Wright stated that about three
of its plates correspond vertically to one interambulacral plate (op. cit. p. 206) ;
but in the plate drawn by Bone, pl. xlvi. fig. 1 d, nine pairs of pores correspond
to two interambulacral plates ; and that is correct.

t Duncan and Sladen, Pal. Ind., ser. xiv., Tert. Echin. of Kachh, p. 17,
{ Lovén, ‘ Etudes,’ pl. xiv. fig. 125.

56 PROF. P. M. DUNCAN AND MR. W. P. SLADEN ON THE

The curved edges of the inner extremity of the peristomial tube,
as seen from below upwards, are not the terminations of the
ambulacra and interradia; for when a section is made transversely
through a test at the ambitus, and the chalk is cleared out from
the actinal portion of the divided mass, plates are seen forming
an internal oblique projection in the position of the perignathic
girdle.

The projection is continuous around and above the peristomial
tube, and is a perignathic girdle of a very remarkable and unique
kind. The girdle is well seen in a specimen at the British Museum
No. 40841 (fig. 8); and it will be observed that the so-called

Fig. 8.

A
vs

oe ae
Voge
Pitee. o>

ribs, ten in number, which pass over the inner surface of the
actinal interradial plates, commence at the outer edge of the
interradial parts of the girdle. The girdle is rather low, and sur-
rounds the peristomial opening in the form of a raised, oblique,
broad, ridge-like ring. The upper surface of the girdle is free,
and consists of flat or slightly irregular slanting surfaces, the
slant being towards the peristome, ending all round and out-
wards in a continuous and wavy free edge. This edge has the
parts which correspond with the ambulacra thin, less projecting
than the other portions, and re-enteringly curved. ‘The parts

ANATOMY OF THE TEST OF DISCOIDEA CYLINDRICA. 57

of the edge of the girdle which correspond with the interradia
are boldly curyed outwards and are larger than the others.

The outer wavy free edge of the whole girdle overhangs the
inner surface of the base of the test, and the ribs just men-
tioned arise from the underpart of the overhanging structure.
The upper surface of the girdle is of course between the edge
just alluded to and the peristome ; and its slanting surface is not
a simple plane one, for on each of the broader interradial portions
there is a slight depression on either side of the centre and an
elevation close to the ambulacral curves. This depression is
probably the relic of a muscular origin, one on each side of the
median line of a plate. No sutures occur in the interradial parts
of the girdle; but it is not satisfactorily shown that there are not
sutures between the ambulacral and interradial portions along
the line of the slight groovings which are on either side of an
ambulacrum high up in the peristome, and at the lower edge of
the inner surface of the girdle—that is to say, in the usual
position of sutures in relation with branchial grooves or cuts*.
There is a distinct median and more or less vertical suture in
every ambulacral part of the girdle, and there are pairs of pores
on the sides of it (fig. 8).

In ambulacrum IIT., zone “db,” there are two pairs of pores
which are placed obliquely, as is the case with the single pairs of
the other ambulacra, and the pairs of the other ambulacra are at
different distances from the free edge of the ambulacral parts of
the girdle; and these different positions are exactly those which
occur in the particular plates around the peristomes of other
regular Echinoidea according to Lovén. The lower part of the
girdle is continuous with the ambulacral and interradial plates of
the actinal part of the test around the peristome. The inner
part of the girdle is the upper continuation of the peristomial
tube.

The anatomy of the perignathic girdle in the Echinoidea was
described by one of us in this Journal (Journ. Linn. Soc., Zool.
vol. xix. p. 179, 1885); and that of Discotdea cylindrica was noticed
as follows (p. 182) :—‘‘ In Discotdea there is a continuous girdle
without arches, although the homologues of the processes exist.”
Reference must be made to the above-mentioned communi-
cation in order to comprehend the terminology. It is evident
that there are no arched parts in Discotdea cylindrica; but it is

* Duncan, Journ. Linn. Soc., Zool. 1885, pl. xxx. fig. 9, letter s,
LINN. JOURN.—ZOOLOGY, VOL. XX. 5

58 PROF. P. M. DUNCAN AND MR. W. P. SLADEN ON THE

true that the ambulacral parts of the girdle are perforated by one
or more pairs of tentacular pores; and therefore the parts thus
penetrated are the homologues of the processes which in other
Gnathostomes (omitting the Cidaride) form the sides to or
processes of the arches (“auricles ” of authors).

The interradial portions of the girdle in Discotdea cylindrica
are the homologues of the ridges of the other regular Hchinoidea,
including the Cidaride, and, as in the other forms, the ridges
are composed of one or more interradial plates.

On comparing the girdles of Cidaris and Discoidea, it will
be noticed that both have the ridges well developed, and clearly
in consequence of the attachment of protractor muscles. In
Cidaride the utility of the retractor muscles is diminished by
the nature of the scaly structure around the peristomial opening ;
and in Discoidea the extremely high and narrow peristomial tube
and the spines attached to tubercles within indicate that wide
opening of the jaw-ends could not occur even as much as in
Cidaris.

Neither in Cidaris nor in Discoidea are the “ processes’’ of the
ambulacral parts of the girdle developed so as to afford origin or
attachment to muscles ; and it is evident that the portions of the
ambulacra in Discoidea which are perforated, and which are the
homologues of the processes of the ambulacral arches of the
Echinide, could not have given attachment to muscles because of
the presence of the pores; moreover, the processes of Hchinide
are not the origins of muscles, but the archesare. The distinction
in structure between the girdles of Discozdea and of the Echinide,
such as the species of the genera Hchinus, Salmacis, Temnopleurus,
Diadema, Celopleurus, Strongylocentrotus, &c., is evident; for
there are no arches and no upward prolongations or processes
to form arches in Discoidea. The greatest affinity in structure
is with the Cidaride; but the difference is important, inasmuch
as the ambulacral parts of the girdle are wanting in Cidarids,
and are high and block the ambulacral path in Discoidea.

No trace of pyramids or of any portions of a dental apparatus
has ever been found in Discoidea ; but it is impossible to reject
the supposition that the genus was gnathostomous, for the pre-
sence of ridges marked with depressions in the girdle would seem
to point to muscular attachments and to the inevitable presence
of jaws. The position of the jaws was probably not so vertical as
in the Cidaride, but slanting more or less; and it is probable

ANATOMY OF THE TEST OF DISCOIDEA CYLINDRICA. 59

that the teeth did not project, but moved as in the Clypeastroids.
It is possible that the dental apparatus was slender, as it is in
Holectypus.

The “Internal Ribs.’’—These ten ridge-like projections on the
interradia within the test are low and narrow, but wider at their
bases than at the free edge. They reach from the outer and
under part of the perignathic girdle to the inner surface of the
test just above the ambitus, and their height is about a millimetre
near the girdle, and 3 millimetres at the opposite extremity. .

There are two ribs to an interradial area, one on each side of
the median line of suture of the interradium ; and each one is
placed along the middle of each row of interradial plates.

The ribs are not additional plates, but simply special upward
growths of the middle portions of the upper surfaces of the inter-
radial plates of the actinal part of the test. The sutures which
are between consecutive interradial plates can be traced over
the ribs. Finally, it must be noticed with regard to the growths
of the inside of the test, that there is a low ridge along each
median suture of the five ambulacra on the inside of the base
of the test; it commences at the ambulacral part of the
girdle, and gradually diminishes towards the ambitus of the test
within *.

The Periproct.—This is small in relation to the size of the test,
and is in a very slight concavity in the odd interradium, being
nearer to the margin of the test than to the peristome. It is
elongated in the direction of the antero-posterior axis of the test,
and varies in shape from the elliptical with rather narrow ends tc
the ovoid witha considerable curvature posteriorly. It is bounded
by four plates in one and five plates in the opposite zone ; aid the
sixth plate from the peristomial margin, not including the first
single plate, forms the inner boundary in one zone, and the seventh
plate in the other zone.

The surface of the test, as has been remarked by nearly every
observer, is remarkably punctate ; and the multitudes of minute
depressions have a corresponding number of small, close. sharp
granules between them. The larger tubercles are in sunken
serobicules which are shallow; and there is decided crenulation
and. perforation.

* Zittel, Handb. d. Pal. Bd. i. 1876-80, p. 514, fig. 873. The ten ribs are
shown and part of the perignathic girdle.

60 PROF. P. M. DUNCAN AND MR. W. P. SLADEN ON THE

The Fifth Basal Plate-—The specimens of Discoidea cylindrica
in the British Museum show that the fifth basal is not a comple-
mentary plate, but a true basal which is not perforated for a
genital duct, but which is penetrated by the madreporite. The
position and dimensions of the plate are those of a normal basal
in other forms; and it is not comparable with the so-called fifth
plate described by Cotteau in one specimen of Hehinoconus albo-
galerus, for that was a part of the left posterior basal. Hcehino-
conus has no fifth basal.

The Evolution of the Fifth Genital Duct and the Perforation of
the Fifth Basal Plate of Species of Discoidea—Lovén has argued
that during the lapse of time the generative organs of species of
Discoidea became more fully developed after the anus moved out
of the apex, and that the fifth plate reappeared and became per-
forated by a genital duct*. There is much to be advanced in
favour of this remarkable generalization, and it is certainly the
case that the oldest species had four basals perforated by genital
ducts, whilst the youngest had five perforated basals. The
oldest species do not, however, obtain a fifth genital duct, and its
perforation during lapse of ages does not occur; for the oldest
and youngest forms of Discoidea cylindrica, for instance, have
only four perforated basals. D. conica, Desor, is a Gault and
Albien species of Europe and Africa, and it follows Lovén’s law,
and has but four basals perforated by the duct, and the fifth is
imperforate. D. subuculus ranges from the Warminster Upper
Greensand into the Lower Chalk ; and it has been described as
having only four or sometimes five basals perforated; there
are two specimens in the British Museum in which all the basals
are perforated.

Discoidea minima, D. Favrina, D. Jullient, and D. Forgemolli
are European and North-African forms, and all have five basals
perforated ; and the age of the fossils is Cenomanien. D. infera
and D. Dizxoni are from the Upper Chalk, and all the five basals
are perforated.

So far as the whole genus is concerned, the generalization of
the appearance of the fifth basal perforation in the later ages of
its lifetime is proved; but the appearance of a fifth per-
forated basal in time has not been proved to occur in the same

* Lovén, On Pourtalesia, Kong]. Svenska Vetenskaps-Akademiens Handlingar,
Bd. xix. No. 7, 1883, p. 68.

ANATOMY OF THE TEST OF DISCOIDEA CYLINDRICA. 61

species. The validity of this interesting observation by Lovén
will have to be tested on other grounds; for it is a matter of
considerable doubt in our minds whether all the species which
have been admitted into the genus Discoidea can remain in it.

D. subuculus differs much from D. cylindrica in the hollowed-out
and tumid nature of the actinal part of the test, in the existence
of low primary ambulacral plates only, and their great crowding
without the formation of compound plates. Again, the madre-
porite is in the second basal only. Nevertheless, we have found
ribs on the inner surface of the actinal part of the test, as in
D. cylindrica; and probably the perignathic girdle will be found.
As yet, we have only detected very indefinite traces of it.

On the Characters of the Genus Lophopus, with Description of a
new Species from Australia. By Stuart O. Riptey, M.A.,
ELS.

[Read 4th November, 1886. ]
(Prats IT.)

ProBaBLy in few groups of the Animal Kingdom have such
unnatural characters been employed for the distinction of genera
and species asin the Phylactolematous Polyzoa. Fewsystematic
zoologists can, for example, have studied the relations of Alcy-
onella and Plumatella without feeling that the current reasons for
separating these two divisions are far from satisfactory, consist-
ing, as expressed by Prof. Allman in his well-known ‘ Monograph
of the Freshwater Polyzoa’ (Ray Society, 1856), chiefly in the
manner of connection between the tubes composing the colony.
*‘ Except in the condition of the dermal system, the structure of
Plumatella differs in no essential point from that of Alcyonella,
This system, however, in the coalescence of the tubes into a
common mass in Alcyonella, while they remain totally distinct in
Plumatella, presents us with a difference which I believe to be
of sufficient importance to justify us in placing the two forms in
separate generic groups ” (/.c, p. 92).

Dr. Jullien (“ Monographie des Bryozoaires d’eau douce,” Bull.
Soc. Zool, France, x. p. 90, published in 1885) has given very
forcible expression to this feeling of dissatisfaction, and has
indeed introduced into the classification modifications of a very

LINN. JOURN.— ZOOLOGY, VOL. XX. 6

62 MR. S. O. RIDLEY ON THE

fundamental character, of which his union of the genus Alcy-
onella with Plumatella, and even the species of the former with
species of the latter genus in a long list of synonyms, is not the
most radical of the changes.

It is, however, my object on this occasion to draw attention to
but one part of the existing systems which seems to require
remodelling.

The diagnosis of Lophopus, as given by Allman (J. c. p. 83),
runs :—“ Coenecium sacciform, hyaline, with a dise which serves
for attachment but not for locomotion; ectocyst gelatinoid ;
orifices scattered. Statoblasts elliptical, with an annulus, but
without marginal spines.”

Jullien (/.¢. p. 189), besides characters derived from the
general relations of the colony as a whole, inserts in his defini-
tion of the genus :—“ Statoblastes elliptiques pourvus d’un anneau
sur le pourtour; anneau terminé en pointe aux extrémités du
grand diametre.”’

The discovery, as detailed below, of a new species of the genus
in Australia, involves the giving up of the last-named character
as of generic importance, and Lophopus remains distinguished
from Cristatella by the absence of spines from the statoblast
and of a locomotor disk from the colony, and from Pectinatella
chiefly by the former character.

Loryoprus LENDENFELDI, 0. sp.

Zoarium forming chain-like or contort linear growths, the
zocecia being aggregated into small linear groups, which are con-
nected with each other by stolon-like lobes. Zooids deeply
divided from each other within the ectocyst, tubular, ending
below in rounded, slightly constricted knobs terminated by a
special thickening of the outermost layer of the endocyst.
Kctocyst perfectly hyaline, about 1 millim. deep, forming low mam-
mille, about ‘02 millim. high, which contain the zocecial orifices.

Tentacles about 50 in number, the longest longer than the
endocyst-body. Hpistome not observed. Ectocyst (in spirit)
hyaline, colourless; endocyst-bodies pale yellow. Statoblast
strictly elliptical in outline, annulus almost flat; body very
convex.

Measurements.—Length of extended polypide (the portion
within ectocyst-sheath) about 1 millim. Diameter of neck of
zocecium (endocyst) about ‘02 millim. Length of lophophore, from

CHARACTERS OF THE GENUS LOPHOPUS. 63

root of arm to tip of furthest tentacle, about 1 millim. Maximum
length of tentacles (in spirit specimens) about 1 millim.

Statoblast about °85-95 millim. long by °7 millim. broad;
annulus about :22 wide at ends, about ‘15 millim, at sides; cells
of annulus about ‘01 to ‘05 millim. broad.

Habitat. Paramatta River, New South Wales.

The chief points distinguishing this species from L. erystallinus
appear to be the absence of terminal angles to the statoblast, and
the knobbed form of the inner end of the endocyst. The ten-
tacles are also probably far longer than in JL. erystallinus. No
other species of the genus is known—that form assigned to it by
Mr. Carter, from Bombay, evidently being rightly removed from
Lophopus by Hyatt and Jullien. I have bestowed upon the
species under notice the name of its indefatigable discoverer,
Dr. von Lendenfeld, not only on account of his discovery, and
of the very remarkable manner in which he has preserved the
specimen, which shows almost all its characters as in life, but also
in commemoration of his generosity in presenting it to the
Natural History Museum, and in allowing me to describe a new
form, systematically and geographically of such high interest.

The particular specimen from which this description is drawn
up coats some thin fasciated vegetable stems for a length of
84 inches, but probably was much more extensive originally ;
the mass thus formed is about 7 millim. in greatest diameter.
The statoblasts are described from a specimen found in a dead
colony, obtained at the same spot, which exhibits the characters
of the ectocyst sufficiently for identification. The conditions
under which the colonies are growing (practically covering the
entire cireumference of the object in which they rest) almost
preclude the possibility of such approaches to locomotion as have
been sometimes attributed to L. crystallinus ; at the same time
their attachment to the base is very slight. The body of the
statoblast is dark umber-brown, the annulus colourless.

_ Minute Structwre.—Staining with borax carmine has enabled
me, in the excellent state of preservation of the specimen, to
demonstrate clearly that the outermost layer of the ectocyst
consists of substellate nucleated cells; these are most usually
roughly oblong in shape, with the branches chiefly at the poles,
and measure about ‘015-02 by ‘005 millim., with a small nucleus
and punctiform nucleolus; but interspersed abundantly among
them are some cells with a globular central mass about ‘01 millim.

64 ON THE CHARACTERS OF THE GENUS LOPHOPUS.

wide, mainly composed of a large nucleus, which contains a
nucleolus, itself about ‘002 millim. broad.

These cells are not described by Allman (Joc. ett.) or in Hyatt’s
papers (Proc. Essex Institute, vols. iv. & vi.), nor have I seen them
noticed in any other paper which I have met with. Their form
would seem to assign a mesodermic origin to them, but hitherto
no overlying tissue has been found, to represent the ectoderm.

Floating Apparatus of the Statoblast.—The cellular structure
of the annulus of the statoblast in those forms in which it is
developed resembles that of the gemmule of true Spongillide,
as elucidated by the researches of Carter, Marshall, and Vej-
dowsky, so strongly that it only requires to be pointed out in
order to be recognized.

Australasian Species of Freshwater Polyzoa.

The only Australasian species of Freshwater Polyzoa recorded
by Jullien (/.¢.), and by Hutton, in his ‘ Catalogue of the Marine
Mollusca of New Zealand,’ is Plumatella Aplini of Macgillivray.
A form assigned to Plumatella repens, var. a of Allman, is, however,
described by A. Hamilton from near Napier in New Zealand
(Trans. New Zeal. Inst. xu. p. 302) ; and Mr. Whitelegge is stated
to have exhibited the same species, Plumatella repens, and Frede-
ricella sultana, from New South Wales, in Proc. Linn. Soe. N. S.
Wales, viii. (18828) pp. 297, 416. The genus Zophopus does not
appear to have been hitherto recorded from Australasia.

DESCRIPTION OF PLATH II.

Fig. 1. Lophopus Lendenfeldi, n. sp. Part of the colony, attached to plant-
stem, showing a subspiral arrangement of the zoarium. Natural size.
2. Part of the zoarium, showing zooids in different states of contraction
or expansion. a. Hyaline (? cuticular) layer observed at termination
of body of polypides. 6. Food-mass in alimentary canal. x40 diam.
3. Portion of the ectocyst, to show structure as exhibited by staining with
borax carmine. a. Stellate cells; b. Globose cells. 600 diam.
4, A statoblast, showing: a, annulus; 0, body; ¢, blastodermic cells*?
x 60 diam.
N.B. These figures are somewhat schematized, except as regards essential
details.

* These bodies appear to consist mainly of refractive granules and of a
large nucleus. Perhaps they represent an early stage in the division of the
germ-cell,

Linn Soc. Journ ZOOL. Vor. XX Pie.

Bl nal svze

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ON THE SENSE OF SMELL IN DOGS. 65

Experiments on the Sense of Smell in Dogs.
By George J. Romaness, LL.D., F.R.S., F.LS.

[Read 16th December, 1886. |

Or all the phenomena presented by the higher evolution of
sense-organs in the Animal Kingdom, to my mind the most
remarkable is the acuteness of olfactory perception which is
exhibited by certain orders of Mammalia. All the other faculties
of special sense are, so to speak, more evenly distributed through-
out the vertebrated series; so that when we compare our own
sense of sight, of hearing, or of taste, with those of vertebrated
animals in general, we at once recognize that they are comparable.
But such is not the case with the sense of smell; for in many of the
Carnivora, Ruminants, &c., this sense has undergone so enormous
a development as to be suggestive of differing from our own, not
merely in degree, but in kind. Any one, for example, who is ac-
customed to deer-stalking must often have been freshly astonished
at the precautions which it is needful to take in order to prevent
the game from getting wind of the sportsman. Indeed, to a
novice such precautions are apt to be regarded as implying a
superstitious exaggeration of the possibilities of olfactory per-
ception ; and it is not until he has himself seen the deer scent
him at some almost incredible distance that he lends himself
without disguised contempt to the direction of the keeper. Yet
among the Carnivora the sense of smell is even more extraordi-
nary. Here, for instance, is an observation upon the subject
which I published several years ago, and which I now quote be-
cause it led to the experiments which it is the object of this
paper to detail :—

“T once tried an experiment with a terrier of my own which
shows, better than any thing that I have ever read, the almost
supernatural capabilities of smell in Dogs. On a Bank
holiday, when the broad walk in Regent’s Park was swarming
with people of all kinds, walking in all directions, 1 took my
terrier (which I knew had a splendid nose, and could track me
for miles) along the walk, and, when his attention was diverted
by a strange dog, I suddenly made a number of zigzags across
the broad walk, then stood on a seat, and watched the terrier.
Finding I had not continued in the direction I was going when
he left me, he went to the place where he had last seen me, and
there, picking up my scent, tracked my footsteps over all the

66 MR. G. J. ROMANES—EXPERIMENTS ON

zigzags I had made until he found me. Now in order to do this _
he had to distinguish my trail from at least a hundred others
quite as fresh, and many thousands of others not so fresh, cross-
ing it at all angles.” *

The object of the experiments about to be described was that
of ascertaining whether a dog, when thus distinguishing his
master’s trail, is guided by some distinctive smell attaching to
his master’s shoes, to any distinctive smell of his master’s feet,
or to both these differences combined.

I have a setter-bitch over which I have shot for eight years.
Having a very good nose, she can track me over immense dis-
tances, and her devotion to me being very exclusive, she consti-
tuted an admirable subject for my experiments.

These consisted in allowing the bitch to be taken out of the
kennel by some one to whom she was indifferent, who then led
her to a prearranged spot from which the tracking was to begin.
Of course this spot was always to leeward of the kennel, and the
person who was to be tracked always walked so as to keep more
or less to leeward of the starting-point. The district—park-lands
surrounding a house—was an open one, presenting, however,
numerous trees, shrubberies, walls, &c., behind which I could hide
at a distance from the starting-point, and so observe the animal
during the whole course of each experiment. Sundry other pre-
cautions, which I need not wait to mention, were taken in order
to ensure that the bitch should have tc depend on her sense
of smell alone, and the following are the experiments which were
tried :—

1.—I walked the grass-lands for about a mile in my ordinary
shooting-boots. The instant she came to the starting-point, the
bitch broke away at her full speed, and, faithfully following my
track, overtook me in a few minutes.

2.—I set a man who was a stranger about the place to walk
the park. Although repeatedly put upon his trail by my servant,
the bitch showed no disposition to follow it.

-3.—I had the bitch taken into the gun-room, where she saw
me ready to start for shooting. I then left the gun-room and
went to another part of the house, while my gamekeeper left the
house by the back door, walked a certain distance to leeward in
the direction of some partridge-ground, and then concealed him-

* ‘Mental Evolution in Animals,’ pp. 92-8; where also see for additional
remarks of a general kind on the sense of smell in different animals,

THE SENSE OF SMELL IN DOGS. 67

self. The bitch, who was now howling to follow me, was led to
_ the back door by another servant. Quickly finding the trail of
the gamekeeper, she tracked it for a few yards ; but, finding that
I had not been with him, she left his trail, and hunted about in
all directions for mine, which, of course, was nowhere to be
found.

4.—I collected all the men about the place,and directed them
to walk close behind one another in Indian file, each man taking
care to place his feet in the footprints of his predecessor.
In this procession, numbering twelve in all, I took the lead,
while the gamekeeper brought up therear. When we had walked
two hundred yards, I turned to the right, followed by five of
the men; and at the point where I had turned to the right, the
seventh man turned to the left, followed by all the remainder.
The two parties thus formed, after having walked in opposite
directions for a considerable distance, concealed themselves, and
the bitch was put upon the common track of the whole party
before the point of divergence. Following this common track
with rapidity, she at first overshot the point of divergence ; but,
quickly recovering it, without any hesitation chose the track
which turned to the right. Yet in this case my footprints in
the common track were overlaid by eleven others, and in the
track to the right by five others. Moreover, as it was the
gamekeeper who brought up the rear, and as in the absence

_ of my trail she would always follow his, the fact of his scent

being, so to speak, uppermost in the series, was shown in no
way to disconcert the animal when following another familiar
scent lowermost in the series.

5.—I requested the stranger before-mentioned to wear my
shooting-boots, and in them to walk the park to leeward of the
kennel. When the bitch was led to this trail she followed
it with the eagerness wherewith she always followed mine.

6.—I wore this stranger’s boots, and walked the park as he
had done. On being taken to this trail, the bitch could not be
induced to follow it.

-7.—-The stranger walked the park in bare feet; the bitch would
~ not follow the trail.

8.—i walked the park in bare feet: the bitch followed my
trail; but in quite a different manner from that which she
displayed when following the trail of my shvoting-boots. She
_ was so much less eager, and therefore so much less rapid, that

68 MR. G. J. ROMANES—EXPERIMENTS ON

her manner was suggestive of great uncertainty whether or not .
she was on my track.

9.—I walked the park in new shooting-boots, which had never
been worn by any one. The bitch wholly refused to take this
trail.

10.—I walked the park in my old shooting-boots, but having
one layer of brown paper glued to their soles and sides. The
bitch was led along my track, but paid no attention to it till she
came to a place where, as I had previously observed, a small
portion of the brown paper first became worn away at one of my
heels. Here she immediately recognized my trail, and speedily
followed it up, although the surface of shoe-leather which
touched the ground was not more than a few square milli-
metres.

11.—I walked in my stocking-soles, trying first with new
cotton socks. The bitch lazily followed the trail a short distance
and then gave it up. I next tried woollen socks which I had
worn all day, but the result was the same, and therefore quite
different from that yielded by my shooting-boots, while more
resembling that which was yielded by my bare feet.

12.—I began to walk in my ordinary shooting-boots, and when
I had gone fifty yards, I kicked them off and carried them with
me, while I continued to walk another three hundred yards in
my stocking-soles; then I took off my stockings, and walked
another three hundred yards on my bare feet. On _ being
taken to the beginning of this trail, or where I had started
m my shooting-boots, the bitch as usual set off upon it at
full speed, nor did she abate this speed throughout the whole
distance. In other words, having been once started upon the
familiar scent of my shooting-boots, she seemed to entertain no
doubt that the scent of the stocking-soles and of the bare
feet belonged to me; although she did not clearly recognize them
as belonging to me when they were not continuations of a track
made by my shooting-boots (10 and 11).

-138.—I requested a gentleman who was calling at the house,
and whom the bitch had never before seen, to accompany me in
a conveyance along one of the carriage-drives. Ata distance of
several hundred yards from the house, I alighted in my shooting-
boots, walked fifty yards beside the carriage, again entered it
while my friend alighted and walked two hundred yards still
further along the drive. The bitch ran the whole 250 yards at

THE SENSE OF SMELL IN DOGS. 69

her full speed, without making any pause at the place where the
scent changed. This experiment was subsequently repeated with
other strangers, and with the same result.

14.—I walked in my ordinary shooting-boots, having previously
soaked them in oil of aniseed. Although the odour of the ani-
seed was so strong that an hour afterwards the path which I had
followed was correctly traced by ‘a friend, this odour did not
appear to disconcert the bitch in following my trail, for she ran
me down as quickly as usual. It was noticed, however, by the
friend who took her to the trail that she did not set off upon it
as instantaneously as usual. She began by examining the first
three or four footsteps with care, and only then started off at
full speed.

15.—Lastly, I tried some experiments on the power which this
bitch might display of recognizing my individual odour as ema-
nating from my whole person. In a large potato-field behind
the house, a number of labourers had been engaged for eight or
ten hours in digging up and carrying away potatoes all the way
along half a dozen adjacent “drills.” Consequently, there was
here a strip of bared land in the field about twenty yards wide,
and a quarter of a mile long, which had been thoroughly well
trampled over by many strange feet. Down this strip of land
I walked in a zigzag course from end to end. On reaching the
bottom I turned out of the field, and again walked up a part of
the way towards the house, but on the other side of a stone wall
which bounded the field. This stone wall was breast high, and
was situated nearly a hundred yards to windward of my previous
course through the potatoes. The bitch, on being led out of the
house, was put upon my trail at the top of the field, and at high
speed picked out my trail among all the others, following roughly
the various zigzags which I had taken. But the moment she
gained the “‘ wind’s-eye’’ of the place where I was standing be-
hind the wall, she turned abruptly at a right angle, threw up her
head, and came as straight as an arrow to the spot where I was
watching her. Yet while watching her I had allowed only my
eyes to come above the wall, so that she proved herself able
to distinguish instantly the odour of the top of my head (without
hat) at a distance of two hundred yards, although at the time
she was surrounded by a number of over-heated labourers.

16.—On another day, when it was perfectly calm, I tried the
experiment of standing in a deep dry ditch, with only the top of

70 ON THE SENSE OF SMELL IN DOGS.

my uncovered head above the level of the surrounding fields.
When she was led within two hundred yards of the place, she
instantly perceived my odour, and ran in a straight line to where 1
had then ducked my head, so that she should receive no assistance
from her sense of sight. This experiment shows that, in the
absence of wind, the odour of my head (and no doubt, in a lesser
degree, that of my body) had diffused itself through the air in all
directions, and in an amount sufficient to enable the setter to
recognize it as my odour at a distance of two hundred yards.

From the above experiments I conclude that this bitch distin-
euishes my trail from that of all others by the peculiar smell of my
boots (1 to 6), and not by the peculiar smell of my feet (8 to 11).
No doubt the smell which she recognizes as belonging distinctively
to my trail is communicated to the boots by the exudations from
my feet ; but these exudations require to be combined with shoe-
leather before they are recognized by her. Probably, however,
if I had always been accustomed to shoot without boots or
stockings, she would have learnt to associate with mea trail made
by my bare feet. The experiments further show that although a
few square millimetres of the surface of one boot is amply
sufficient to make a trail which the animal can recognize as mine,
the scent is not able to penetrate a single layer of brown paper
(10). Furthermore, it would appear that in following a trail
this bitch is ready at any moment to be guided by inference as
well as perception, and that the act of inference is instantaneous
(12 and 18 as compared with 2, 8, and 11). Lastly, the experi-
ments show that not only the feet (as these affect the boots), but
likewise the whole body of a man exhales a peculiar or individual
odour, which a dog can recognize as that of his master amid a
crowd of other persons (15); that the individual quality of this
odour can be recognized at great distances to windward (15),
or, in calm weather, at great distances in any direction (16);
and that it does not admit of being overcome by the strong smell
of aniseed (14), or by that of many other footprints (4).

MR. F. DAY ON THE LOCH-LEVEN TROUT. 71

On the Loch-Leven Trout (Salmo levenensis).
By Francis Day, C.LE., F.LS.

[Read 2nd December, 1886. ]

SALMO LEVENENSIS, Walker.

Salmo levenensis, Walker, Wernerian Memoirs, i. p. 541 (1808),
apud Neill ; Walker, Posthumous Essays on Natural History
(1812) ; Yarrell, Brit. Fishes, (ed. 2) ii. p. 117, (ed. 8) i. p. 257;
Gunther, Catal. of Fishes, vi.p.101; Couch, Fishes Brit. Isles, iv.
p. 243, pl. cexx.; Houghton, Brit. Freshwater Fishes, p. 128, c. fig.;
Day, British and Irish Fishes, ii. p. 92, pl. exvi. figs. 2 & 2 a.

Salmo taurinus, or Loch-Leven Bull-Trout, Walker, Essays, l. ce.
(large examples).

Loch-Leven Trout, Richardson, Fauna Bor.-Americana, 1886,
p-143 ; Knox, Proceedings Linnean Society,vol.i. p.854, Dec. 1854.

Salmo cecifer, Parnell, Fishes of the Firth of Forth, p. 3806
pl. xxx., and Wern. Mem. vi. p. 146, pl. xxx.

Among the general public, anglers, and fishermen it has, from
almost time immemorial, been a subject of argument as to
whether the Loch-Leven trout should be considered a species
distinct from the burn-trout (Salmo fario) ; and also, supposing
it to be a distinct species, whether it may not be the descendant
of a marine form which, having ascended the river Leven and
obtained access into the loch from the sea, has been unable to
return there. Scientific men have joined in this discussion and
given or refused specific rank to the Loch-Leven trout; in the
meantime, the form in question has been selected as the stock-
fish for the justly celebrated Howietoun fish-farm of Sir James
Maitland, which is within 25 miles of Loch Leven and at about
the same elevation above the sea, and here facilities have existed
for studying the race more alopehy, perhaps, than any other of
our British trout.

In Sir Robert Sibbald’s history of Kinross-shire, 1710, we
read :—“ Loch Leven abounds with fine fish, suchas the salmonds*,

* The term salmond was used so vaguely by some authors as applicable to
both the salmon and sea-trout, that the simple name being given is hardly
sufficient evidence of the presence of Salmo salar. Thus Sir R. Sibbald, in his
‘Scotia Illustrata,’ 1684, divided salmon from salmoneta, and referred to the
latter as follows:—“Salmoneta, qui nostratibus the Salmon-trout” (p. 25).
He also observed, “The Grey trout, or Bill-trout, some of them as large as a
salmond”; but, as I shall show, this grey stage is not the livery of old speci-
mens, and none have been recorded over 10 lb. in weight, it would therefore
seem he referred to sea-trout; again, silvery trout in Scotch lochs are often
classed as sea-trout.

72 MR. F. DAY ON THE LOCH-LEVEN TROUT.

taken in the summer ... . The Grey-trout or Bill-trout, some |
of them as big ay a salmond; greyish skinned and red fished,
a foot long, taken all the year over. Cendue or Camdue in
Trish, Blackhead, having a black spot on the top of its head, is
fat, big as a Dunbar herring, red fished, much esteemed.”

Pennant, in 1769, went to Loch Leven, and observed :—“ The
fish of this lake are pike, small perch, fine eels, and most excel-
lent trouts, the best and the reddest I ever saw; the largest
about 6 lb. in weight ”’ (Journ. 4th ed. p. 69). In his ‘ British
Zoology, 1776, he did not refer to any distinct species existing
in Loch Leven ; but after remarking on the large trouts of Lough
Neagh in Ireland, locally termed Buddaghs, he continued, “ 'Trouts
(probably of the same species) are also taken in Hulse-water, a
lake in Cumberland, of a much superior size to those of Lough
Neagh. These are supposed to be the same with the trout of
the lake of Geneva, a fish I have eaten more than once, and
think but a very indifferent one” (iv. p. 299).

The Reverend A. Smith, ‘Statistical Account of Kinross,’
1793, remarked that “In Loch Leven are all the different species
of hill, burn, and muir trout that are to be met with in Scot-
land, evidently appearing from the diversity of manner in which
they are spotted; yet all three different kinds, after being two
years in the loch and arriving at ? lb. or 1 1b. in weight, are red
in the flesh, as all the trout of every kind in the loch are, except,
perhaps, those newly brought down by the floods, or such as are
sickly. The Silver-grey trout, with about four or five spots on
the middle of each side, is apparently the original native of the
loch, and in many respects the finest fish of the whole. The fry
of all kinds are white in the flesh till they come to the size of a
herring, about the beginning of the third year..... Those
called bull-trout are believed to be the old ones. In spring,
1791, a large one was caught that weighed 10 lb.”

Dr. Walker, in his posthumous ‘Essays on Natural History
and Rural Economy,’ 1812, observed of the trout in Loch Leven :—
“The first most frequent is called at the place Grey Trout, and is
a fish not distinctly described by naturalists; it is found usually
from 1 lb. to 2 lb. in weight, at times considerably larger. ‘This
is supposed to be Salmo levenensis, N. The second, called by
the inhabitants Bull-trout, Salmo taurinus, N., supposed to be a
distinct species ; but there is reason to suppose this is the male
of the above. These two are generally known in Edinburgh as

6 MR. F. DAY ON THE LOCH-LEVEN TROUT. 73

Loch-Leven trout. The third is called at Kinross the Camday,
is 8 in. to 10 in. long, and reckoned a distinct species ; but is
only the grey trout at an early age.” He likewise referred to
three more species as the Burn trout, the Highland or Muir
trout, and another form of bull trout, which he does not appear
to have seen, found in the deep parts of the lake, attaining to
7 1b. or 8 lb. in weight, and with yellow flesh.

Graham, ‘ General Review of the Agriculture of Kinross and
Clackmannuan,’ published about the commencement of the present
century, after giving an account of the fish found in Loch
Leven, remarked, “Flounders are also found in Loch Leven,”
which demonstrated that at this period sea-fishes were able to
obtain access up the river Leven into the lake.

In the year 1874, Mr. R. Burns Begg, the ex-president of the
Kinross Fishing Club, compiled an interesting account of the
Loch-Leven trout, and of the locality which it inhabited. The
Loch-Leven lake, prior to 1830, covered a superficial area of 4312
acres ; it is situated 860 feet above the sea-level, and receives the
waters of the Garny and of the north and south Queich ; while the
mean flow from it throughout the year amounts to 4000 cubic
feet a minute, which goes into the river Leven, and this river,
after a course of 14 miles, falls into the Firth of Forth. In
December, 1830, the loch was diminished to three fourths of its
original dimensions, or to 3548 acres, by an extensive drainage
operation, which permanently reduced its natural level to the
extent of four and a half feet, and means were likewise devised
by which, when desired, another four and a half feet can be
drawn off. Fleming made a careful inspection of the loch
during the years 1834 and 1835, in order to ascertain what
effect the drainage had had upon its fisheries, and he concluded
that they were permanently diminished one-third in their value,
the sluices acting injuriously to young fish by reason of the strong
current at the outflow ; and that the margin of the lake had under-
gone a change unfavourable to its piscine inhabitants, owing to
the peculiar barrenness of the shore rendering the new margin
ill suited for supplying them with food. In the lake itself, how-
ever, the water-snails were found not to have been destroyed.

Many have supposed that the superior flavour of Loch-Leven
trout is a consequence of the quality and abundance of the food
which they could obtain there.

In the ‘ New Statistical Account of Scotland,’ mention is made

74 MR. F. DAY ON THE LOCH-LEVEN TROUT.

of a trout taken April 27th, 1841, that weighed 10 lb., being
27 inches long and 17 inches in girth. We are likewise told of
the fish in this lake, that their superiority in quality is not
confined solely to the Loch-Leven trout proper, but is to be
observed in the common trout, and even in the pike, perch, and
eels; also that the trout of Loch Leven do not continue to
exhibit the same distinctive superiority when they are removed
to other waters. In new quarters, however favourable such may
appear to be, they are said to invariably deteriorate and lose
much of their quality.

The peculiarly excellent food in the water at Loch Leven has
been supposed to consist of a small reddish-coloured mollusk,
believed to be restricted to the shallow shingly beds lying near
to the shores (the form here alluded to would seem to be a
Limnea), and the sessile-eyed crustacean, “screw” or “* water-
shrimp,” Gammarus. Mr. Wilson concluded that it was owing
to the abundant and perpetual breeding of these and other
living creatures that the trout in question owed their supe-
riority. A fisherman, however, who had the management of the
curing of the trout, and had observed the food taken from their
stomachs, remarked that he had never observed any small shells,
but mostly worms, minnows, perch, and young trout. Further-
more, evidence was adduced by fish-dealers and others who had
been regularly supplied with trout, both before and since the drain-
age, who distinctly stated that they could observe no deteriora-
tion whatever in the fish. Parnell, however, held a diiterent
view; and there cannot be a doubt that the stock of fish largely
diminished from some cause.

Whether this form is or is not the Salmo cumberland of La-
cépede, in his ‘ Histoire Naturelle des Poissons,’ vol. v. p. 696,
cannot now be determined from the meagre description which
has been handed down to us; but that author described it ag
having a small head, white flesh, and being externally of a grey
colour. A correspondent of Loudon’s ‘Magazine of Natural
History,’ 1832, vol. v. p. 317, remarked upon a form of trout
which was found in Ulswater and Windermere, termed by the
residents a “grey trout” and having the habits of a char, which
he likened to Lacépéde’s fish, and asserted was captured up
to 20 lb. weight. Parnell in 1838, l. c., appears to be the first
who scientifically investigated this form of trout, and from
his remarks we learn that he considered “the differences that

MR. F. DAY ON THE LOCH-LEVEN TROUT. 75

exist between S. cecifer (as he termed this form) and S. fario
are very striking. The pectorals of S. cecifer, when expanded,
are pointed; in S. fario they are rounded. The caudal fin in
S. cecifer is lunated at the end; in NS. fario it is sinuous or
even. S. cecifer has never any red spots; S. fario is scarcely
ever without them. The caudal rays are much longer in SN. ce-
cifer than in 8. fario, in fish of equal length. In S. cecifer the
tail-fin is pointed at the upper and lower extremities ; in S. fario
they are rounded. ‘The flesh of S. cecifer is of a deep red, that
of 8. fario is pinkish or often white. The cecal appendages in
S. cecifer are from 60 to 80 in number; inS. fario I have never
found them to exceed 46.” He also observed that this fish does
not appear to be peculiar to Loch Leven, as he had seen speci-
mens that had been taken in some of the lakes of the county of
Sutherland.

Sir John Richardson, in the‘ Fauna Boreali-Americana,’ /. c.,
remarked that in “ external form, the proportional size of various
parts of the head and gill-covers, the size of the scales and the
dentition, agrees with S. lemanus..... Three individuals of
the Loch-Leven trout that were dissected had each 73 pyloric
cca, and in one of them 59 vertebre were counted.” Yarrell
added nothing to the previous descriptions. Knox, ‘ Lone Glens
of Scotland,’ 1854, observed of this trout of Loch Leven, that it
“ig a beautiful silvery dark-spotted trout, imagined by some to
be peculiar to the lake. This, however, is not likely, since trout
quite resembling those of Leven are found in many northern
lakes” (p. 36). He concluded, after citing some of the opinions
of others, that he was ‘“ disposed to think that two species of
trout inhabit Loch Leven, independent of the common river
trout; namely, the trout which lives on entomostrace, and comes
into season in December, January, and February ; and the trout,
which, feeding on the buccinum, and on flies, worms, and all the
common food of the common river trout, comes into season later
in the spring” (p. 37). In the ‘ Proceedings of the Linnean
Society ’ (Dec. 19th, 1854), Dr. Knox remarked that at first he
thought this a specific form, “although anatomical investigation
has not hitherto confirmed it.”’

Dr. Giinther, 7. c., gave a fuller description than the previous
authors whom I have quoted. He observed of this fish that it
has the “ body much less stout than in S. fario.... In the
male sex a mandibular hook has never been observed. Maxillary

76 MR. F. DAY ON THE LOCH-LEVEN TROUT.

much longer than the snout, but much narrower and more feeble .
than in 8. fario (see figures, p. 6); in specimens 13 inches long
it extends to below the hinder margin of the orbit, and at no
age does it reach much beyond it.... The teeth of the body of
the vomer form a single series, and are persistent throughout
life. ins well developed, not rounded.’ He found from 49 to
90 cecal appendages. At page 6, l. c., are figured two maxillary
bones, stated to be from 8. fario and S. levenensis*, but the
drawings not being completed at their proximal extremities
render it almost impossible to understand what they are intended
to represent. The supplementary bone would seem to be where
the most difference exists. The same author likewise remarked
(p. 7) on the question of species in Salmonide, and gave his
reasons for admitting certain forms to that rank, stating that
“whenever the zoologist observes two forms distinguished by
peculiarities of organization such as cannot be conceived to be
the effects of an internal or external cause, disappearing with
the disappearance of that cause, and which forms have been
propagated and are being propagated uniformly through all the
generations within the limits of our observation, and are yet
most probably to be propagated during the existence of mankind,
he is obliged to describe these two forms as distinct, and they
will commonly be called species.”? Dr. Ginther has also stated,
at a meeting of the Zoological Society, that the late Sir J.
Richardson had informed him that he believed the true Loch-
Leven trout had disappeared from that lake.

In giving a decision on the well-known “ Orange-fin ” case, in
1872, the Sheriff Substitute found that “in reference to the out-
ward silvery appearance of the fish in question, both Dr. Giinther
and Professor Young state that the silvery coat with which these
fishes is clothed is to be regarded as a distinctive mark of their
being migratory fish of the salmon kind. The assumption of the
silvery coat... . in the case of river fish, is to be held an almost
infallible test of a migratory and sea-going habit. Nor is this
inconsistent with the well-known fact, that in the case of certain
fish which inhabit lochs having now no communication with the
sea, a similar silvery appearance is to be seen. In the case of

* The teeth in the maxilla of S. Jevenensis, in Dr. Giinther’s figure, are
shown as directed forwards and inwards ; the base of each tooth appears as if
resting on the skin with its point turned towards the maxillary bone!

= 7 = SO oe

MR. F. DAY ON THE LOCH-LEVEN TROUT. TF

the Loch-Leven trout, which affords the most notable example
of the phenomenon referred to, it must, however, be kept in view,
as having an important bearing on the character of this fish, that
the loch which it inhabits had, most probably, at one time a
communication with the sea, and that the fish themselves possess
in a most remarkable degree the features of the salmon and the
sea-trout .... The fact of one of these trout having, in the
course of the present trial, been regarded by Professor Young as
a sea-trout, after examination, is a very strong testimony to the
difference between the characteristic features of the Loch-Leven
trout and those of all the non-migratory river-fish ” (pp. 166, 167).

In the year 1873 Sir James Maitland commenced fish-culture
at Howietoun, and selected as the form of trout which he con-
sidered would prove best adapted for this purpose the true
Loch-Leven breed, the eggs of which he obtained at the lake,
and from which his present stock originated *. For some years
he has assisted me, and allowed every facility for examining
the fish in his establishment, while I have given very particular
attention to the following question, Js the Loch-Leven trout a
distinct species or merely a local race?

The first inquiry will therefore be, On what grounds has the
Loch-Leven trout been regarded as a distinct species? Canany
persistent differences from other trout be shown in its external
form, its internal organization, its tints, or the colour or taste of
its flesh ?

As to external form, the Loch-Leven trout has been said to be
much less stout, its head shorter, its fins more pointed, while the rays
in its caudal fin are longer than those in the burn-trout ; the poste-
rior extremity of this fin is also said to be lunated and pointed at
both its upper and lower angles, and its pectoral fin is likewise
pointed. Also that the male has no hook on the lower jaw; that its
maxillary bone is more feeble than in any other form of trout, and
that it never extends posteriorly beyond the hind margin of the
orbit.

As to its body being less stout than that seen in burn-trout,

* Others have likewise stocked pieces of water with these fish, but with vary-
ing success. Thus Knox (‘Lone Glens of Scotland,’ 1854) remarked upon “ the
artificial Lake of Prestmannan, into which, some years ago, the beauteous Trout
of Loch Leven had been introduced. Under circumstances highly disadvanta-
geous they throve, notwithstanding, tolerably well, and even bred at the entrance
of a small stream which mainly supplies the lake” (p. 35).

LINN. JOURN.—ZOOLOGY, VOL. XX. 7

78 MR. F. DAY ON THE LOCH-LEVEN TROUT.

this is certainly sometimes the case, but such a character is not
persistent. In the form of the body there is no difference percep-
tible in those fish reared in Gloucestershire, as I shall presently
show, or at Mr. Andrews’s establishment at Guildford, from eggs
obtained from Howietoun, and young of brook-trout raised from
local brook-trout eggs. As this is the case in fish reared from Loch-
Leven eggs it is evident that their form ultimately depends upon
local circumstances or conditions ; for if they are removed to an-
other locality where the conditions are different, the brook-trout
form is at once seen. The same observations apply to the length
of the head, which in some well-fed examples, and, in fact, gene-
rally in the young raised at Howietoun, is a little shorter in pro-
portion to the length of the body than usual in the brook-trout,
but this changes on their being transferred to a new home.

As to the length of the rays of the caudal fin* being longer
than in the brook-trout, I have been unable to find that such is
the case, either in specimens from Howietoun, from Loch Leven,
or in those in the British Museum, as they seem to be absolutely
identical in the two forms. In a skeleton of a female 20 inches
long, I find the middle caudal ray is 2°1 inches in length, and the
longest outer ray 29 inches ; but were Dr. Giinther’s figures, as
noted below, to be applicable to these fish, the outer ray should
be 41 inches in length. Such proportions I have never seen in any
of the thousands of these fish I have observed at Howietoun or
elsewhere, not omitting those in the British Museum. Specimens
having the angle of this fin pointed would appear to be young
fish, often males, kept, as at Howietoun, where they are not dis-
turbed; but in the old fishes this fin is invariably rounded at its
posterior extremity.

The statement that the pectoral fin is pointed is partially cor-
rect in small specimens, as it also is in small brook-trout, but in
old and well-preserved examples it is as rounded as in other
races of freshwater trout f.

As to the male having no knob on the lower jaw, that likewise

* “Tn specimens 13 inches long, the middle caudal rays are not quite half as
long as the outer ones, and in older ones they are half as long.” (Giinther.)
In a specimen 133 inches long the middle caudal ray was 1 inch in length and
the outer or longest one 13.

+ In order to demonstrate this, dried examples of the pectoral fin taken from
fishes of this race at various ages were shown at the Meeting when the paper
was read,

MR. F. DAY ON THE LOCH-LEVEN TROUT. 79

is a most erroneous assertion *; in the one figured as a diagram,
which measures 14°6 inches in length, and was 44 months old
when caught in October 1886, the knob is very well deve-
loped. This knob is constantly seen in all old males of this form ;
while even in some old females at Howietoun a small one is
occasionally perceptible.

As to the maxillary bone being “ much narrower and more feeble
than in S. fario, in specimens 13 inches long it extends to below
the hinder margin of the orbit, and at.no age does it reach beyond
it” (Gunther), Is this so? This statement as to where the
maxillary bone extends posteriorly, first made by Dr. Giinther,
is not borne out by an inspection of Howietoun fish, in which in
large specimens it extends from one to two diameters of the orbit
posterior to the eye, and this is of normal occurrence. In an
exainple 26 inches long it reaches to 13 diameter of the orbit
behind the eye; the longest fish in that establishment having a
total length of 27 inches. For as they become more or less
sterile at from 8 to 10 years of age, to which breeding males
rarely if ever attain, older fishes are not present.

Doubtless the maxilla and teeth with which it is armed are not
so strong at Howietoun as in some (not all) brook-trout of the
same size; and the cause of this feebleness in the jaws, which
ceases under altered conditions, is immaterial to discuss, because
we have no evidence pointing as to how it first commenced.
Although perhaps it may be likened to what is seen in some sea-
trout, it may also be observed in many loch-trout in the north
of Scotland, with whose jaws I have compared those of the Loch-
Leven fishes. The question, in fact, now is, what will be the result
on the form and strength of the jaws and teeth when these fishes
are transferred to a new locality to battle their own way in the
world? Anyhow, figures of the comparative strength of the jaws
in a Loch-Leven female trout and one of the brook-trout of the

* Since this paper was read, the following observation appeared in the ‘ Bir-
mingham Daily Post’ of Dec. 11, illustrating how erroneous statements as to
specific differences in species may give rise to a possibly false conclusion :—
“The big trout which was recently captured in the Birmingham Corporation
Reservoir at Shustoke has been mounted by Cooper, of London, and may be
seen, during next week, at Keeling’s fishing-tackle shop, Digbeth. It weighed
when caught 84 lb., and its length was 27 inches. The formation of the lower
jaw shows that it was an old fish, and not one of the Loch-Leven trout with
which the reservoir was stocked three years ago.”

80 MR. F. DAY ON THE LOCH-LEVEN TROUT.

same size and sex as published by Dr. Giinther are very mis-
leading ; for I cannot find such examples in the British-Museum
collection showing what he has represented, neither can I in
nature *.

Having thus seen that in its external form either the differ-
ences which have been stated to exist between this fish and the
brook-trout are erroneous, or else lable to alteration when
the fish is removed to another locality, all must allow that such
non-persistent differences are no basis upon which to found a
species. |

As to external colour, we find Parnell asserting, as among its
specific characters, “ body without red spots;” and that these
fish are generally seen without them up to a certain age is of
very common occurrence, they being of a grey colour densely
spotted with black, and if males, with the fins almost black. At
Howietoun, three main types of colour are observable amongst
these fish—a slaty or greenish grey, becoming lighter beneath,
and the upper two thirds of the body and dorsal fin spotted with
black, and the fins generally greyish black. This form of colour
is prevalent up to the end of the fourth season, and may almost
be looked upon as equivalent to the silvery stage of the salmon
smolt or grilse, but I have never seen one over four years of age
continuing this livery. The second*¢ or adult form is of a general
purplish golden, densely covered with black spots, among which
some red ones are usually to be seen, and many old females get
a dark line along the middle of the belly, which, as well as the
under surface of the head, is more or less black in males. In one
female 18 inches long, on November 24th, three bright orange
spots were present on the adipose dorsal fin, which as a rule is
of a lead-colour, with two or three black spotst. The third form,
which will have to be again referred to, consists of small under-

* Specimens and diagrams were shown at the Meeting.

+ When old enough to feed on clams, which are about the size of marbles,
this yellow colour shows itself. In some small examples, hatched in 1883, the
“ finger-marks” were very distinct on netting the pond at the end of November
1886.

t We must not forget that brook-trout vary greatly in colour even when in
the same locality ; thus “Ephemera” in 1853 remarked of those in the Wandle
that such as “ feed under the cover of the trees, or lie perdu under banks or arti-
ficial ‘ hides’ during sunshine, are dark brown and yellow; those that frequent
the unshaded streams with a clear sandy bottom are of a silvery hue” (p. 274).

'
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MR. F. DAY ON THE LOCH-LEVEN TROUT. 81

sized fish, which, owing to sickness or some other cause, have the

colours of the brook-trout, with orange-tipped adipose dorsal

fins.

The question here arises whether these colours are perceptible
in these fishes when removed to fresh localities. In the year
1868, the late Mr. MclIvor, of the Government Gardens at Oota-
camund in the Madras Presidency, succeeded in introducing some
Loch-Leven trout and other European fish to that elevated
region, where they are or were doing well (see Journal of the Lin-
nean Society, Zoology, vol. xii. p. 562). In January 1876, Mr.
Thomas, F.L.S., of the Madras Civil Service, sent me a specimen
from the Hills which was 64 inches long, and on its body were red
spots. In this instance it was clear that if a young Loch-Leven
trout could assume red spots when removed to Asia, there was no
reason why any similar movement in Europe might not occasion
the same results.

The assumption of the general colours of the trout in any
given locality by introduced breeds is of very common occurrence,
at least after the third year. Now this is the period at which
the young of the imported forms would be in a condition to be
observed by the fisherman, whether angler or netter.

This is generally asserted to be owing to the imported fish
having interbred with the local race, and the hybrid (as it is
wrongly termed) or mongrel form has the local colours. It is
therefore interesting to ascertain whether, were eggs removed to a
given spot quite distinct from the waters where the parents reside,
the young which emerge from those eggs would retain the colours
of their parents or assume those peculiar to the locality; for
if this latter occurs, it must be evident that such has been con-
sequent upon local surroundings.

Ten thousand yearlings from Howietoun were turned into
Loch Goldenhove, about two miles away, and fed by the same
stream which passes through the fish-farm ; this loch is nine acres
in extent, and averages six feet in depth. In July 1886, I exa-
mined some of these introduced fish, and found them of a pur-
plish colour shot with gold, and covered with black ocellated spots,
but no redones. Dorsal fin spotted with black, but without any
white edging, its outer surface greyish ; a little orange upon the
adipose dorsal fin. The colours, in fact, of these fish were not what
is seen in the Howietoun ponds, but nearly approaching those
in the Loch where they had been placed. A few had some red

82 MR. F. DAY ON THE LOCH-LEVEN TROUT.

spots. As the water in the two localities was the same, food would
seem to have been the principal reason of this change in colour.

Although the last experiment does not prove very much, it
shows that some alteration in colour may follow new environ-
ments ; buta still more conclusive result as to the change in colour
which may take place in these fish under like circumstances has
lately occurred in Gloucestershire. The present proprietor of
Cowley had two ponds in a wood on his estate, each about an
acre in extent, and from the lower of which springs a small
stream ; these he wished to have stocked with some good kinds
of sporting fish. The two ponds are supplied by underground
springs, while there is a fall of about 16 feet from the outlet of
the upper pond, and a rather greater one from that of the lower
where the stream commences. It will thus be apparent that no
fish could obtain access from above, neither could they ascend
the 16-feet perpendicular fall from the stream to the lower pond.
It was determined to try the Loch-Leven trout; so these ponds
were drained, mudded, and then puddled with elay.

During December 1884 and January 1885 one thousand yearling
Loch-Leven trout were received from Mr. Andrews of Guildford *,
and these were placed in the ponds by Mr. Ogden, of Cheltenham.
In August this year (1886) I was informed that, it having become
necessary to remove these fish to a more suitable locality, they
had been capturing them, and very great differences were percep-
tible both in size and colour among the two sets of fish—those
in the upper pond being silvery with a few black spots, whereas
those in the lower pond were of a much larger size, covered with
spots, and having purple and golden reflections. Having obtained
leave, we visited these ponds on August 25, and first examined
the temperature of the water by means of thermometers, when
we found that they scarcely differed. The lower pond was the
deeper, and in it were large quantities of the American weed,
Anacharis Alsinastrum, also some Chara, while on the surface
was a considerable amount of the water crowsfoot (Ranun-
culus aquatilis); whereas in the upper pond there were fewer
weeds, but some Pimpinella Saxifraga was present near its upper
end.

* Mr. Andrews, of Westgate House, Guildford, wrote (September 14, 1886) :—
“ The yearling fish supplied to Mr. Ogden, of Cheltenham, in 1885, were Loch-
levens reared from eggs which were sent me from Howietoun. ‘There can be

no doubt of their being from ova from Stirling, as they were put in a pond
quite distinct from the others on a different water-shed.”

MR. F. DAY ON THE LOCH-LEVEN TROUT. 83

It seemed, so far as we could ascertain, that more animal pond-
life was present in the lower pond among the greater amount of
vegetation, especially the sessile-eyed crustaceans Gammarus
pulex and water-snails (Limnea ovata, variety peregra). These
forms were, however, also present in the upper pond, where a
small water-newt was also netted. During the month of April
this year, when investigating the stream which issues from the
lower end of these ponds, I found enormous numbers of tadpoles
and the larve of Ephemeride and their allies.

A fish captured in the upper pond was seven inches long, its
colours generally silvery with a golden abdomen, and a few black
spots along the sides, three of which were below the lateral line
and two on it, as well as three red ones ; cheeks silvery yellow,
abdomen golden. Some spots on the rayed dorsal fin, which
had a white black-based edge at its upper angle; while the
ventral and anal fins had a very distinct white black-based
edging. Upper and lower rays of caudal fin and the upper
end of the adipose dorsal orange-edged. 52 cecal appendages.
A male, but sterile. The appearance of this fish as to colour
was, Mr. Ogden observed, similar to the others removed from
the pond.

Two fishes were taken from the lower pond, one ten, the other
eleven inches long. They were generally purplish, with golden
reflections. The side of the body (of one which was most criti-
cally examined) from the upper edge of the pectoral fin to and
above the lateral line was closely dotted with ocellated black spots,
while there were also some red ones, five of which were on the
lateral line. Cheeks golden ; abdomen golden, becoming white
on the chest. Dorsal fin with numerous spots, and a pink black-
edged upper angle. Adipose dorsal with a red edge and several
black spots. Pectoral and ventral chrome with white edges, base
dark. 62 cecalappendages. A male, with the generative organs
well developed. 22 small shells of Limnea in its stomach.

As regards the colour * on the adipose dorsal fin, I examined
at Howietoun, on August 15th, 1886,a number of two-year-olds,

* An interesting occurrence has taken place at Cowley among these fish,
which would seem to prove that Lochlevens throwing back to brook-trout
may be consequent upon a diminished supply of food causing deterioration.
During the last week in November Mr. Ogden was near this lower pond and saw
a large trout rise ; so he returned to the house, and having obtained his fishing-

tackle, made a cast over it and captured it at the first throw. The fish proved to
be 15 oz. in weight and in good condition, but was described to me as having

84 MR, F. DAY ON THE LOCH-LEVEN TROUT.

with the result that there was merely a trace of orange visible in
some, but the black white-edged margin to the rayed dorsal and
anal fins was not uncommon.

On July 5th, 1886, I examined the colour of the adipose dorsal
fin in many Lochlevens, from pond 9; the first was rising 2-years,
and was five inches long. It had two black spots on the adipose
dorsal but not a trace of orange, and no red spots on the body.
Several more subsequentiy looked at were the same. Three had a
slight orange tinge on the adipose fin, and likewise a few red spots
on the body ; in fact this fin was orange-tinted in all, wherein red
spots existed on the sides, but the presence of these red spots
was the exception, not the rule. Passing on to the nursery-
ponds, we examined a few undersized Lochlevens which had not
fed well; all had their bodies red-spotted and also red on the
adipose dorsal fin, while the rayed dorsal had a more distinctly
black white-edged margin than was generally seen. Some of
the fish had 2, 3, or 4 black spots on this fin. These fish would
at once have the brook-trout livery and not pass through
the normal silvery stage—a stage, we have been erroneously
informed, which is an intallible test of a trout being migratory
and sea-going.

Having paid a visit to Mr. Andrews’s* well-known fish-cultural
establishment at Guildford, I was shown the yearling Lochlevens
which had red spots and red edges to the upper margin of the
adipose dorsal fin and on each lateral margin of the tail-fin.

Here I must shortly digress to describe a hybrid specimen
raised at Howietoun, which possibly affords one of the most
convincing proofs that could be adduced of the identity or close

been nearly black, while it was the largest fish that has been seen. From the
fact that this pond lately had had very few fishes in it (they having been
removed), a more abundant supply of food was obtainable, and consequently
this trout had grown larger and taken on the Loch-Leven trout colours.

* Mr. Andrews (MS., Noy. 1886) finds at Guildford that ‘‘eggs of the
Loch-Leven trout from Howietoun do very well with him; they are hatched
in water coming from chalk, and reared where it comes from a gravelly soil.
The young grow more rapidly and are deeper in form than seen at the same
age in their native home. ‘The yearlings have the edge of the adipose dorsal
fin and sides of the tail-fin red, and there are also some red spots on the
body, in common with young of the brook-trout and some reputed as S. ferox.”
Mr. Andrews continues that he ‘knows of no unmistakable peculiarities ob-
servable in the foregoing three varieties of British trout;” and I question if
any one else is able to point out their existence.

MR. F. DAY ON THE LOCH-LEVEN TROUT. 85

relationship of the Loch-Leven and burn-trout. In November,
1883, some eges of a Loch-Leven trout were milted from a salmon-
parr at Howietoun (which specimen I still possess in spirit), and
hatched in March 1884. As I have elsewhere related, most of
the young died of dropsy, but a few lived, and some are still at
Howietoun in pond 16, very many being small, but a few of fair
size, Some even giving eggs and milt this season. On November
24th I removed one of these fish, a beautiful parr, showing the long
pectorals and large caudal fin of the salmon, but having only nine
finger-marks; while as to colours it was of a beautiful silvery
glossed with gold, the rayed dorsal fin rather densely spotted
with black, some of the spots having a deep scarlet edging, and a
white margin with a dark base being present at the front upper
corner of this fin as well as of the anal. Adipose dorsal red-edged ;
caudal straw-coloured, with red upper and lower edges. Numerous
black spots in the upper half of the body, also some red ones, but
most of the latter along the lateral line and some below it. A
large black spot on opercle and some smaller ones.

Here was a hybrid showing the number of bars of the trout,
and also most of the trout-colours, but with this remarkable varia-
tion from the Loch-Leven breed, that the orange edging was
present on the adipose dorsal * and the light edging on the dorsal
and anal fins. In fact the spots on the dorsal fin closely resembled
those seen in the burn-trout in Sutherlandshire. Unless the
Lochlevens have burn-trout blood in them, how could these fish
possibly throw back to the colours of the latter race ?

It is normal for the adipose fin of the Loch-Leven trout to have
black spots upon it, but no orange margin. ‘The reason is pro-
bably similar to that which causes these fish to have no orange
spots on the body, because if orange spots exist upon the body
the adipose dorsal is also orange-tipped. Evidently whatever
causes this coloration in one part of the body equally does so in
the other.

As to the internal organization of the Loch-Leven trout, we
have been told that it possesses from 49 to 90 cecal appendages,
and that the teeth along the body of the vomer “ form a single
series, and are persistent throughout life.” In fact from the time
Parnell first ascertained that these trout often possess a larger

* Not only is the adipose dorsal normally destitute of an orange edging or

orange spots in both young salmon and young Lochlevens, but this colouring is
present in many sea-trout with which young Lochlevens have been compared.

86 MR. F. DAY ON THE LOCH-LEVEN TROUT.

number of these appendages than are usual in brook-trout until »

the present day this has been held conclusive evidence as to their
specific difference from other forms. Many who would admit that
variations in external colour or in that of the flesh, or even altera-
tions in form, may be dependent on local surroundings, will be slow
to believe that structural differences are not of much greater value.
Hence we must first inquire whether the number of these cecal
pylori are constant in the Loch-Leven race of trout, whether they
ever vary in the brook-trout, and, lastly, if any facts can be pro-
duced proving them to be inconstant. |

Among the local Loch-Leven forms we are told by Parnell that
the cecal pylori are from 60 to 80 in number. Sir J. Richardson
found 78 in each of the three which he dissected, and Giinther from
49 to 90; and although in the description of the species the latter
writer says, “Czcal pylori normally 60 to 80,” he instances seven
females in the British-Museum collection as follows :—“ Females,
from 12 to18inches long. Purchased, said to be from Loch Leven.
Caught in April. Cecal pylori 65, 63, 60, 54, 54, 53, 49;
vertebra 58-59. These specimens have the pyloric appendages
fewer in number than is generally stated ; yet these cxca are so
wide—so much wider than in S. fario, that the reduction of their
number has evidently been caused by a confluence of several
ceca into one” (Catal. vi. p. 101).

From the foregoing statements it is evident that the number
of these appendages is very variable, for we have them stated as
being from 49 to 90. If, however, we turn to the writings of most
authors who have counted the cecal pylori in 8. fario, we find them
enumerated as follows :—‘“ I have never found them to exceed 46”
(Parnell, ‘ Fish Firth of Forth,’ p. 308). Thompson in 18386 ex-
amined the so-called S. ferox, and found in four examples 49, 45,
839, and 86 (‘ Nat. History of Ireland, iv. p. 157). Gtinther
among his other five non-migratory freshwater forms enumerated
them as varying from 38 to 49.

Having thus shown that these appendages in the Loch-Leven
trout have been recorded as between 49 and 90, while in othernon-
migratory freshwater forms they have been found to be between
33 and 49, I propose enumerating some which I have counted in
examples of this fish. Among males, in specimens varying from 7
to 204 inches in length, I have found them as follows :—At Howie-
toun, 8 examples of fertile fish averaged about 67 cecal appen-
dages, founded on these numbers—82, 74, 74, 73, 65, 62, 62, 48.

MR. F. DAY ON THE LOCH-LEVEN TROUT. 87

At Cowley, in Gloucestershire, one fertile male had 62, one
which was sterile 52. Among females, varying in length from 12
to 22 inches, the number of these appendages in 7 examples of
fertile fish averaged about 58, derived from the following numbers
—66, 64, 62, 59, 57, 55, 45; while one large female from Loch
Leven had 47. Asa rule these ceca appear to be larger in females
than in males, while in one of the latter a single one of these
tubes was abnormally shortened near the pylorus.

The foregoing figures show a variation at Howietoun in the
number of these appendages, ranging from 48 to 82 in male fish,
and from 45 to 66 among female fish ; while in one male which
was fertile, examined in Gloucestershire, there were 62, and in
another, not so well fed, from the same locality, and sterile, there
were 52. In only one is the number seen to approach 90 (82)
as given by Dr. Giinther, and with that exception 75 was the
largest number counted, and from that down to 45, clearly
showing that this is an unstable character, prone to change,
and consequently unsuitable for discriminating species. Also
that, away from Loch Leven, these appendages have diminished
in number, and still more so in examples from the ova hatched
at Guildford and reared in Gloucestershire, where the smallest
fish were sterile and had the fewest of these appendages. As to
the diameter of the cxca, the difference was not apparent, except
so far as I have mentioned above.

Possibly the number of the cecal appendages may, under certain
circumstances, be found of value in ascertaining whether the food
on which the fish subsist is of such a nature that they will thrive
or deteriorate.

If these appendages decrease when the fish are transferred
to other localities, it may be asked if instances can be adduced
where they have ever been found to increase in trout when
removed to better feeding-grounds or improved conditions of
lite*. The eggs sent from the brook-trout of Hampshire and
Buckinghamshire by Mr. Frank Buckland and Mr. Francis
Francis to Tasmania have developed into a large race, in which
the cxcal pylori seem to have reached the normal number of 52,
showing a considerable augmentation, and again proving the
number of these organs to be inconstant.

Respecting the vomerine teeth being in a single series in Loch-

* Since this paper was read (viz. in March 1887) young rainbow-trout
(Salmo iridens), 22 months old, raised at Howietoun, have been examined. In

California, whence the eggs were received, these fish are said to possess about 40
excal appendages; in one dissected at Howietoun I found 71.

88 MR. F. DAY ON THE LOCH-LEVEN TROUT.

Leven trout, but in a double row in burn-trout of the same size, we —
have first to consider whether the facts as stated are correct. All
trout and salmon (not charr) when young, irrespective of the teeth
on the head of the vomer, have a double row along its body ; but
these in all are dependent on age or rapidity of growth, and fall
out, commencing behind and extending forwards. In salmon and
sea-trout, which are forms that grow most rapidly, these teeth are
shed the earliest, while the Loch-Leven trout, which is likewise a
rapid grower, loses them rather sooner than the burn or loch
form; hence to say that in the mature examples they are invariably
in a single row is erroneous. At the same time it is not here
advanced that rapidity of growth is the sole cause of this, for
the deciduousness of the teeth appears to be owing to the absorp-
tion or narrowing of the tooth-bearing ridge on the vomer, in
consequence of which the teeth, originally placed in pairs, become
ultimately ranged in a single row and finally fall out.

The condition of the vomerine dental system in specimens of
Loch-Leven trout may be thus recorded :—(1) 2 20 inches
long, 2 teeth exist on the hind edge of the head of the vomer,
and 3 along the front half of its body, the first two of which
are almost opposite one another. (2) ¢ 9 inches long, 2
teeth at hind edge of vomer, 2 at the front end of the shaft,
and 7 in an irregular zigzag line, almost in one continuous row,
while their points turn alternately to the right and left. (8) 2
10°9 inches long, 2 at hind edge of head of bone, 2 in a line
at the commencement of the body, and 8 as in the last but more
distinctly in pairs. (4) 2 18°5 inches long, 2 at hind edge of
head of vomer, then 4 in a single row, next a pair turning one to
each side, and lastly 4 more single ones. (5) ¢ 14 inches long,
2 teeth at hind edge of head of vomer, 9 in a single row along
the body of that bone; of these the two central ones form a pair.
(6) 2 19°2 inches long, 4 teeth at hind edge of body of the
vomer, 12 along the body, among which are three pairs. (7) 2
23 inches long, 3 at hind edge of head of vomer, 8 along its body
in a single row, some turning one way and some another.
The foregoing seven specimens show that it is by no means an
invariable rule that all the teeth along the body of the vomer are
in a single row.

Having examined a considerable number of burn or loch trout
in Sutherlandshire, I found that the maxille were not stronger
than in the Lochlevens, while a specimen* taken at random

* Exhibited when the paper was read.

—ssveye= =

12 2. Ft. &

MR. F. DAY ON THE LOCH-LEVEN TROUT. 89

shows the same dentition as in the form under discussion : burn-
trout, ¢ 11 inches long, 2 teeth at the hind edge of the head
of the vomer, 10 in almost a single line along the body of that bone,
turning alternately right and left at their points, but in only one
instance are they ina pair. Further south, as in the Windrush
in Gloucestershire, the maxille are stronger and the vomerine
teeth are more nearly in a double line than in more northern
specimens; but an entire series, showing all these grades of
variation, may be traced in freshwater trout in most localities
where I have searched for it.

The colour of the flesh of the Loch-Leven trout is said to be
deep red, and it is reputed to be very good eating. Whether
the flavour of these fish has or has not deteriorated since
the partial draining of the lake, as asserted by some and con-
tradicted by others, must ever remain unsolved, because how
the fish were cooked, the degree of hunger in the partakers
of the food, and many other circumstances would have also to
be taken into account; while deciding such a question from
recollection would be a rather doubtful proceeding. There is a
legend that in olden times these fish never took a fly; and an
anonymous writer in 1886, commenting upon the bad luck which
had attended an angling competition, observed that fly-fishing on
Loch Leven had been in existence for about 25 years, but
previous to that time these fish showed no disposition for winged
prey. Granting the general accuracy of this statement would
seem to partially confirm the opinion of Parnell and some
others, that the local food has diminished in amount, and therefore
these fish will now take the fly. Why the charr has disappeared
from this lake is not material to the present inquiry.

As food Parnell held that at Loch Leven the flesh of this form
of trout is of a dark red, but in the common loch or burn-trout
pinkish or often white. This, however, cannot be held as distinctive
of species, for some trout captured on the same day at Loch
Assynt, in Sutherlandshire, showed all variations in the colour of
their flesh, from white to red, and were all equally well tasted.
Parnell also observed that “ James Stuart Monteith, Esq., of
Closeburn, caught a number of small river-trout, and transferred
them to a lake (Loch Ettrick), where they grew rapidly ; their
flesh, which previously exhibited a white chalky appearance,
became in a short time of a deep red, while their external
appearance remained the same from the time they were first put
in” (op. cit. p. 307).

90 MR. F. DAY ON THE LOCH-LEVEN TROUT.

Mr. Ffennell, writing of the Lochlevens at Mr. Andrews’s .
(Times, Oct. 14th, 1886), observed that “the Loch-Leven trout
were no strangers to me; I have killed many, and as a fresh-
water fish I hold them in high esteem: and I certainly think that
those I took from the roadside pond in Surrey were the very best
I had ever placed before me.”

The question whether the Loch-Leven trout is a local race
or a distinct species, is one of considerable practical im-
portance to the fish-culturists of this country, quite irrespective
of its scientific interest. If it is a species distinet from the
brook-trout, its introduction into our streams and dissemination
through our fresh waters would be a great source of hybridization
among our indigenous forms, and this would tend towards sterility
of the offspring. On the other hand, if it is merely a local race,
its crossing with the brook-trout would be merely the inter-
breeding between two varieties of one species, which, instead of
being a cause of sterility, is more commonly a means of improving
a breed.

I assume it as granted that the Howietoun fish are in every
respect similar to those of Loch Leven, whence the breed was
derived. This variety is sometimes, not always, finer shaped
towards the tail, and has a rather shorter head, as in the sea-
trout, S. albus, than the ordinary brook-trout, while its form
is much like that of the Salmo lemanus or loch-trout of the
Luke of Geneva and elsewhere, which has been shown to be
a variety of S. fario. Its maxillary bones are somewhat finer
than in our ordinary river-trout ; its cecal appendages are more
numerous; and its colour differs, being as a rule silvery with black,
but having no red spots up to its fourth or fifth year. Dr.
Ginther’s observations that the male never has a knob on the
lower jaw, that its fins are never rounded, that the teeth along the
body of the vomer are always in a single row, very different from
those of the brook or burn trout, may be dismissed as erroneous
statements, probably made owing to the want of opportunity of
examining specimens. The Loch-Leven trout is doubtless a
rapid grower in its northern home, and the race at Howietoun
has been much improved by selection of breeders ; but removing
the eggs to a new locality and then rearing the young has shown
that the form and colour of the local race of trout is, as a rule,
assumed, while even the number of cecal appendages becomes
altered, owing to changed condition of life.

f
.
"
:
:
i.

MR. E. C. BOUSFIELD ON THE GENUS DERO. 91

The Natural History of the Genus Dero. By Epwarp C. Bous-
FIELD, L.R.C.P. Lond. (Communicated by Dr. J. Murte,
F.L.S.) |

[Read 4th November, 1886.]

(Puatrs ITI.-V.)

At the meeting of the British Association in 1885, at the request
of Prof. McIntosh, I presented a summary of the results of
my study of the Annelids of the genus Dero, with descriptions
of some new species which had come under my notice, and
remarks as to the identification of others. From the necessary
limits of such a communication, I was compelled to omit
much detail; and lack of time and material led to a certain
want of accuracy in my collation of the work of other writers,
and the comparison of the examples which came under my own
notice with their descriptions.

Enlarged facilities of reference having now enabled me to
acquire a tolerably complete knowledge of all that has hitherto
been written on this genus, and a considerable supply of material
from various sources having given opportunities for extended ob-
servation and closer comparison, I venture to offer the following
as the most complete account of the genus allowed by the present
state of knowledge. Four new species having been met with in
the course of the eighteen months or more during which I have
been working at these Annelids, it would appear probable that
others still remain to be discovered ; and it is to be hoped that
the publication of a summary of what has at present been accom-
plished may facilitate future observation.

History and Bibliography.

The Annelids with which this paper deals, though their biblio-
graphy covers a period of a century and a quarter, have hitherto
been the subject of very little original work, owing probably to
their habits of life, which prevent their presence in a state of
nature from being easily discovered. In view of the great advances
which have been made in methods of research, and especially in
optical appliances, the very evident way in which the observations
of earlier writers, and their interpretations (not unfrequently
erroneous), have been copied by later authors, in many cases
almost verbatim et literatim, is not a little surprising.

92 MR. E. C. BOUSFIELD ON THE

The first description of any species of Dero is to be found in |

the ‘ Insecten-Belustigung,’ iii. Theil, of Résel von Rosenhof (24*),
published in 1755. Several figures and a lengthy description
are there given of a species identical with Dero furcata, under
the name of “das geschmeidige Wasserschlinglein mit zwey
Gabelspizen.” The remarkable feature of Rosel’s account is
the moniliform arrangement of the two long palpi by which
this species is distinguished, which he described. This character
has not been noticed by Grebincky, Semper, or myself; but
the explanation is not far to seek. One of the first signs of
approaching dissolution in the Naid family is the separation of
the cuticle from the cells of the epidermis, with formation of
bulle; and under these circumstances something of the kind
has been observed by the writer. It is somewhat curious
that up to 1877, when the last.description of this species was
written, every observer who described the form gave it a fresh
name; so that it has received trivial names from its discoverer
and from Miller, and scientific ones from Oken, Grebincky,
Leidy, and Semper. Résel observed the process of transverse
fission in an advanced stage, and described it as a “curious
method of copulation.” He also described the result of transverse
sections of the worm.

Following Rosel, the distinguished naturalist O. F. Miller (18),
in 1771, gave a remarkably excellent account of one or two
species; all the main features having been grasped, and the
branchial processes (the characteristic feature of the genus)
having had their proper function assigned to them. In specific
characters, however, Miller was less fortunate; and to him must
be given the blame of the confusion in the nomenclature of the
genus which so long existed. In consequence of the small scale
on which his figures are drawn, there is great difficulty in identi-
fying the species represented ; but two, if not three, distinct ones
are certainly shown, all being included under the general name
of Nais digitata. Rosel’s species is recognized as distinct,
and named “the eyeless Naid with pronged tail,” the other
being called “the blind Naid with fingered tail.” The distinc-
tion thus formulated is too slender to enable me to decide
as to the specific differences; and in view of the impossi-
bility of ascertaining the particular species to which Miller
intended to apply the name digitata, there seems to be no

* The numbers in brackets refer to the Bibliography, p. 106.

_i i.

TT ———

NATURAL HISTORY OF THE GENUS DERO. 93

better course open than to entirely reject it. Up to the present
time the name is occasionally used on the Continent; but as the
worm designated may be any one of three species at least, and as
no clear definite description has yet been given of any of these in
particular under the name in question, this cannot be used as an
argument against the course proposed.

In acknowledgment of Miiller’s work, I have attached his own
name to the last new species discovered.

The name by which the genus is now known was bestowed upon
it by Oken (20) in 1815, and the species described by Rosel dis-
tinguished as Dero furcata, Miiller’s appellation being retained
for the other species. This was apparently the extent of Oken’s
knowledge of the genus, as his figure was taken from Miller,
and his diagnosis is as scanty as possible.

Dutrochet (9) rediscovered the genus, and being ignorant of
the work of earlier writers, renamed it Xantho, remarking that it
appeared either not to have been observed before or to have
been confounded with Mais. He also described the branchial
funnel as an organ of progression and retrogression and of pre-
hension; the branchial processes being spoken of as veritable
non-articulated limbs, resembling the feet of cephalopodous
Mollusca and the arms of the polyps! The number of branchial
processes is given as ten in one species and six in another,
to which the names Xantho decapoda and X. hexapoda are respec-
tively given. As the figures which accompanied the original
paper appear to have been lost, any attempt to identify Dutro-
chet’s species must be mere guesswork.

Blainville (1) gave a somewhat unintelligible account of two
species, named Nais digitata and NV. decapoda. The former is
credited with the possession of six pairs of fleshy lobes, the
latter with five, reference being also made to a drawing in the
‘Encyclopédie Méthodique’ of a species with no less than eight
pairs. Having had an opportunity of examining the latter work,
I found that the plate in question is merely a replica of Miiller’s
(18) ; while the peculiarly unmethodical character of the arrange-
ment of the text renders any search for the article referring to
the figures hopeless. However, as none of Miller’s figures show
more than four pairs of lobes, Blainville has evidently interpreted
them wrongly.

Blainville (2) in a second article in the ‘ Dictionnaire des Sciences
natureiles’ and a writer in ‘ Rees’s Cyclopedia’ (23) quoted from

LINN. JOURN.—ZOOLOGY, VOL. XX. 8

94, MR. E. Ce BOUSFIELD ON THE

Fabricius the description of a worm which he described as Nais
quadricuspida, which appears to have some affinity with the genus
Dero. Apparently this species has not since been observed,
though from the description given by Fabricius (10) it seems
sufficiently well marked *.

Gervais (11) reclassified the Naidide, and renamed the genus
Dero; Uronais being the designation applied to it, apparently
from the peculiarity of the tail. He appears to have been the
first to remark that Miiller’s figures evidently represented two
distinct species. In common with some other writers, he
included the Nats barbata of Miiller, in spite of the fact that the
latter is described as having a simple truncated tail.

Girsted (19) gave only a very short account of the genus,
calling it Proto (Oken).

Grube (12), whilst contributing little or nothing to our know-
ledge of the genus, made some interesting remarks on its syste-
matic position, and referred to two marine forms, described by
Dujardin (8) and Dugeés (7) respectively, which he regarded as
allied to Dero. Having carefully compared the papers referred
to by Grube, I am unable to agree with his conclusion. Grube
also remarked on the confusion in the nomenclature, professing
himself unable to decide the question of the origin of the name
Protot. It should be remarked that Grube does not mention
having seen Dero.

Bosc (4) mentioned, under the name of Nais auricularis, a worm
found in Carolina with a tail formed by a large tubercle, in the
middle of which is the anus,—a description which would well
apply to a Dero in a contracted state.

Pennant (21), Shaw (27), Stewart (28), and Turton (80) gave
descriptions of Nais digitata which afford no assistance in iden-
tifying the form; and the same may be said of a long paper by
Houghton (13), who, baving found the genus in England, instead

* “Nais verrucis lateralibus bifidis setosis cirris abdominalibus et cauda
quadrifida.”

+ The origin of this name remains unknown. (CErsted attributed it to Oken,
in whose work no trace of it is to be found; nor is it Jikely that he would have
given to one genus two different names. Another writer, in spite of Grube’s
remark, attributed the name to him. The only feasible explanation appears
to be that some unknown writer, between the time of Gervais and Céirsted
(i. €. 1838 and 1848), originated it in a paper which has been lost. ‘The ‘ Nomen-
clator Zoologicus’ of Agassiz gives both names, and attributes both to Oken, but
evidently wrongly. The etymology is there given as dépw, cutem exuo.

ee ee

NATURAL HISTORY OF THE GENUS DERO, 95

of observing it for himself, did little more than copy Miller’s
description. Johnston (14) merely mentions the genus to
throw doubt on its right to a position in the British fauna
at all.

In 1855, for the first time since Miiller noticed the genus
under consideration, we meet with an attempt to give something
hike an exact account of a new species. In that year a paper
appeared from the pen of D’Udekem (6), which must be regarded
as the starting-point of all modern work on the subject. Up to
that time Dero digitata and D. furcata (the latter of which had
almost been lost sight of) had held the field alone; but D’Udekem’s
contribution contained a description of a new species, Dero
obtusa, clearly expressed, and accompanied by a figure which
rendered it easily recognizable.

In 1872 Perrier (22) published a very interesting and exhaustive
account of a species which he identified with that described by
D’Udekem; and the fact that he was wrong on this point in no
way detracts from the value of his observations, whilst his figures
are almost all that could be desired. In acknowledgment of
Perrier’s work, I have named the species which he studied Dero
Perrieri, it being new.

Prof. Leidy (15), in a paper of which he has kindly sent me
a reprint, describes two worms, one of which he calls Dero limosa
and the other Auwlophorus vagus. The former appears to be
identical with a species found abundantly round London; the
latter is considered by Prof. Leidy to be identical with that
described by Rosel (loc. cit.), the Dero furcata of Oken; and
apparently Prof. Leidy’s distinction is based upon the fact that
he found the latter free.

Semper (26) has described two species under new names, his
Dero philippinensis being apparently the same as Dero limosa just
referred to, whilst Dero Rodriquezii is undoubtedly the same as
the Aulophorus vagus of Leidy, Dero furcata of Oken, and Dero
palpigera of Grebincky.

Tauber’s work (29) contributes nothing to our knowledge of
the genus, being merely a catalogue of Danish Annulata. He
remarked that the genus is rare in Denmark.

The magnificent work of the Bohemian Professor Vejdovsky
(81) contains a full list of names and synonyms of the species of
Dero known up to the date of bis publication. Unfortunately
he observed only a very few examples of the genus, the species

96 MR. E. C. BOUSFIELD ON THE

being one which he identifies with Dero digitata. As there is,
however, an absence of anything like a diagnostic description of
the species in question, the identity of Dero digitata is still, and
probably will remain, unsettled.

The last contribution to the literature of the subject is the
abstract of a paper read by me at the Aberdeen Meeting of the
British Association (5). The conclusions therein arrived at hold
good only in so far as they are corroborated by the present
communication. |

Habits, and Methods of Observation.

The species of the genus Dero, with the doubtful exception of
the form described by Fabricius in 1778, are all freshwater
dwellers. They pass their lives entirely below the water-level
in tubes which they secrete, and into the composition of which
foreign matter does not enter. The tubes are as a rule either
constructed in the mud, or along the stems of aquatic plants, or
even within decayed stalks.

To this method of life is probably owing the small amount of
attention which the genus has attracted. If, however, these
worms be present in any locality, they may be easily detected as
follows :—

A portion of the mud, within an inch of its surface, is placed in
a bottle to about the depth of half an inch and water added. After
the mud has subsided the bottle is allowed to stand for about 12
hours, when, if any samples of Dero be present, they will have
constructed their tubes in contact with the glass, either in the mud
or on the sides of the bottle, and may be removed for examination
by pressing on the end from which the head protrudes (as the tail
is very easily injured) with a camel-hair pencil. Under this
treatment the worm backs rapidly out of the tube, and may be
readily secured with a pipette. The species of the genus Dero
are sociable in their habits, as many as half a dozen tubes being
often placed side by side, and it is quite the exception to find
single ones.

Having secured the worm, the question arises how best to
examine it. It is almost impossible to determine the species of
any given example when ordinary methods, such as the com-
pressorium or the live-trough, are alone employed. In the first
case the pressure, even if slight, prevents that full expansion of
the brancbial area which is absolutely necessary for exact obser-
vation ; whilst in the second case the restlessness of the worm is

NATURAL HISTORY OF THE GENUS DERO. 97

so great that it can only be kept in the field of a power too low
for needful details to be made out.

For general observation the method which I have found
most suitable is to transfer the worms to a live-trough, with
a sufficient depth of mud for them to form their tubes
(about z inch), when they may be observed under perfectly
natural conditions; the hinder end of the worm, carrying the
respiratory apparatus, being kept protruded upward, whilst the
head is occupied below in ingesting the mud which forms the
food of these creatures. If the tubes have been formed amid
vegetable débris, the best plan is to secure a portion in the
compressorium under slight pressure, or m a small zoophyte-
trough, when powers as high as Zeiss’s B B or an English §-in.
may easily be employed. To make out the histological details,
nothing is better than the cotton-wool trap used for wandering
Rotifera; with this and a judiciously regulated pressure, a ;),-in.
objective may be safely used. The form of compressorium
adopted by me is that known as Beck’s parallel compressorium,
and I have found nothing to equal it for the facilities it offers of
increasing or diminishing pressure without removal, and of
viewing an object on both sides.

General Characters.

In general outline the species of Dero closely resemble their
relatives of the genus WVais. The following marked differences,
however, obtain :—

1. They are destitute of eyes.

2. They are furnished with decidedly red blood.

3. The perivisceral fluid is devoid of corpuscles.

4. They inhabit fixed tubes.

5. They possess a highly specialized respiratory organ on the
last segment of the body.

The general form of the body is more or less cylindrical, the
head being obtusely pointed. The thickness gradually increases
from the head for about two fifths of the length of the worm, after
which it diminishes gradually again, being narrowest in the last
segment but one.

As in Nais, the mouth-segment is destitute of organs of
motion, whilst the four following have them only on the ventral
surface *,

* Except Dero furcata, of which see description.

98 MR. E. C. BOUSFIELD ON THE

These organs are known as sete, their form being more or less
that shown in Pl. IV. fig. 10 and Pl. V. fig. 16, curved like a
long f, with a central shoulder and bifurcate outer end. Each of
the bristle-bundles in the first four ventral pairs contains from four
to six such sete, in length considerably exceeding those of the body
generally, though there is but little difference in the general form.

In the dorsal bristle-bundles, the first pair of which is found
in the sixth segment, bristles of two forms are found. The
first and most evident are tapering, projecting throngh the
epidermis to a distance about equal to the diameter of the
worm, and of these one is found in each bundle. The second
are short, stout, notched at the outer extremity, and barely
penetrate the skin. Notwithstanding their inferior length they
are always the first developed, and if there should by chance be
two tapering bristles in a bundle, each of them is accompanined
by its own shorter one.

The developmental relation between these two forms was
pointed out by Perrier, and is additionally proved by the fact that
in the posterior segments the tapering bristles are altogether
wanting, the short ones alone remaining; these at length are
reduced to mere points in the last few segments, and finally
disappear.

Illustrations of the various forms of sete will be found in the
figures already referred to.

The integument of the worm is smooth, and shows but few
palpocils, except at the head and tail, and even there not many ;
they are connected with pyriform cells in the epidermis in the
manner described by me in a previous paper *.

The digestive canal conforms closely to the Naid type in its
exsertile proboscis, its pharynx beset with mucous glands, and
its long pharynx dilated at one point to form a gastric enlarge-
ment, passing on into an intestine, capacious, closely beset with
the so-called hepatic cells, and having a very strong inward
ciliary current for a great part of its length. The segmental
organs, which begin in the sixth segment, are essentially simple
convoluted tubes with very narrow Jwmen, the dilated inner
extremity being thickly clothed with fine short cilia. The
external condition of these organs varies greatly. Frequently —
they are as described above, without any appendages. In other
instances in the same species, at the same period of the year,

* “On Slavina and Ophidonais,” Journ. Linn. Soc., Zool. vol. xix. p. 265, 1886.

S66 Oe eae as ee:

NATURAL HISTORY OF THE GENUS DERO. 99

they are embedded, except for a short distance at each end, in a
mass of large, clear, spheroidal cells. When these masses are
not found on the segmental organs, they may occur on the septa
between the segments, or they may be absent altogether. Ihave,
however, been unable to discover the conditions which govern
their appearance.

It has been stated that the internal end of the segmental
organs is a ciliated funnel; they end externally in a somewhat
dilated portion, which communicates freely with the surrounding
element. Their function appears to be purely mechanical—
that of preventing undue distention of the body by the
fluid which passes through the wall of the intestine, and is
doubtless charged with effete material from the blood-vessels
which run in contact with it. Accordingly I find that, con-
trary to the general accepted opinion, the lumen of the tube of
the segmental organ is not ciliated, but that along one side of it
is attached a membrane whose undulations have the effect of
driving the coutained fluid in an outward direction. During full
activity it is not easy to see the edge of the membrane, though
the character of the movement even then is such as to suggest
some action quite different from that of cilia, and much more
nearly resembling that seen in the vibratile tags of the rotifers.

When vitality is at a low ebb, it is by no means difficult, with
a power of about 500 diameters and suitable illumination, to see
the edge of the membrane. , The same conditions exist in the
case of other Oligocheta, and indeed it is not easy to see how
cilia could act to advantage in a tube of such extremely small
calibre, nor am I aware of any instance in other forms of animal
life where such is the case *.

The nervous system of Dero is excessively difficult to make
out, but appears to closely resemble that of avs in its arrange-
ment. Perrier’s account of the species which he observed
stops short at this point, his description being very meagre, and
in the few details given there is nothing to indicate any special
arrangement.

* The observations which led to the above conclusions were made on
Tubifex, Dero, Nais, Stylaria, Chetogaster, and Afolosoma. In the case of
Chetogaster only was there any difference. In the latter genus I have never
seen any movement whatever in the tubular portion of the segmental organs, but
on one occasion observed in connection therewith an organ exactly resembling
the vibratile tags already referred to.

100 MR. E. C. BOUSFIELD ON THE

Branchial Area and Cireulatory System.

The branchial apparatus is by far the most important character
of Dero. Though it is found in a modified form in the Awlophorus
of Schmarda (25), yet in its full development it is present only in
the genus under notice. In all the Oligocheta a strong inward
current is visible in the hinder part of the intestine, which no
doubt subserves a respiratory purpose, as it commences at the
point where the arterial system receives the blood from the
venous; and at this point, if nowhere else, both vessels run in
close contact with, or are embedded in, the intestinal wall, so that
the most favourable conditions for interchange are combined. In
those forms which live and move free in the water, this is
doubtless sufficient provision for their needs; but in the case
of Dero, which, though not fixed, is yet stationary, a special pro-
vision seems required, such as is found in the branchial area.

This is essentially, and in its simplest aspect, an opening-out
of the hinder part of the intestine, supported by a layer of
epidermis, the space between the two being occupied by muscular
elements and blood-vessels. Figs. 4 and 5 in Pl. LV. (the former
taken from Perrier’s monograph, the latter from life) will show
the generalarrangement. In fig. 5, which gives the area in sec-
tion, the relative proportions between its two constituents are
shown by the shading, the dark portion being integumental,
whilst the light shading indicates the continuation of the mucous
membrane of the intestine, which supplies the respiratory
element. The form of the area differs in different species; but
in all cases a number of branchial processes arise from the floor
of it, their number being four and ne more. This may seem
a surprisiug statement, in view of the considerably greater
number, e¢ght pairs according to one writer already referred to,
which have previously been described. Reference has already
been made to the difficulty of determining the species of any
given example of the genus, and similar difficulties, not overcome,
have been the cause of the remarkable divergencies and dis-
crepancies to be found in the descriptions heretofore given.
These have obtained down to the latest period: thus both Semper
and Leidy have described appearances which can easily be re-
produced, but which are not by any means normal. If an apology
be needed for such an assertion, I hope it will be found in the
fact that I have devoted myself almost exclusively to the study

NATURAL HISTORY OF THE GENUS DERO. 101

of this genus for nearly two years, have examined hundreds of
examples of its various species, and have done my utmost to make
sure of every fact which I advance. It is true that in three
species supplementary branchie are present, but these are
much smaller than the primary ones, and are always placed at
the angles of the dorsal lip of the area. Where more than three
pairs are attributed to any species, the incurved margins of the
area, which are ciliated on both surfaces, have been mistaken
for additional branchiz.

The integumental and mucous layers of the area are not
equally extensive ; the mucous layer is wanting (Pl. LV. fig. 5) at
the tip, and encroaches on the margin of the integumental layer
at every other part. The branchiz are simple elevations of the
mucous layer, containing loops of blood-vessels one in each, and
are lined by a layer of stellate muscle-cells continuous with the
layer which underlies the mucous membrane of all the other
portions of the area. This muscular network is capable of great
expansion and contraction, and in the latter condition fully
realizes Bose’s description of Nats auricularis. The area is
extremely sensitive, the slightest jar causing contraction, and,
what is somewhat remarkable, it appears to be endowed with
sensibility to actinic light *.

In some species, in addition to the muscular network of stel-
late cells already spoken of, an arrangement of radiating cells,
spindle-shaped, with forked extremities and a central nucleus, is
found connecting the dorsal lip of the branchial area with the
intestine beneath it.

In most cases the branchial area is concave in full dilatation ;
but in Dero Mulleri it is capable of such expansion as to become
altogether convex, and to stand at right angles to the axis of the
body. ‘The same is the case toa less extent in D. latissima.

The chief blood-vessels are, as usual, two in number, a dorsal
(arterial) and a ventral (venous). ‘The dorsal vessel in Dero can

* I have several times tried to photograph the tail of Dero, but hitherto
with small success, the ignition of the magnesium-ribbon employed as
the source of light causing contraction in nearly every case. The same phe-
nomenon occurs in Hydra. In order to overcome this difficulty a drop-shutter
was placed between the source of the light and the object, but the result was the
same; and the marvellous rapidity of the contraction was shown by the fact

that, although fully expanded up to the instant of exposure, the photograph i in
every case showed the polyp in a contracted condition.

102 MR. E. C. BOUSFIELD ON THE

only be so called for convenience; since for the greater part of
its length it runs along the side or lower part of the intestine
embedded in its wall, only becoming free when it reaches the
junction of the latter with the stomach. From this point it is
connected by a varying number of contractile loops with the
ventral vessel, and terminates exactly as in Waits. In the bran-
chial area the circulation between the dorsal and ventral vessels
is carried on as follows:—The abdominal vessel, running back-
ward undivided to the boundary of the mucous layer of the area,
divides into two branches which run right and left round its
margin. Hach of these main branches gives off at least three
secondary ones, one to each of the branchia of its own side, which
runs to its summit and down again without breaking up into a
capillary network ; whilst the third, also undivided, runs obliquely
across the area, the continuation of the main branch running on
round the margin. AJl these branches again unite to form a
common trunk, which joins with a similar one from the opposite
side to form the commencement of the dorsal vessel.

The specialization of the termination of the hinder portion of
the intestine does not appear to have any effect in abrogating the
general respiratory function of that tube; doubtless an advantage
~ to these worms, which spend so large a portion of their lives in
crawling in: the mud in search of food. It should be remarked
that the tube is much too narrow to admit of the branchial area
being drawn into it when expanded; nor is the tube wide enough
to give room for the bristles, the latter being always much curved
when within it; and though the worm is able, as a rule, to turn
round with facility, yet it sometimes happens that it becomes
fixed in the tube in so doing, and is quite unable to extricate
itself.

Distribution of Species.

With the exception of Dero Milleri, received from Mr. Bolton,
of Birmingham, I have found all the known forms within a short
distance of London. The richest hunting-grounds are without
doubt the tanks at Kew and at the Royal Botanic Society’s
Gardens, Regent’s Park. The latter locality has furnished a
species, the only one whose indigenous character is doubtful, viz.
the beautiful D. furcata, the finding of which and the difficulty .
of obtaining any reliable information about it, first led the writer
to undertake the study of this genus. Here also he first found

ee , ne

NATURAL HISTORY OF THE GENUS DERO. 103

D. latissima, as also in a pond between Sutton and Redhill on
the high road, and later D. limosa. The latter is especially
abundant in the Lily-tank at Kew. D. obtusa and D. Perrieri
are to be found in the large pond on Wandsworth Common; and
both species, as well as D. latissima and the rare D. acuta, have
been found in the pond at Greenwich Park. The last-named
species was also found for the first time by my wife in the
Crystal-Palace basin.

Abroad the only species known to have been found are D.
furcata, D. limosa, D. obtusa (Belgium only), and the doubtful
D. digitata. Of these, the first two are widely distributed,
ranging from the United States to South Russia and the Phi-
lippine Islands. It will be seen, therefore, that at present the
British Isles are apparently by far the richest in species of any
known locality. Others doubtless remain to be discovered ;
and to this end I am anxious to obtain samples of mud in a
moist state from various localities abroad *; and South America
especially should prove fertile in this direction. Much remains
to be done before our knowledge of the minute Oligocheta can
be considered even fairly complete ; and it is quite possible that
systematic observations in tropical and subtropical regions would
reveal forms still more interesting and remarkable than those
included in the genus Dero.

My thanks are due to Mr. T. Bolton, whose kindness in sup-
plying me with specimens has resulted in the discovery of one
new species which has not yet been found elsewhere.

The following is a systematic account of the various species at
present known.

Family Narnomorpna, Vejdovsky.
Genus Dero, Oken.
Proto.—Xantho, Dutrochet.—Nais digitata, Willer.

Char. Minute Annelids of from 40 to 100 segments, red-
blooded, eye-less, inhabiting tubes secreted from the body, without
aggregation of foreign material. The last segment bears an expan-
sion from which arise four processes. Perivisceral fluid free from

* I shall be much obliged for samples from any freshwater lake, pond, or
tank, or even slow streamabroad. ‘The best time for collecting would probably
be before the rainy season in such localities as have one; in those which have

not, probably the end of summer would be best. The samples may be sent to
me, care of the Linnean Society.

104 MR. E. C. BOUSFIELD ON THE

corpuscles. First five segments destitute of dorsal sete; thesixth —

and following having both dorsal and ventral sete, the latter
hooked, the former straight, and one forked bristle in each bundle.
Contractile loops in the sixth and some following segments, one
in each.

a. Without secondary branchie.

1. Dero tatisstma, Bousfield (5). Pl. LV. fig. 8.

Segments 80-40. Contractile loops 4. The branchial area
with entire margin, well developed, width in full expansion
exceeding length, capable of complete eversion, then becoming
convex. Branchie long, well developed, plano-cylindrical.

2. Dero Perrtert, Bousfield. Pl. IV. figs. 4-7.

Dero obtusa, Perrier (22).

Segments 25-35. Contractile loops 3-5. Branchial area with
entire margin, trefoil-shaped in full expansion. Branchial pro-
cesses well developed, cylindrical, the posterior pair longer than
the anterior.

3. Dero oprusa, D’ Udekem (6). PI. III. figs. 1-3.

Segments 45-50. Contractile loops usually 4 (4-6). The
branchial area somewhat irregular, and apt to be unsymmetrical.
Branchie rather short, foliate; the posterior margin nearly
straight, the anterior curving outward at the middle. A dis-
tinct dorsal lip, divided from the alar portions of the area by
a deep groove on each side.

This species shows the first trace of the modification which
leads to the formation of supplementary branchial processes in
the distinct demarcation of the dorsal lip, at the angles of which
they are borne by those species which possess them.

4. Dero Miruert, sp. nov. Pl. IV. figs. 9, 10.

Segments 70-95. Contractile loops usually 7 (6-8). The
branchial area with separate dorsal lip. Branchial processes
oblong, quadrangular, the attached border the longest, superior
angles rounded.

The largest of all known species. Furnished to the writer by
Mr. Bolton. Chains of two zooids frequently consist of 1830-140
seyments, and measure nearly an inch in length. Branchial area
disproportionately small.

—_—-

NATURAL HISTORY OF THE GENUS DERO. 105

3. With secondary branchie.

5. Duro t1mosa, Letdy (15). Pl. V. figs. 11-16.

Dero philippinensis, Semper (26).

Segments 55-60. Contractile loops 5, the last much the
smallest. The branchial area with dorsal lip, each angle bearing
a secondary branchia, short, cylindrical, and containing a loop of
blood-vessels. Branchial processes as in D. obtusa, but longer
and narrower. |

The foregoing species appears to me, for reasons already
stated, to be identical with Prof. Leidy’s; and there is no
difference which cannot be explained on the grounds laid down
(p. 100). Should the species under notice hereafter prove to
be distinct from Prof. Leidy’s, it will stand as new, unless
indeed it be the same as Prof. Semper’s D. philippinensis, as on
the same considerations seems probable ; though the latter may
possibly be identical with the species next to be described, a point
which cannot be settled in the absence of a figure of Prof. Semper’s
worm.

6. Drro acuta, Bousfield (5).

Characters nearly as in D. limosa; but the branchie much
longer, obovate in form, with pointed apices; length about 4 or
5 times the greatest breadth.

Only two examples of this species have come under my notice
from the localities already mentioned (p. 103). A careful search
during the present season has been unsuccessful; so that no
figure can be given; but there will be no difficulty in recognizing
the worm from the description above.

y. With secondary branchie and palpi.

7. Dero Furcata, Oken (20). PI. V. figs. 17, 18.

Das geschmeidige Wasserschlinglein mit zwey Gabelspizen,
Résel (24)—Dero palpigera, Grebincky.—Dero Rodriguezii,
Semper (26).

Segments 35. The branchial area more funnel-shaped than in
the other forms; the dorsal lip not divided off, but bearing sup-
plementary branchie at its junction with the ale. The integu-
mental layer at the tip of the branchial area prolonged into two
palpi of varying length, non-ciliated, extra-vascular. Branchiw
nearly cylindrical ; contractile loops tive. The first dorsal bristle-
bundles in the fifth segment.

106

MR. E. C. BOUSFIELD ON THE

The most beautiful of all the species of Dero; but somewhat »
anomalous in the position of the first dorsal bristles and in
the possession of palpi. Only found among the fibrous bundles
of the dead stems of Cyperus (in the Royal Botanic Gardens),
which must be pulled apart to obtain the worms.

Note.—The above descriptions all apply to the asexual forms

only.

—
SiO COMI Oy Ge to

Pig. 1,

A description of the sexual forms is reserved.

BIBLIOGRAPHY.

. BLAINVILLE.—Dict. des Sci. Nat. t. xxxiv. p. 131.
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. BousFiELD.—Report Brit. Assoc. Adv. Sci. 1885, p. 1097.
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. Stewart.— Elements Brit. Zool. p. 391.

. TauBeR.—Annulata Danica, i. p. 79.

. Turron.—British Fauna, p. 137.

. Vespovsky.—Syst. u. Morph. der Oligoch. pp. 27 & 117.

DESCRIPTION OF THE PLATES.
Puate III.

Dero obtusa. The whole worm, enlarged from a photograph of the
living object.

A dorsal view of the branchial} area and processes of D, obtusa, drawn
from life. x 75.

. The same. A copy of D’Udekem’s original figure.

a ne te aaa ae TS

A.0.Walker.

Crowther lith.

XANTHO SCABERRIMUS,G@.

Linn. Soc. Journ. Zoou. Vou. XX. Px. 7.

Mintern imp.

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OF CRUSTACEA FROM SINGAPORE, 1

Carapace transversely ovate, very slightly convex, smooth, and
shining ; regions strongly defined, especially behind ; edge fringed
with long hairs. Front slightly prominent in the middle. Che-
lipedes very large, equal, densely covered on the upperside with
long hairs, smooth and shining white on the underside; wrist
shorter than the hand and hollowed out on the inner side. Next
three pairs of legs short and weak, covered with long hair on the
upperside ; dactylus very small and curved; merus strong, as
long as the two following joints. Last pair of legs long, slender,
and almost naked; the last joint polished and horn-coloured,
with a brush of hairs at the extremity. Length ;4, in. (=8 mil-
lim.). Width =; in. (=10 millim.).

DESCRIPTION OF THE PLATES.

Puate VI.
Fig. 1. Maia Miersi, ¢.
2. Ditto, underside of orbital region.
3. Ditto, abdomen.
4. Doclea tetraptera, 3.
5. Ditto, underside of orbital region.
6. Ditto, left chelipede.
7. Ditto, abdomen.
8, 8*. Ditto, hairs (magnified).

Pruate V)I.
Xantho scaberrimus, 3.

Prate VIII.
. Porcellanella picta.
. Ditto, left chelipede.
. Onychomorpha lamelligera.
. Goniosoma inequale.
. Petrolisthes corallicola, var., chelipede.
. Diogenes avarus, cephalothorax.
. Ditto, chelipede.

Fig.

eV Woe

JID oO Pp

Puate IX.
. Polyonyx cometes.
. Front view of ditto.
. Right chelipede of ditto.
. Caphyra Archeri.
. Chelipede of ditto.
. Lyphlocarcinus villosus.
. Left chelipede of ditto.
. Abdomen of ditto, ¢.
. Sesarma Bocourti, chelipede of 2.

LINN. JOURN.—ZOOLOGY, VOL. Xx. 11

Fig.

COWIHopod oe

118 SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS.

Observations on Ants, Bees, and Wasps.—Part XI.
By Sir Joun Luszock, Bart., M.P., D.C.0., LL.D., F.R.S.

[Read 1st December, 1887.]

On THE RELATION BETWEEN Formica sanguinea AND
ITs SLAVES.

Ip is well known that Polyergus rufescens is entirely depend-
ent on its slaves. Huber long ago found that this ant will
starve in the midst of plenty, and will not even put food into its
own mouth. I have shown that isolated specimens will live for
months if they are allowed a slave for an hour or so every two or
three days to clean and feed them. It is said, on the contrary,
that our only slave-making species, Formica sanguinea, can
manage without slaves. Indeed, it appears that nests are
sometimes found in which there are mistresses alone, entirely
without slaves. Forel thinks that he has observed in such
nests generally a larger proportion of small individuals than
in nests which possess slaves. This would be interesting as
tending to show that in such nests the young are less well
nourished than when they have slaves to attend upon them.

The question remains, of what advantage are the slaves to the
F. sanguinea? Forel says, I do not quite understand why, “Je
ne veux pas trop rechercher le motif qui pousse les sanguinea a
se faire presque toujours des esclaves.” ‘‘ Peut-étre,” he adds, “‘le
sentiment de leur force, et le désir de travailler moins, pour faire
plus a leur aise la chasse aux Lasius flavus et L. niger, est-il le
mobile qui les pousse a cet acte. Celui-ci leur serait peu 4 peu
devenu plus ou moins instinctif puisqu’il était avantageux a
la conservation de leur espéce.”’ This suggestion seems very
probabie, and may be partially correct ; it is not, however,
I think, a complete explanation. I have had under observation
several nests of I. sanguinea. One of them I owe to the kind-
ness of M. Forel himself, who sent it to me in June 1882.
There was no queen, and, though the nest was very healthy, of
course the numbers gradually diminished. At the beginning
of January 1886 the last slave died, and there then remained
only about 50 F. sanguinea. Under these circumstances the
F. sanguinea began to die off rapidly ; by the middle of the year
only six remained alive, and these, no doubt, would not have
survived long. On July lst I got some pupe of &. fusca and
placed them outside the nest. The sanguineas soon discovered

SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS. 119

them, carried them into the nest, and from that day until De-

cember 1887, more than six months, there was only one other
death. [Two of the & sanguinea are still (August 1888) alive. ]
Although then it may be true, as to which I express no opinion,
that there are nests of F. sanguinea without slaves, still this
observation seems to indicate that the slaves perform some im-
portant function in the economy of the nest. It still remains
to be determined in what exactly this function consists.

Ant-GUuESTS.

Dr. Wasmann has recently published * an interesting memoir
on certain of these ‘‘ Ant-guests.”? His observations relate ex-
clusively to some of the beetles which live with ants.

He confirms V. Hagen’s statement that the specimens of
Atemeles emarginatus which live with Myrmica levinodis, a
yellow ant, are paler in colour than those which share the nests
of the black Formica fusca. He entirely confirms the statements
of previous observers that the Atemeles is actually fed by the
ants, who also clean them just as they do their own fellows. The
Atemeles also, on their part, perform the same kind offices
for the ants. He also repeatedly saw the ants licking the
bunches of golden hairs on the abdomen of the Atemeles.

The Atemeles has adopted very closely the habits of the ants
with which it lives. They pair, moreover, in the nests of the
ants. Still, they are not entirely dependent on their hosts, like
some of the other ant-guests, but are able to feed themselves.
Indeed the Myrmicas seem to drive them out of the nest to-
wards the beginning of May. Dr. Wasmann is disposed to
attribute this to the anxiety of the ants for their young. In
Myrmica the pup are naked, and he thinks the ants are afraid
that the Atemeles would be unable to resist the temptation of
eating them. In support of this suggestion, he observes that in
the nests of Formica sanguinea, whose pupe spin a silken cocoon
and are therefore protected, he has found Atemeles as late as the
end of June. He has not been able to satisfy himself whether
the larve of Atemeles are brought up in the ants’ nest or not;
but inasmuch as while the Atemeles are far from rare, he has
only found among them a single Jarva which could belong to the
species, and even this was not certainly identified, it seems

probable that the larval stage is passed elsewhere.

* Deutsche Entom. Zeitschrift, 1886, p. 49.
LI*

120 SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS.

Lomechusa strumosa has been recorded from the nests of For-:
mica sanguinea, Myrmica rubra, Formica congerens, and F. rufa;
but Dr. Wasmann, like V. Hagen and Forel, has never found it
except with F/ sanguinea. It is fed by the ants in the same man-
ner as Atemeles, and has very similar bundles of golden hairs on
the abdomen, which are licked by the ants like those of Atemeles.
While, however, the ants seem to communicate, with the Atemeles
mainly by means of the antenne, in the case of Lomechusa the
parts of the mouth are brought more into play. He has found
Lomechusa in the ants’ nests up to the end of June.

Dinarda dentata is a still more frequent inmate in the nests of
I. sanguinea, but plays a very different part. The ants seem in-
different to them, and when they take any notice it is of an
unfriendly character. Nor can this be wondered at, for according
to M. Wasmann the principal food of the Dinardas consists of
any ant which may chance to die, or any other weak insect
which falls in their way. The ants seem thoroughly to distrust
them, and it is curious that they should be tolerated. Grimm,
indeed, thought be had seen ants licking the Dinardas, as they
do Atemeles and Lomechusa. Wasmann, however, considers that
this was a mistake; at any rate he has never seen anything of the
kind. If an ant meets a Dinarda she either treats it with in-
difference or threatens the beetle with her jaws, and the Dinarda
then raises its abdomen, which appears always to drive away
the ant. It is possible that the Dinarda has the power of pro-
ducing an odour distressing to the ants, or perhaps they eject
poison like the Formicas themselves. They seem always to re-
main in the nests of the ants, and pass through their transform-
ation in them. Sormica sanguinea, like #. rufa and F. congerens,
changes its nest periodically twice a year; such, at least, is said
to be the case on the continent ; I am not aware whether the
same habit has been observed in this country. The summer
nests are looser and opener, the winter ones lower and more
compressed. In their migration from one nest to the other,
which are occasionally at some distance apart, the ants are
accompanied by the Dinardas. On one occasion, when the ants
were flitting, Dr. Wasmann in 20 minutes captured among
them thirteen specimens of Dinarda; while under other circum-
stances he never saw one outside the nest. |

Heterius ferrugineus, belonging to a totally different family of
beetles, the Histeridx, and which inhabits the nests of Polyergus |

SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS. 121

rufescens, Formica sanguinea, F. pressilabris, F. fusca, F. rufi-
barbis, F. rufa, F. exsecta, Lasius niger, and Tapinoma erraticum,
appears to agree in its habits with Dinarda, and to devour dead
and wounded ants, as also do the Myrmedonie.

Dr. Wasmann confirms entirely my observations, in opposition
to the statements of Lespés, that while ants are deadly enemies
to those of other nests, even of the same species, the domestic
animals, on the contrary, may be transferred from one nest to
another and are not attacked.

He justly observes that, no doubt, many interesting dis-
coveries are in store for us as to the relations between ants and
their guests. The marvellous and grotesque antenne of the
Paussidez will doubtless, one of these days, tell a wonderful story
to sume patient and fortunate observer.

Mimicry amone AnrTs.

Prof. C. Emery has published in the Bull. della Soc. Ent.
Italiana, 1886, a short, but interesting note on the habits of
Camponotus lateralis. Of this species there are two varieties :
one black, like its nearest allies; the other red, with the abdomen
and part of the thorax black. They live in small colonies, and
make expeditions up trees to collect honey-dew from the Aphides.
The black type (C. foveolatus, Mayr, C. ebeninus, Emery) some-
times goes in troops, but generally a few join the troops of other
black ants, such as Formica gagates and Camponotus ethiops.
Prof. Emery suggests that, their numbers being small and their
sight not very good, they find it convenient to accompany other
ants which live in larger communities, and that they perhaps
escape detection from the similarity of colour.

This suggestion derives some support from the fact that the
red variety accompanies in a similar manner the troops of
Oremastogaster scutellaris, which is red and black, and at first:
sight curiously like the red variety of C. lateralis. Cremasto-
gaster scutellaris lives in immense communities among the pine
woods along the shores of the Mediterranean, and is, as I know
to my cost, a very pugnacious species. Prof. Emery suggests
that the black form of C. lateralis is the original type, resem-
bling as it does its nearest congeners ; and that the red variety
has the advantage, from its similarity to Cremastogaster scutel-
laris, of using that species as its guide and of sharing, undetected,
in the produce of its flocks and herds. Prof. Emery observes

122 SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS.

that he only suggests this explanation. The facts he mentions

are very interesting, and it is to be hoped that he will continue
his observations.

ON THE COLOUR-SENSE OF ANTS.

Prof. Graber * has published an interesting memoir on this
subject. He confirms my observations on ants and Daphnias, in
which I showed that they are sensitive to the ultra-violet rays,
by similar observations on earthworms, newts, &c. It is inter-
esting, moreover, that the species examined by him showed
themselves, like the ants, especially sensitive to the blue, violet,
and ultra-violet rays. Prof. Graber, however, states that he
differs from me, inasmuch as I attributed the sensitiveness to the
ultra-violet rays exclusively to vision ; that it is “‘ ausschliesslich
durch die Augen vermittelt.”” I would not, however, express that
opinion as applying absolutely to all animals, though it is, I
believe, true of ants, where the opacity of the chitine renders it
unlikely that the light would be perceived except by the medium
of the eyes and ocelli.

Graber has demonstrated in earthworms and newts, and Plateau
in certain Myriapodst, that these animals perceive the difference
between light and darkness by the general surface of the skin.
But more than this, Graber appears to have demonstrated that
earthworms and newts distinguish not only between light of
different intensities, but also between rays of different wave-
lengths, preferring red to blue or green and green to blue. He
found, moreover, as I did, that they are sensitive to the ultra-
violet rays. Harthworms, of course, have no eyes, but thinking
that the light might perhaps act directly on the cephalic ganglia,
Graber decapitated a certain number, and found that the light
still acted on them in the same manner, though the differences
were not so marked. He also covered over the eyes of newts,
and found that the same held good with them. Hence he con-
cludes that the general surface of the skin is sensitive to light.

These results are certainly curious and interesting; but,
even if we admit the absolute correctness of his deductions,
I do not see that they are in opposition to those at which
I had arrived. My main conclusions were that ants, Daphnias,

* «Fundamental Versuche iiber die Helligkeits- und Farben Empfindlich-
keit augenloser und geblendeter Thiere,” Sitz. Kais. Akad. der Wiss. Wien, 1883.
t+ Journ. de l’Anatomie et de la Physiologie, 1886, p. 431.

all

SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS. 123

&e., were able to perceive light of different wave-lengths,
and that their eyes were sensitive to the ultra-violet rays
much beyond our limits of vision. His observations do not in
any way controvert these deductions: indeed the argument by
which (p. 234) he endeavours to prove that the effect is due
to true light and not to warmth, presupposes that sensations
which can be felt by the general surface of the skin are still
more vividly perceived by the special organs of vision.

Prof. Graber’s observations have been followed up by
M. Forel*. He took fifteen specimens of Camponotus ligni-
perdus, which is a large species and, moreover, possesses the
advantage, for this purpose, of having no ocelli, and carefully
covered the eyes with opaque varnish. He then placed them in
a box with ten normal specimens of the same species (to which
he subsequently added five more), and covered over one half of
the box with cardboard and the other half with a layer of water.
In this way the one half of the box was darker than the other,
but the temperature of the two sides was approximately equal.
In four experiments the numbers were as follows :—

Under the cardboard. Under the water.
Hoodwinked Ants. Normal Ants. Hoodwinked Ants. Normal Ants.
3 9 12 1]
13 i 3 3
9 2 5 1
3 5 12 2
28 33 32 Yi

It will be seen that a very large majority of the normal ants in
every case went under the cardboard ; while it was practically
indifferent to the hoodwinked ants in which side of the box they
rested. Moreover, every time the water and the cardboard were
transposed, the normal ants were much excited and began run-
ning about to avoid the light, while the hoodwinked ants were
quite unaffected.

These experiences therefore proved that the varnish did, in
fact, render the ants temporarily blind, their instincts being in
other respects unaffected.

He then replaced the cardboard and water by a solution of

* Ree. Zool. Suisse, 1887.

124 SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS.

esculine, which is impervious to the ultra-violet rays, and a glass
of deep cobalt, which stopped most of the other rays but
permitted the ultra-violet to pass. The results then were :—

Under the Esculine. Under the Cobalt glass.
Hoodwinked Ants. Normal Ants. Hoodwinked Ants. Normal Ants.
11 8 3 q
i a 13 SI 2
9 12 D 3
5 13 9 2
10 12 } :
3 pul 12 3
12 13 3 1
61 82 40 15

Thus, then, a very large proportion of the normal ants preferred
to avoid the ultra-violet rays by going under the esculine. To
the varnished ants, on the contrary, it was indifferent whether
they were under the esculine or the cobalt. The slight prepond-
erance in favour of the esculine was probably partly due to having
started the experiments with a larger number of ants in the side
of the box then covered with esculine, and partly from the fact
that the hoodwinked ants would have a tendency to accompany
the others.

From these and other experiments M. Forel comes to the same
conclusion as I did, that the ants perceive the ultra-violet rays
with their eyes; and not, as suggested by Graber, by the skin
generally.

EXPERIMENTS with Platyarthrus.

In connection with this subject I may add that I do not at all
doubt the sensitiveness to light of eyeless animals. Jn experi-
menting on this subject I have always found that though the
Platyarthrus, which live with the ants, have no eyes, yet if part
of the nest be uncovered and part kept dark, they soon find
their way into the shaded part. It is, however, easy to imagine
that in unpigmented animals, whose skins are more or less semi-
transparent, the light might act directly on the nervous system,
even though it could not produce anything which could be called
vision.

SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS. 125

On THE FuNcTION oF THE Compound EyEs and OCELLI.

Forel agrees with Réaumur, Marcel de Serres, and Dugés, that
in insects which possess both ocelli and compound eyes the
ocelli may be covered over without materially affecting the move-
ments of the animals; while, on the contrary, if the compound
eyes are so treated, they behave just asinthe dark. For instance
Forel varnished over the compound eyes of some flies (Calli-
phoria vomitoria and Lucilia cesar), and found that if placed on
the ground they made no attempt to rise, while if thrown in the
air they flew first in one direction and then in another, striking
agaiust any object that came in their way, and being apparently
quite unable to guide themselves. They flew repeatedly against
a wall, falling to the ground and unable to alight against it as
they do so cleverly when they have their eyes to guide them.
Finally, they ended in flying away straight up into the air and
quite out of sight.

Johannes Miller inclined to the opinion that insects saw near
objects with their ocelli. Plateau satisfied himself that the move-
ments of insects are not affected by the ocelli being covered over,
and hence concluded that they are rudimentary organs. The com-
plexity of their structure, however, seems fatal to this conclusion.

Forel confesses that the use of the ocelli still remains an
enigma, but he is disposed to think that they enable their
possessors to see in comparative darkness. He observes* that
they are specially developed in insects which require to see both
in bright light and also in comparative obscurity. Aerial insects
do not generally require or possess ocelli.

Lebert expresses the opinion ¢ that in spiders some of their
eight eyes—those which are most convex and brightly coloured—
serve to see during daylight ; the others, flatter and colourless,
during the dusk. Pavesi has observed { that, while the species
of Nesticus possess normally eight eyes, in a cave-dwelling species
(Nesticus speluncarum) there are four only, the four middle eyes
being atrophied. This suggests that the four central eyes serve
specially in daylight.

SENSE OF SMELL.

In my previous memoirs I have recorded a few experiments
which convinced me that ants are gifted with a very highly
developed sense of smell, and that this resides in the antenne.

Ped. p, 181. +t Die Spinnen der Schweiz, p. 6.

+ “Sopra una nuova specie di Ragni appartenente alle collezioni del Museo
Civico di Genova,” Ann, Mus. Civ. 1878, p. 344,

126 SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS.

Forel, Graber, Lefebvre, Perris, and other recent writers have

come to the same conclusion, and there can, I think, be no rea-

sonable doubt that in very many insects the antenne serve as
organs of smell. At the same time it does not necessarily follow
that the sense of smell should be confined to them. Even in
ourselves it is not always easy to distinguish the sense of taste
from that of smell.

Graber deprived a beetle (Silpha thoracica) of its antenne, and
then tested it with oil of rosemary and asafcetida. A beetle
of the same species, but with antennz, showed its perception by
movement in half a second to one second in the case of the oil of
rosemary, and rather longer, 1 second to 2 seconds, in the case of
the asafcetida. The St/pha without antennz showed its percep-
tion of the oil of rosemary in 3 seconds on an average of eleven
times, while in no case did it show any indication of perceiving
the asafoetida even in 60 seconds.

Prof. Graber infers, “dass der eine Geruchsstoff (Assa-feet),
der nichts weniger als ein sehr feiner ist, nur durch das Medium
der Fihler perzipiert Bewegungen auslost, wihrend der andere
(Rosmarin6l) ahnliches auch ohne Vermittlung dieser angeblich
spezifischen Geruchsorgane bewirkt.”

Graber questions some of the experiments which seemed to meto
demonstrate the existence of a sense of smellin ants. He says:—

‘Da Lubbock noch hinzufigt, dass keiner, der das Benehmen
der Ameisen unter diesen Umstindén beobachten wirde, den ge-
ringsten Zweifel an ihrem Geruchsvermogen haben konnte,
wihlte ich auch diese Methode, um zu erforschen, wie sich etwa
der Fiuhler beraubte Ameisen verhalten, wiirden. Ich war
nicht wenig tberrascht zu finden, dass auch diese (es handelt
sich um Formica rufa) vor dem Riechobjekt umkehrten. Um
ganz sicher zu gehen, versuchte ich’s aber noch mit dem gleichen
Arrangement aber mit Weglassung des Riech-stoftes, und siehe
da! sie kehrten auch jetztnoch um! Bei genauerer Beobachtung
der von einer Ameise vom Anfang an auf dem Papiersteg zu-
rickgelegten Strecke stellte sich auch bald heraus, dass es sich
bei dem gewissen Umkehren lediglich um ein versuchsweises
Abschreiten oder Ausprobiren des unbekannten Weges bandelt,
oder dass sich die Ameisen ahnlich benehmen wie wir selbst,
wenn wir etwa auf einem schwanken Brette eine tiefe Gebirgs-
kluft tberschreiten sollen.’’*

* V. Graber, ‘‘ Verg]. Grundversuche ber die Wirk. und d. Aufnahmestellen
chem. Reize bei den Thieren.” Biol, Centraiblatt. vol. xiii. p. 449 (1885-6),

SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS. 127

M. Graber’s observation is, I doubt not, quite correct, but his
inference is not well founded, nor was his experiment the same as
mine. Itis quite true that if an ant be started off along a narrow
paper bridge, she will after awhile turn round and come back
again. I do not, however, think that this is due, as he suggests,
to any sense of giddiness. Ants which habitually climb trees
are not likely to be affected by any such sensation. It is rather,
I believe, that they feel they are being sent on a fool’s errand.
Why should they start off and run straight forward into a strange
country ? They turn round in hopes of finding their way home,
whether the bridge is high or low, broad or narrow, or indeed
whether they are on any bridge at all. M. Graber has not
observed that I expressly stated that in each case they stopped
exactly when they came to the scented pencil.

SENSE OF DIRECTION.

Fabre has made a number of experiments from which he con-
cludes that bees have a certain sense of direction. My own
experiments led me to the opposite opinion. Ihave now repeated
some of them, and made others, which all led to the same con-
clusion. For instance I put down some honey on a piece of
glass, close to a nest of Lasius niger, and when the ants
were feeding I placed it quietly on the middle of a boaid 1 ft.
square and 18 inches from the nest. I did this with 13 ants and
marked the points at which they left the board. Five of them
did so on the half of the board nearest to the nest, and 8 on
that turned away from it; I then timed 3 of them. They all
found the nest eventually, but it took them 10, 12, and 20 minutes
respectively. Again, I took 40 ants which were feeding on some
honey, and put them down on a gravel-path about 50 yards from
the nest, and in the middle of a square 18 inches in diameter,
which I marked out on the path by straws. They wandered
about with every appearance of having lost themselves, aid crossed
the boundary in all directions. J marked down where they left
the square and then took them near the nest, which they joyfully
entered. Two of them, however, we watched for an hour.
They meandered about, and at the end of the time one was about
2 feet from where she started, but scarcely any nearer home; the
other about 6 feet away, and nearly as much further from home.

I prepared a corresponding square on paper, and having indi-
cated by the arrow the direction of the nest, | marked down the
spot where each ant passed the boundary. They crossed it in all

128 SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS.

directions; and if the square were divided into two halves, one
towards the nest and one away from it, the number in each
was almost exactly the same.

RECOGNITION OF FRIENDS.

In the interesting memoir already cited Forel says:—“ Lubbock
(1. c.) a cru démontrer que les fourmis enlevées de leur nid a
létat de nymphe et écloses hors de chez elles étaient néanmoins
reconnues par leurs compagnes lorsqu’on les leur rendait. Dans
mes Fourmis de la Suisse, javais cru démontrer le contraire.
Voici une expérience que j’ai faite ces jours-ci: le 7 aout, je
donne des nymphes de Formica pratensis prés déclore a quel-
ques Formica sanguinea dans une boite. Le 9 aotit quelquesunes
éclosent. Le 11 aoit, au matin, je prends l’une des jeunes
pratensis agée de deux ou trois jours seulement et je la porte
a la fourmiliere natale dont elle était sortie comme nymphe
seulement 4 jours auparavant. lle y est fort mal recue. Ses
nourrices d’il y a 4 jours l’empoignent qui par la téte, qui par le
thorax ; qui par les pattes en recourbant leur abdomen d’un air
menacant. Deux d’entre elles la tinrent longtemps en sens inverse
chacune par une patte en l’écartelant. Enfin cependant on finit
par la tolérer, comme on le fait aussi pour de si jeunes fourmis
(encore blanc jaunatre) provenant de fourmiliéres différeates.
J’attends encore deux jours pour laisser durcir un peu mes
nouvelles écloses. Puis j’en reporte deux sur leur nid. Elles
sont violemment attaquécs. L’une d’elles est inondée de venin,
tiraillée et tuée. L’autre est longtemps tiraillée et mordue, mais
finalement laissée tranquille (tolérée?). On m’objectera l’odeur
des sanguinea qui avait vécu 4 jours avec la premiere et 6 jours
avec les deux dernitres. A cela je répondrai simplement par
Vexpérience de la page 278 4 282 de mes Fourmis de la Suisse,
ou des &. pratensis adultes séparées depuis deux mois de leurs
compagnes par une alliance forcée avec des F. sanguinea, alliance
que j’avais provoquée, reconnurent immédiatement leurs anciennes
compagnes et s’alliérent presque sans dispute avec elles. Je
maintiens done mon opinion: les fourmisapprennent a se connaitre
petit a petit a partir de leur éclosion. Je crois du reste que
c’est au moyen de perceptions olfactives de contact.”*

I have, however, repeated my previous observations with the
same results.

* August Forel, “ Expérience et Remarques critiques sur les Sensations des
Insectes.”” Recueil Zool. Suisse, tome iv. (1887), pp. 179-180,

SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS. 129

i
i j

Part of one of my Ant-cases, showing circular nest of Tapinoma, with path-
way leading to the opening. The Ants enter the circular fortification by one
or two tunnels not visible in the figure.

SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS. 131

At the beginning of August I brought in a nest of Lasius
niger containing a large number of pupe. Some of these I placed
by themselves in charge of three ants belonging to the same
species, and taken from a nest which I have had under observa-
tion for rather more than 10 years. On the 28th August, I took
12 of the young ants, which in the meantime had emerged from
the separated pupz, selecting some which had all but acquired
their full colour. Four of them I replaced in their old nest, and
four in that from which their nurses were taken.

At 4.30. In their own nest none were attacked.
In their nurses’ nest one was attacked.

ye In their own nest none were attacked.
In their nurses’ nest all four were attacked.
8. In their own nest none were attacked.

In their nurses’ nest three were attacked.
The next day I took six more and marked them with a spot
of paint as usual, and at 7.30 replaced them in their own nest.
At 8 Ifound 5 quite at home. The others I could not see,
but none were attacked.

8.30 a s re a
9 ” 3 ” ” ”

10 ” 4 ” ee) ”

i ye ‘9 ,» y»

12 ae, is : -
1 99 3 ? 9 ”
4, ” 4 ” ” ”
(4 ” i ” ” ”
9 ” 2 ”

The next morning I could only see two, but none were being
attacked and there were no dead ones. It is probable that the
paint had been cleaned off the others, but it was not easy to find
them all among so many. At any rate none were being attacked
nor had any been killed.

These observations, therefore, quite confirm those previously
made, and seem to show that if pupx are taken from a nest, kept
till they become perfect insects, and then replaced in the nest,
they are recognized as friends.

When we consider the immense number of ants in a nests
amounting in some cases to over 500,000, it is a most remarkable
fact that they all know one another. If a stranger, even be-

132 SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS.

longing to the same species, be placed among them, she will be
at once attacked and driven out of the nest. Nay, more, I have
already shown that they remember their friends even after more
than a year’s separation, and that it is not by any sign or pass-
word, because even if rendered intoxicated, so as to be utterly in-
sensible, they are still recognized. As regards the mode of re-
cognition, Mr. McCook considers that it is by scent, and states
that if ants are more or less soaked in water, they are no longer
recognized by their friends, but are attacked. He mentions a case
in which an ant fell accidentally into some water :—

‘She remained in the liquid some moments and crept out of it.
Immediately she was seized in a hostile manner, first by one,
then by another, then by a third; the two antenne and one leg were
thus held. <A fourth one assaulted the middle thorax and petiole ;
the poor little bather was thus dragged helplessly to and fro fora
long time, and was evidently ordained to death. Presently I took
up the struggling heap. Two of the assailants kept their hold ; one
finally dropped, the other I could not tear loose, and so put the
pair back upon the tree, leaving the doomed immersionist to her
hard fate.”

His attention having been called to this, he noticed several
other cases, always with the same result. I have not myself
been able to repeat the observation with the same species, but
witb two at least of our native ants the results were exactly re-
versed. In one case five specimens of Lasius niger fell into water
and remained immersed for three hours. I then took them out
and put them into a bottle to-recover themselves. The following
morning I allowed them to return. They were received as
friends, and though we watched them from 7.30 till 1.80 every
hour, there was not the slightest sign of hostility. The nest was
moreover placed in a close box, so that if any ant were killed we
could inevitably find the body, and I can therefore positively state
that no ant died. In this case, therefore, it is clear that the
immersion did not prevent them from being recognized. Again,
three specimens of Formica fusca dropped into water. After
three hours I took them out, and after keeping them by themselves
for the night to recover, I put them back into the nest. They
were unquestionably received as friends, without the slightest
sign of hostility, or even of doubt. I do not, however, by any
means intend to express the opinion that smell is not the mode
by which recognition is effected.

°
——

SS EE —Ee—eEeEee

.
f
:

SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS. 133

LONGEVITY.

It may be remembered that my nests have enabled me to keep

ants under observation for long periods, and that I have identified
workers of Lasius niger and Formica fusca which were at least
7 years old, and two queens of Formica fusca which have lived with
me ever since December 1874. One of these queens, after ailing
for some days, died on the 30th July, 1887. She must then have
been more than 13 years old. I was at first afraid that the other
one might be affected by the death of her companion. She
lived, however, until the 8th August, 1888, when she must have
been nearly 15 years old, and is therefore by far the oldest
insect on record.
» Moreover, what is very extraordinary, she continued to lay
fertile eggs. This remarkable fact is most interesting from a
physiological point of view. Fertilization took place in 1874 at
the latest. There has been no male in the nest since then, and,
moreover, it is I believe well established that queen ants and
queen bees are fertilized once for all. Hence the spermatozoa
of 1874 must have retained their life and energy for 138 years, a
fact, I believe, unparalleled in physiology.

In some plants (Rues) the pollen-tube takes as long as two
years to reach the ovule. Indeed the pollen has some claims to
be regarded asa separate organism, for it certainly possesses the
power of growth and of assimilating nourishment. There is not,
however, so far as I am aware, any other case which can compare
with that of my queen ant as regards the longevity of the male
element. One is even tempted to wonder, under such cireum-
stances, whether there is any multiplication of the spermatogenic
cells.

Moreover the case is not altogether isolated. I had another
queen of Formica fusca which lived to be 13 years old, and I have
now a queen of Lasiws niger which is more than 9 years old, and
still lays fertile eggs which produce female ants.

Ants AND SuEps oF Melampyrum pratense.

M. Lundstrom has recently called attention to the interesting
fact that the seeds of this plant closely resemble pupe of ants
in size, shape, and colour, even to the black mark at one end.
He has suggested very ingeniously that this may be an advantage
to the plant by deceiving the ants, and thus inducing them to
carry off and so disseminate the seeds. There seemed, however,

LINN. JOURN.—ZOOLOGY, VOL. XX. 12

134 SIR JOHN LUBROCK ON ANTS, BEES, AND WASPS.

some improbability in the idea that ants should be deceived |

as to their own sisters. M. Liindstrom has found seeds of this
species in ants’ nests, but has not actually seen ants carrying
them off, and I thought it would be worth while to determine
this.

Accordingly I took 10 seeds and placed them just outside one
of my domesticated nests of Lasius niger. A certain number of
ants were outside, and I saw several come up to the seeds, but
they took no notice ofthem. I left them lying there for two days.
I then tried them with another nest, the roof of which consisted
of two plates of glass, side by side, but with an interval between
them. I placed the seeds in this interval, and uncovered one of
the sides. The ants immediately began carrying the pupe which
were thus exposed to the light to the other, covered part of
the nest, in doing which they necessarily passed close to the
seeds, but they did not take the slightest notice of them. This
operation was finished by 11 a.m., and I left them undisturbed
till 12, the seeds remaining unnoticed and untouched. I then
moved the cover from one half of the nest to the other, and the
ants immediately began transporting the pupe to the shaded
half. One or two of them examined the seeds, not one of which,
however, was moved. This took about an hour. At 4 PoM.,
however, three of the seeds had been carried in, and the next
day, at 7 a.M., two more seeds had been carried in. I then
removed them, and put them just outside one of my nests of
F. fusca.

Aug. 31. 7 a.m. None have been touched. I now put the
covering clese to, but not over them. The ants took no notice
of them.

Sept. 2. I now placed them just in the entrance of the nest
and covered over a part just outside. The ants collected as
usual under the cover. I then removed the cover just inside the
nest, so that the ants to reach it had to pass among the seeds.
They, however, came in, but did not move a single seed. I once
again moved the cover outside, and they followed it as before,
but without moving the seeds.

So far as these observations go it would seem that Ff. fusca
takes no notice of these seeds, but that they really are under
certain circumstances carried off by Lasius niger.

Wasps.

Mr. and Mrs. Peckham have published in the ‘ Proceedings of

SIR JOHN LUBBOCK ON ANTS, BEES, AND WASPS. 185

the Natural History Society of Wisconsin’* a very interesting
paper on the special senses of wasps, and their conclusions concur
closely with mine.

It appears from their observations that some wasps stay out
all night and return early in the morning before the others begin
coming out. For instance, on the 18th Aug. the first wasps left
the nest at 7.25 ; 10, however, had already returned, 3 of them
before 5 a.m. It appears from their observations that the
average time a wasp is absent from the nest, that is the average
length of each excursion, is 43 minutes. They observe that this
may appear inconsistent with my observations, when the trips
were shorter and more numerous, one of my wasps having paid
me 116 visits in 15 hours and a half. But, as they justly observe,
the cases are not comparable. My wasps and theirs were like
Jacob and Ishmael—mine haa everything ready prepared for
them, theirs had to hunt for themselves.

As regards the sense of hearing, they repeated some of my
experiments with the same results. They seem to consider that
as regards the sense of colour their conclusions are somewhat
at variance with mine.

As regards the supposed sense of direction they say +:—‘“‘ Sir
John Lubbock, in dealing with the sense of direction in ants, con-
cluded, after a number of observations, that they were endowed
with this sense in a high degree. Subsequently he discovered,
quite accidentally, that the ants found their way by observing
the direction in which the light was falling.” My conclusion
was, however, the result of many observations carried on under
varied conditions, and I should hardly call it an accident.

They came to the conclusion, as I had done, that wasps have
no sense of direction, that is to say in the form of a mysterious
additional sense, but that, if they do not know where they are, they
rise higher and higher into the air, circling as they do so, until they
discover some high treetop or other object that had before served
them as a landmark, and that in this way they are able to make
their way home. This entirely tallies with my own conclusion. It
is interesting as showing that the vision of wasps must be good”
for somewhat distant objects.

They also found, as I had done, that their memory varied
greatly in different individuals.

* April 1887.
+ Proc. Nat. Hist. Soc. Wisconsin, April 1887, p. 118,
12"

186 DR. J. RAE ON THE BIRDS AND MAMMALS

Lastly, 1 give two illustrations which will convey an idea of »
some of my ant-nests.

That on p. 129 represents about a quarter of one of my frames.
The shaded part represents the earth, which will be seen to have
been arranged by the ants into a sort of circular fortification, or
zereba, access to which is obtained by one or two tunnels, not
visible in the illustration, and to which a pathway leads from the
entrance.

The second (facing this page) represents a nest of Lasius niger.
It shows the entrance, a vestibule, and two chambers, in the outer
and larger one of which the ants have left some pillars, almost
as if to support the roof. The queen is surrounded by workers,
those in her immediate neighbourhood all having their faces
turned towards her. There is a group of pups, and several of
larvee, sorted as usual according toages. There are alsoa number
of the blind woodlice (Platyarthrus Hoffmanseggi).

Notes on some of the Birds and Mammals of the Hudson’s Bay
Company’s Territory, and of the Arctic Coast of America.
By Joun Raz, M.D., LL.D., F.R.S., &. (Communicated by
G. J. Romanss, F.L.S.)

[Read 16th February, 1888. ]

Durine twenty years’ residence in various parts of the Hudson’s
Bay Company’s Territory, embracing the extreme south of the
shores of James’s and Hudson’s Bays, and north to the Arctic
Sea, I have had, as a sportsman, many opportunities of devoting
considerable attention to the habits and peculiarities of animals,
especially birds, over a very extensive field of observation, the
result of which I shall attempt to give in the following remarks,
some of which may possibly be new, other points disputed or
perhaps already well known.

My first ten years were spent at Moose Factory, the principal
depot of the Hudson’s Bay Company in the Southern Depart-
ment, lat. 51° N., long. 81° W., where the marshes along the
coast form the favourite feeding-grounds of a variety of geese,
ducks, &c. on their migrations to and from their breeding-places
in the north. A great part of my spare time at these seasons,
spring and autumn, was spent in shooting these birds, and at
the same time acquiring some knowledge of their peculiarities.

[Zo face p, 136.

illars, and inner room; the queen surrounded by workers ;
nd the blind woodlice (Platyarthrus Hoffmanseggit).

Linn. Journ., Zool. SV oliecse [To face p. 186

vestibule, main chamber with pillars, and inner room; the queen surrounded by workers ;

Nest of Lasius niger, showing the entrance,
g to ages; and the blind woodlice (Platyarthrus Hoffmanseggii).

a group of pups, and several of larvx, sorted accordin

Sag 7 i, ieee ae Se al a
Po <a eS ae — Ts 1. —_ = 7
ms a ' <i e o a ~~ > = “ _; — ws

OF THE HUDSON’S BAY TERRITORY. 137

First let me notice that magnificent bird the Canada goose (Anser
canadensis), probably one of the finest of its kind in the world.
This is the earliest arrival of the waterfowl migrants in spring, and
makes its appearance at Moose with extreme regularity on the
23rd of April, St. George’s day. So much is this the case that,
during the ten years of my residence there, we had on every St.
George’s day a goose for our mess dinner, first seen and shot
on that day, and this I learnt from older inhabitants had been
the case for many previous years. I may add that this bird
arrives with equal punctuality at York Factory in lat. 57° N.,
450 miles further north, but a week later.

The Cree Indians, both at Moose and York Factory, assert
positively that a small brown bird uses this goose as a convenient
means of transport to the north, and that they have been often
seen flying off when their aerial conveyance was either shot or
shot at. - The little passenger has been pointed out to me, but I
have forgotton its name, and it certainly makes its appearance on
the shores of Hudson’s Bay at the same date as this goose, which,
by the way, is the only kind that is said to carry passengers *.
The natives of the McKenzie River, more than 1000 miles to
the north-west, tell the same story. From my observation I am
led to believe that there is another species of the Canada goose,
much larger, but less numerous. The male of this larger bird
is distinguished by a ruddy-brown colour of plumage on the
breast, by the greater loudness and sonorousness of its call,
and by its much greater size, so that a difference is made in
the quantity served out as rations to the men. The line of
flight is also different, as they generally pass by Rupert’s River
about 100 miles east of Moose, but a few are sometimes to be

* Since this paper was read, an article by J. HE. Harting, on “ Small birds
assisted on their migrations by larger ones,” has appeared in the Natural
History columns of ‘The Field’ of March 31st, 1888, in which will be found
much additional information on the subject.—J. R.

t Baird, Brewer, and Ridgway in their ‘Water Birds of North America’
recognize two species of Canada Goose—a large species with 18 to 20 tail-feathers,
and a smaller one with 13 to 16 tail-feathers. Each of these supposed species
they subdivide into two races, a grey and a brown one. Dr. Elliott Coues,
in his ‘ Key to North-American Birds’ (2nd ed. 1884, p. 689), remarks ‘“‘ there
seems little probability of establishing good characters for more than one
species of the canadensis group, with probably four varieties :—(1) large, no
collar (702, canadensis) ; (2) small, no collar (704, hutchinsi); (3) large,
collared (702 a, occipitalis) ; (4) small, collared (703, lewcopareia).” ‘The two
larger ones both have 18 tail-feathers ; the two smaller ones 16 only.—Eb.

138 DR. J. RAE ON THE BIRDS AND MAMMALS

obtained at the latter place, which afforded mean opportunity of
comparing them with the more common or smaller kind. This
Anser canadensis (major ?), instead of being seen feeding in the
marshes as the others do in autumn, chiefly frequents the higher
and more rocky grounds on the eastern shores of James’s Bay,
where its principal food consists of berries of various kinds.

By far the most numerous of the goose tribe that visit the
Moose marshes in the autumn are the snow goose, or white-
wavy (Anser hyperboreus), and the blue-winged goose of
Edwards (Anser cerulescens). These birds resemble each other
very much in size, call, and form, but not in colour; and as
they often feed in proximity, the blue goose was for a long
time supposed to be merely the. young of the snow goose; an
erroneous opinion, which I endeavoured to correct in a little
book published in 1850 by Boone, entitled ‘ Expedition to the
Shores of the Polar Sea 1846-47.’ *

These snow and blue-winged geese have a peculiarity I have
never noticed in any other species. Previous to taking their
southern flight from Hudson’s Bay some time in October,
they remain for several days almost constantly on the open sea,
washing themselves, taking sudden and rapid flights, apparently
having a “‘ happy time,” but they are never seen feeding. They
are at this time very fat, and when shot, their stomachs and
intestines are found to be entirely empty, resembling in this
respect salmon, I am told, prior to, and in preparation for, their
hard work in ascending rivers to their spawning-beds. After
this period of fasting, ablution, and exercise has been gone
through, the birds are evidently ready to start on their flight
of some hundreds of miles. On the first favourable opportunity,
which means a northerly wind, they take wing in batches of
fifty or more, circling round until they attain a safe altitude,
and then bear away on a true southerly course, never resting
until they reach winter-quarters, on the shores or swamps of the
Southern States fT. |

* The specific distinctness of these two geese is generally admitted by modern
ornithologists.—Ep.

t I may mention that 45 years ago the blue-winged and white-wavy geese
visited Moose in about equal numbers, as they still do ; whereas at Albany, 100
miles to the north, there were great numbers of the white bird and scarcely a
blue-wing to be seen. Now the two kinds are about equally abundant there,

whilst at Rupert’s River, 100 miles east of Moose, now, as formerly, the blue-
winged birds are alone met with.—J. R.

OF THE HUDSON’S BAY TERRITORY. 139

The Canada goose, on the contrary, stops by the way to feed,
especially on the lakes and swamps where there is wild rice,
which makes both geese and ducks much finer eating than any
other kind of food I know. Both the white and blue wavy
are excellent eating, and one of them with a pound of flour
or bread, is given as a day’s rations, and is much liked by the
men, especially when fresh. Many thousands are annually cured
with salt, and packed in barrels for use at the Hudson’s Bay
Company’s stations on the coast; and the Indians bone and
dry a great number for winter food.

All species of grouse in British North America have a well
known habit of passing the night under the snow, during the
winter, to protect themselves from the cold; but possibly a practice
which most of them follow more or less when the snow is not too
hard packed may not have been generally observed. The bird
is not content to make its resting place close to the door by
which it has entered the snow, but usually bores a tunnel a few
inches under the surface, three or more feet in length, before
settling down for the night. The cause for going through so
much, apparently, useless labour was at first difficult to under-
stand, for its bed would have been equally warm had the bird
remained within a foot of where it had entered the snow, but
a little more experience taught me to admire the acuteness and
intelligence of the proceeding, for during my walks in the woods
I frequently came to places where a fox, lynx, or other carnivore
had in the night approached cautiously (judging by the short
steps) and made a long spring on the entrance hole ; the occupant
was not there, however, but had flown up 3 or 4 feet off, as seen
by its exit in the snow, and was thus saved from almost certain
death.

The prairie-hen, which is fairly numerous near Moose, shows
great carefulness in this respect, and in very cold weather takes
its “siesta’’ between breakfast and supper under the snow,
out of which I have often seen them pop their heads, without
taking wing, before I had got within shot, no doubt to observe
if an enemy were approaching.

Without counting the small white grouse peculiar to the
Rocky Mountains, I believe there are three other species to be
found at or near the Arctic coast.

First and chief among these is the Willow-grouse (7. salicetz),
by far the most numerous, and forming an important article of

140 DR. J. RAE ON THE BIRDS AND MAMMALS

food for the Indians living near the coast of Hudson’s Bay. These
birds extend their breeding-grounds up to the Arctic shores of
America, but as a rule do not extend their migrations to the
large islands further north.

Dr. Bell says :—‘‘ The summer plumage of Tetrao saliceti, the
cock bird, is exactly the colour of the English cock pheasant
with the exception of the wings, which have a good deal of
white, and in winter the white of the living bird has a beautiful
delicate rosy tint, which forms a considerable contrast with the
surrounding snow.” * ‘This description is somewhat misleading.
The plumage of the cock willow-grouse in summer resembles as
nearly as possible that of the Scottish grouse, with the exception
that the primary feathers of the wings of the former are always
white. The “delicate rosy tint of the white plumage” is rarely
seen, and only in beautiful warm sunny winter or spring days,
mever on a cold winter day. In the spring, or pairing season,
the call and peculiar habits of the Willow and Scottish cock
Grouse exactly resemble each other.

The Rock Grouse (Tetrao rupestris) is so well marked by
its smaller size, its more slender beak, and the black patch
extending from the angle of its mouth to the eye in the male,
that it cannot be mistaken for any other.

A third species differs considerably from Tetrao saliceti and
rupestris, being fully as large as the former, but the bill seems
shorter, its feet smaller, and its call perfectly different from either
of the others; it is also found further to the north. I saw a
good many males (the hens were nesting) on Wollaston Land,
lat. 69° N., in May and early June, and managed to shoot a few,
although they were very wild, possibly with the intention of
leading me away from the nest +. |

At Toronto, Lake Ontario, Canada, an island forms an excellent
harbour. Along the outer side of this island an immense number
of a small sandpiper, called “black heart” (the Dunlin, Zringa
alpina pacifica), pass northward every season on the 23rd April
(St. George’s day) and are not seen on any other day, except,
perhaps, some wounded ones on the 24th that cannot continue

* See “‘ Notes on Birds of Hudson’s Bay” by Robert Bell, M.D. Proceedings
of the Royal Society of Canada for 1882, vol. i. p. 49.—J. R.

t There is, I think, a specimen of this bird in the Natural History Museum
at South Kensington called Tetrao mutus, but certain distinguished naturalists

do not believe in it. I brought one or two specimens from the Arctie regions
in 1847, which were presented to the British Museum.—J. R.

OF THE HUDSON’S BAY TERRITORY. 141

their flight. This flight is so well known that many sportsmen
line the shores of the island on the day named, and hundreds of
these pretty little birds, which are good eating, are shot.
Persons who have resided many years in Toronto have told me
that they did not remember an instance of variation in the date of
the arrival of these birds.

Over almost every part of the wooded country of British
North America, east of the Rocky Mountains, the American hare
(Lepus americanus), usually called the “ rabbit’? by the Hudson’s
Bay Company’s people, is to be found in more or less abundance,
and it may not be generally known that every ten years these
animals are attacked by an epidemic so fatal, that from being
very numerous they gradually die off until scarcely one is to be
seen. The survival of the fittest then begins to increase, and at the
end of ten years they are again at their maximum. Ihave myself
seen two of those cycles, and know men in the Hudson’s Bay
Co’s. service, who have witnessed four or five of such events.

The latest years of abundance were 1885 and 1886, the hares
having increased gradually from 1880-81, which were years of
scarcity. The curious thing is that this takes place in the same
years over an extent of country about as large as one fourth of
Europe. It has been asserted by distinguished naturalists, among
them by my friend Sir John Richardson, that the hares migrate ;
but this cannot be the case, for it is not known where they go to,
besides they are found sitting in their “forms” dead, usually under
small pine or spruce trees, the branches of which grow close to the
ground. Iaccount for the disease in this way. The hares do not
spread themselves broadcast all over the country, but live in
colonies extending over a square mile or more, where the trees and
plants on which they feed are abundant, and here they become so
crowded together that the ground gets poisoned by their excreta,
as is the case with domestic poultry when kept too long on the
same land without being cleaned or shifted, and hence disease.
The grouse disease in Scotland I attribute to the same cause,
when too large a stock has been left on the moors. When the
grouse “pack,” they have in winter some favourite resort to
which they fly during storms (chiefly from the west) for shelter,
and I have seen such places perfectly covered with droppings,
even in Orkney, where grouse are never very numerous.

The effect of these epidemics among the hares is peculiar, and
affects both the Indians and some of the fur-bearing animals,
as 1 shall endeavour to explain.

142 DR. J. RAE ON THE BIRDS AND MAMMALS

When the hares are abundant, an Indian and his family pitch
their tent among them in winter, and cut down a number of the
trees, part of which forms the hares’ favourite food, then make
barriers of small pine trees and brush, through which gaps are cut
to allow the hares torun through. Allowing thema short time to
fatten up on the abundant food provided for them, a hundred
snares, or more, are set in the openings of the barriers, and these
Snares are attended to by the wife and children of the hunter,
whilst he sets up a number of traps in two or three directions to
the distance of perhaps eight or ten miles from his tent, each of
which he visits two or three times a week to bring home the fur-
bearing animals caught, chiefly fox, lynx, fisher, and marten,
taking with him on each visit a supply of fresh baits. The
Indian is thus carrying on his winter hunt in the most advan-
tageous manner, the hares attracting the carnivora above named
to his traps, whilst at the same time they supply, without any
difficulty, an abundance of food and the most comfortable winter
blankets known. The making of these blankets is peculiar ; the
hare skins, after being cut into strips, are stitched end to end,
and plaited so loosely that the finger can be poked through
them in any direction, yet a person can sleep comfortably
wrapped up in one of these on the coldest night, with the tem-
perature say 40° below zero, without any fire.

When the hares become scarce, not only has the Indian to
travel about;in search of large game, or go fishing to obtain
food for himself and family, but the fur-bearing animals have
also to wander abroad; consequently the Indian cannot catch
so many hares, and they have time to increase and multiply until
a season of abundance again comes round.

The house-building habits of the muskrat in nearly every part —
of British North America are well known, but there is one plan
to which it sometimes resorts under certain circumstances which
appears to show great intelligence in enabling it to get its food
more readily. The muskrat, when about to build its house,
selects a pond or swamp of good pure water, on the bottom of
which grow the plants which constitute its winter supply of food.
If the pond or swamp is of considerable extent, and the house a
large one containing many rats, they, when the water begins to
freeze in early winter, keep several holes open in the ice in
different directions, and at a distance from the house, and build
a little hut of mud and weeds (just large enough to hold one rat
comfortably) over each hole which-—especially when covered with

OF THE HUDSON’S BAY TERRITORY. 143

-snow—prevents it freezing up. These huts euable the rats to
extend their feeding ground to all parts of the pond, which could
not be reached at all, or with difficulty, from the house if they
had to swim home every time with a mouthful of food, to eat it.
With these little shelters they are saved a great amount of labour
and are enabled to reach all the food in the pond.* I remember;
when on a snow-shoe journey, one of my men went very quietly
up to one of these miniature mud huts, and knocked it over with
his axe, disclosing a live rat with some of the food it had been
eating. The practice of building these little eating huts is by
no means common, and does not seem to be resorted to when
the pond is of moderate dimensions, and all parts of it can be
reached from the house without difficulty.

I am not aware if it is generally known that the lemmings
(Myodes hudsonicus, &c.) of North America migrate much in
the same manner as do those of Norway and Sweden. When
travelling in June 1851 southward from the Arctic coast along
the west bank of the Coppermine River, and north of the Arctic
Circle, we met with thousands of these lemmings speeding north-
ward, and as the ice on some of the smaller streams had broken
up, it was amusing to see these little creatures running back-
wards and forwards along the banks looking for a smooth place
with slow current at which to swim across. Having found this,
they at once jumped in, swam very fast, and on reaching the
opposite side gave themselves a good shake as a dog would, and
continued their journey as if nothing had happened. At that
date the sun was above the horizon all hours of the 24, and we
were travelling by night to avoid the snow-glare in our eyes, the
sun being then in our rear. As the lemmings appeared to travel
only by night, we should not have seen them had we been
travelling in the daytime, for they then hide themselves under the
snow, orstones. The man who was carrying our cooking utensils
and small supply of provisions, having, when fording a stream,
been swept into a deep hole by the current, whereby his whole
load was lost, we had, for a day or two, to live chiefly on
lemmings roasted between thin plates of limestone, and found
them very fat and good. Our dogs easily killed as many as they
required. Prior to this, whilst on the coast, crossing the ice to
islands some miles distant, a lemming was noticed defending

* The beaver, especially when its dam is large, scrapes holes in the banks

from under water upwards unti! above the water-level, to which it retires to feed
instead of going back to its house.—J. R.

144 ON THE MAMMALS OF THE HUDSON’S BAY TERRITORY.

itself most gallantly against the attack of two large gulls, which
continued swooping down at it, but were kept at bay by the
brave little animal turning on its back and squealing loudly. I
ran up and was in time to drive away its dangerous opponents,
and place it in safe shelter under a piece of ice.

Occasionally large numbers of lemmings are found drowned
along the shores of James’s Bay, but as they are generally seen
after a very high tide, it is uncertain whether they are then
migrating, or merely caught by the high tide on their native
grounds. As soon as snow falls to any considerable depth, the
lemming leaves its summer quarters in the ground, and builds
a bed of grass and moss as a winter shelter, from which it
bores under the snow in one or more directions to obtain food.
They do not seem to hibernate during the winter, for when
wintering in a snow-hut at Repulse Bay in 1853-4 I used to
hear them scratching tracks through the snow throughout the
cold season*.

The Arctic hare (Lepus glacialis), of which I have seen and
shot a good many, shows a considerable amount of intelligence
in its efforts to throw foxes, wolves, and other enemies off the
scent. It seems to have been eminently successful in this re-
spect with the crew of McClure’s ship when wintering in Prince
of Wales’s Strait, if it were not from bad shooting on the
part of the men, for we are told that although hundreds of
hares were seen, the total bag in a month was only seven
hares !

The Arctic hare, after its night’s feed, usually goes some dis-
tance from its feeding-ground before settling down in its form

for the day. When following up the track, I was at first ex- |

tremely puzzled to find all at once a complication of tracks, and
on going a little further found no track at all. On retracing my
steps and carefully examining the snow, I observed two very small
marks, at least 20 feet distant, from the main track. A succes-
sion of two or three similar long jumps followed, and a very little
experience taught me that the hare was always crouching near ;
usually close to some large stones or rock uncovered by snow.
These long jumps were invariably made to leeward for obvious
reasons, because if to windward a fox would have scented his prey.

* Many lemmings were seen at the winter quarters of the Nares Arctic
Expedition, in lat. 82°, and a large “ cache” of dead ones was found, made by
a white fox.—J. R.

— —

COPEPODA OF MADEIRA AND THE CANARY ISLANDS. 145

Even after becoming acquainted with this artful dodge, the hare
must be stalked with caution. Being all white except the tips
of its ears, it is not easily seen when nearly buried in the snow,
and when discovered the sportsman must dissemble, and pretend
that he has not seen it, walking in an oblique direction, ap-
proaching, but at the same time as if passing by, never looking
directly at the game. When near enough he should wheel
suddenly round and fire. If this plan be not adopted, the hare
will very generally bolt round the rock, and escape under its
shelter. This may be called pot shooting, but the best of sports-
men have often to do the same thing with ptarmigan in the
Scottish mountains, when these birds fly round a rock and are
out of sight in an instant, if not shot before taking wing.

Copepoda of Madeira and the Canary Islands, with Descriptions
of New Genera and Species. By Isaac C. THompson,
F.R.M.S. (Communicated by Prof. Hzerpmay, F.L.S.)

[Read 17th November, 1887. ]
(Puates X.—XIITI.)

Dvrine the spring of 1887, in company with Mr. W. 8. M‘Millan,
of Liverpool, I visited Madeira and the Canary Islands with the
object of collecting and examining the pelagic fauna at the
various convenient stopping-places.

We took dredge and tow-net, and all the necessary appliances
for the examination and preservation of specimens, and were for-
tunate in securing a large mass of material.

It is intended in this paper to treat of the Copepoda only ; and
as the dredging operations were neither so practicable, nor fruitful
in results as the tow-netting, it was to the latter that we de-
voted chief attention, all the Copepoda collected being free-
swimming species.

We used one of the very fine-meshed tow-nets of the ‘ Chal-
lenger’ pattern, and immediately preserved the captures in a
medium which I have always found useful for small Crustacea,
composed of glycerine, alcohol, and water in the following pro-
portions :—

Glycerine...... 1 part

Proof-spirit ... 2 pr | adding 1 per cent. of carbolic acid.
Water ......... 1 part

146 MR. I. C. THOMPSON ON THE COPEPODA OF

In this solution the colours (and they are very varied in the
Copepoda) are well preserved, and the tissues are rendered
sufficiently transparent for observation under the microscope
without further treatment.

We longed to cast a tow-net while traversing the fifteen hundred
miles of Atlantic waters that intervened before reaching our first
destination, Madeira, and improvised a long tough canvas bag,
weighted, with a wide-necked bottle at the bottom end, for the
purpose ; but the speed of the vessel was too great to allow of
any captures. The mails could not give way to the study of
biology, so our genial captain would not allow any temporary
stoppage; but the death of a poor fireman on the fourth day
out necessitating a funeral pause of a few minutes, we took
advantage of the opportunity by getting a haul; and the tow-net
brought up sufficient to occupy us and relieve the monotony of
the rest of the voyage. The haul was almost entirely composed
of Copepoda: Calanus jfinmarchicus, Centropages typicus and
C. brachiatus, Dias longiremis, and a few other species, nearly all
found on our British coasts.

The deep-blue transparent waters of Madeira proved very
fruitful both in number and variety of Copepoda, Funchal Bay
being our hunting-ground during two visits. Thence to Teneriffe,
where we anchored for a few hours in the Santa Cruz harbour,
and, as at Madeira, collected enough Copepoda to occupy us
many months in examining and working out.

Grand Canary was our next destination, and there we spent
many days dredging, collecting, and tow-netting at various parts.
of the island. The absence of any shelter or harbour at Las
Palmas rendered dredging difficult, for we had to take eight
men to manage the boat, the sea being seldom at all smooth.
And, from some cause or other, the tow-netting results obtained
here were certainly much poorer than those of the more shel-
tered bays of Madeira and Teneriffe.

Returning to Teneriffe, we crossed the island from Santa Cruz
to Orotava, grandly situated on the north side directly under
the Peak, and famous for its splendid climate and scenery. We
found the waters of the ocean here very plentiful in Copepoda,
and on various occasions collected a large amount of them. One
of these occasions was long after sunset, and it is worthy of note
that the Copepoda then taken were little different in point of
number or species from those taken in daylight, although with

MADETRA AND THE CANARY ISLANDS. 147

the night-haul were a largely increased number of Schizopoda,
some of them highly phosphorescent.

From a pretty thorough examination of the material obtained
at the various islands of the Canaries and Madeira, it seems
evident that their Copepodan fauna varies in quantity rather than
specifically. For, excepting several cases where one or two
specimens only of a species were found, which probably indi-
cated the rarity of that species, their geographical distribution
appeared to be general amongst the islands.

Sixty-four species in all were obtained, as enumerated below.
Of these six are new to science, and three of them required new
genera. (See Plates X. to XIII.)

Of the sixty-four species, twenty-two are known in British
waters; and of these, thirteen belong to the family Har-
pacticide.

Following the classification given by Brady*, the sixty-four
species are distributed into families as follows :—

alenidm ........... 30 species.
Ovclopid@::.)......... Batis
Harpacticide ...... re
Corycxide .......... 1 as
Artotrogide ......... ane

Description of the Species collected.

Family CALANIDA.
CALANUS FINMARCHICUS, Gunner.
C. vateus, Brady.

C. proprnquus, Brady.

This species, like the two preceding, is widely distributed. A
high power of the microscope ( x 400) shows very fine hairs on
the inner margin of the terminal spines of the swimming-feet, not
mentioned by Brady in his description of the species.

C. ronsus, Brady.

C. aractntis, Dana.
Both found very sparingly.

* ‘Monograph of British Copepoda, Ray Society, 1876-78 & 1879.

148 MR. I. C. THOMPSON ON THE COPEPODA OF

CaLANvS Pavo, Dana.

This species occurs in considerable numbers in the gatherings
from Madeira and Teneriffe ; but the elegant peacock-tail plumes
figured by Dana (Crustacea of U.S. Exploring Expedition) have
become detached in all the specimens.

PARACALANUS PARVUS, Claus.
One or two specimens only of this rare form were taken at
Orotava, Teneriffe.

EUCALANUS ATTENUATUS, Dana.

EK. setiGer, Brady.

Brady remarks upon the “ mimetic resemblance”’ of this spe-
cies to Calanus finmarchicus. Our specimens do not bear out
this remark ; and the absence of terminal spines to the swimming-
feet of this species is a distinguishing feature.

RHINCALANUS CORNUTUS, Dana.

R. aieas, Brady.

PsEUDOCALANUS ELONGATUS, Boeck.
PLEUROMMA ABDOMINALE, Claus.
LEUCKARTIA FLAVICORNIS, Claus.
ScoLEectrHRix Dana, Lubbock.

S. minor, Brady.

EUCHETA PRESTANDRED, Philippi.

CANDACE TRUNCATA, Dana.

C. NigROCINCTA, n. sp. (PI. X. figs. 1-6.)

Length 54, inch. Head-somite distinct from thorax. Body oval,
rounded in front; posterior segment terminated by two lateral
spines. Anterior antenne 23-jointed, bearing several spinous
processes on inner margin, and short sete at the termination of
each joint. The 8th, 9th, 10th, and 11th joints (fig. 1) are deeply
pigmented witha blackish-brown colour, the same pigment tinging
the setze of the posterior antenne and the terminal spines of the
swimming-feet. Posterior antenne (fig. 2) similar to that of
C. truncata, but with fewer terminal sete. Anterior foot-jaws
(fig. 3) 2-jointed, large and powerful, with two small claw-like
spines on basal joint, and five large ones on second joint. Pos-
terior foot-jaws small, 7-jointed. First four pairs of swimming-

MADEIRA AND THE CANARY ISLANDS. 149

feet (fig. 4) alike; inner branch has one joint only; outer edge
of main branch is finely serrated. Pigment does not extend
above the terminal spine, which is somewhat bent and finely ser-
rated and clothed with dark hairs on dorsal side. Fifth feet of
male (fig. 5) each 3-jointed, the right foot having extension on
inner side of middle joint, both terminated by two small claws.
Abdomen of male (fig. 6) 3-jointed, the third being nearly equal
in size to the other two; caudal segments about twice as long as
broad, terminated by short strong sete.

Several specimens of this strongly marked form were taken
at Orotava, Teneriffe, all of which appear to be males.

Jt .has strong points of resemblance to both Candace pachy-
dactyla and C. truncata, Dana, but is evidently distinct from
both. The dark rings on the anterior antenne distinguish it at a
glance.

CANDACE BREVICORNIS, 0. sp. (Pl. X. figs. 7, 8.)

Several specimens of a Candace were taken at Orotava and at
Santa Cruz, Teneriffe, which, although corresponding in most
respects with C. truncata, Dana, differ in some particulars which
are of sufficient importance to render it specifically distinct.

In the first place the anterior antenne are composed of eighteen
joints only, while C. truncata has twenty-four. Then the first
four pairs of swimming-feet (fig. 8) are the same as in C. truncata,
except that in our specimens, to which I propose to give the name
C. brevicornis, the sete are all elegantly plumed, much resembling
minute black feathers. No fifth feet can be made out in any of the
specimens obtained. Abdomen only 2-jointed, the caudal seg-
ments being long and straight, while in C. truncata they are
somewhat stumpy and divergent. The basal joint of the abdo-
men is less triangular than in C. truncata. Our specimens are
probably all females.

Dias tonarremis, Lilljeborg.
ACARTIA LAXA, Dana.

A.. DENTICORNIS, Brady.

In all our specimens of both species of Acartia there is a long
recurved spine at distal end of first joint of each anterior antenna,
which does not appear to have been previously described.

Drepanorpus FurcAtUs, Brady.

Several females of this species were found, but no males; a
LINN. JOURN.—ZOOLOGY, VOL. Xx. 13

150 MR. I. C. THOMPSON ON THE COPEPODA OF

similar experience to Brady’s, His specimens of this species
(‘ Challenger’ Report on the Copepoda, p. 77) were sparingly
found in three remarkably distinct areas.

Tremora DUBIA, Lubbock.

Very abundant in all the gatherings.

Istas CLAVIPES, Boeck.

First abdominal segment of female has two remarkable trifid
spines on ventral side, not described by Boeck or Brady. I first
noticed this point in specimens taken in Liverpool Bay. The
fifth feet of male also differ considerably from Brady’s drawing.

CENTROPAGES BRACHIATUS.
C. TYPICUS.
C. VIOLACEUS.

Our specimens of the latter are most profusely and elegantly
adorned with violet plumose setz.

MEcYNOCERA, n. gen.

Cephalothorax 6-jointed; head united with thorax; abdomen
4-jointed in the male, 3-jointed in the female. Anterior antenne
23-jointed, very long... Outer branch of posterior antenne 3-
jointed, the inner branch 7-jointed. Mandibles and maxille
well developed. Anterior foot-jaw 3-jointed and very muscular,
and, like the posterior, clothed with strong plumed sete. Swim-
ming-feet alike in both sexes, fifth feet entirely wanting.

Mecynocera Cravsi,n. sp. (Pl. XI. figs. 1-4.)

Length ;; inch. Rostrum bifid and very slender. Anterior
antenne 23-jointed, alike in both sexes, about twice the length
of the entire animal (fig. 1); the 11th, 12th, and 18th joints of
left antenne only edged with fine saw-teeth (fig. 2). Basal
portion of both antenne bear several short setz and a few long
ones, and at intervals, and especially at apex, are several long
whip-like sete. Posterior antenne very muscular; the outer
branch 3-jointed, the inner 7-jointed and terminated by spread-
ing sete. Mandibles finely toothed. Maxille well developed,
with two spreading setiferous branches and broad rounded palp.
Anterior foot-jaw 3-jointed, and, like the posterior, which is
small, bearing a large number of plumose sete.

Swimming-feet (fig. 3) have small hooked spines at ends of
joints, but no terminal spines besides the strong spinous sete.

MADEIRA AND THE CANARY ISLANDS. L151

Fifth feet wanting in both sexes. Abdomen of male (fig. 4) 4-
jointed, female 3-jointed, the basal joint of latter being somewhat
heart-shaped, and having doubled circular genital apertures.
The third abdominal joint is divided longitudinally, the inter-
vening span being filled with hyaline membrane. Caudal ter-
minations in female divergent, in male less so; both terminated
by four plumous setz on each side. Colour reddish brown.

Males and females were both plentiful, and taken by surface
tow-net at all the places visited. Indeed this species seemed to
be more widespread than any other.

Claus, in his memoir ‘ Die freilebenden Copepoden’ &c., gives
a figure (pl. xxxu. fig. 17) of a form which is evidently this species,
although the anterior antennw are net correctly represented.
He has informed me in a letter that the material at his disposal
was not sufficient to enable him to describe the species. I have
therefore much pleasure in naming this form, which is so
abundant at the Canary Islands, after the distinguished natu-
ralist who first discovered it.

Family CYCLOPIDZ.
OITHONA CHALLENGERI, Brady.
O. spINnrFRONS, Boeck.
O. PLUMIFERA, Dana.

O. sETIGER, Dana.

All four species of this delicately organized genus were found
fairly plentifully and generally distributed. The long feathery
red-coloured plumes of O. plumifera give the species a very grace-
ful appearance, and realily distinguish it from the others,

Family HARPACTICID 2.
LONGIPEDIA CoRONATA, Claus.

SETELLA GRactnis, Dana.

Various specimens of this species collected differed consider-
ably from each other in colour and form, but not sufficiently to be
considered specific.

EUTERPE GRACILIS, Claus.

This minute species, hitherto considered very rare, we found
widely distributed but not numerous.

i3”

152 MR. I. C. THOMPSON ON THE COPEPODA OF

Detavatia roBusra, Brady § Robertson.

DIOSACCUS TENUICORNIS, Claus.
One specimen only found at Orotava, Teneriffe.

LAOPHONTE OCURTICAUDATA, Boeck.
L. SERRATA, Claus. |
DacryLopus TIsBorpEs, Claus.

THatestris Mysts, Claus. | All found generally

T. nuFocincta, Norman. > distributed, but none

|
WESTWOODIA NOBILIS, Baird. | plentiful.

HLARPACTICUS CHELIFER, JZuller.
PORCELLIDIUM VIRIDE, Philippr. |

Ipya FuRcATA, Baird. J

MacHAIROPUS, 0. gen.

Head longitudinally rounded, terminating in long thin bifid
rostrum. Anterior antenne 12-jointed, about two thirds the
length of cephalothorax; sparingly setiferous. Posterior an-
tenne long; the outer branch 3-jointed, the inner 2-jointed.
Mandibles have very fine claw-like teeth; the palpi is two-
branched, with long fine hairs. }

Maxillw have four strong serrated curved spines and numerous
sete. Posterior foot-jaw 2-jointed, having long sword-like spines
with enlarged bases. Anterior foot-jaw small. Abdomen 2-
jointed.

MacHAIRrOPUS SANCTE-CRUCIS, n. sp. (PI. XII. figs. 1-5.)

Length ;4, inch. Rounded head (fig. 1) of a deep pink colour.
Anterior antenne (fig. 2) 12-jointed. Posterior antenne (fig. 3)
has long whip-like set, extending to the length of the anterior
antenne. Swimming-feet (fig. 4) have three joints to outer
branch, and two joints to inner. Terminal spine narrow and
finely serrated. Basal joints of swimming-feet have long spine
on inner margin. Fifth feet (fig. 5) have one joint terminated by
a long, stout, curved spine, with a small one at each side. First
joint of abdomen is small, the second long, and wide in the
middle. Caudal segments are about four times as long as broad,
and are terminated by short non-plumose sete.

MADEIRA AND THE CANARY ISLANDS. 153

One specimen only, sex undetermined ; was taken by tow-net
at Santa Cruz, Teneriffe.

ScUTELLIDUM TISBOIDES, Claus.

Family CORYCATID &.

Corycaus varius, Dana.
C. peLLucipuUs, Dana.

C. rimBatus, Brady.

C. venustus, Dana.

C. spectiosus, Dana.

All five species of Coryceus we found in fair abundance and
widely distributed.

CoPrmLiIA MIRABILIS, Dana.
Found very sparingly.
LUBBOCKIA SQUILLIMANA, Claus.

Several specimens of both sexes of this striking species were
taken at Teneriffe and Grand Canary.

~-Oncma optusa, Dana.

The remarkable variety of colour in this species is worthy of
note, especially when fresh. It ranges from olive-green to
purple, blue, magenta, scarlet, &c. The colours are mostly
retained if mounted in Farrants medium soon after capture.

SAPPHIRINA INHQUALIS, Dana.
Fairly plentiful throughout

S. serrata, Brady. Pe oralinarinds
gs.

S. METALLINA.

Family ARTOTROGID.

ACONTIOPHORUS ANGULATUS, n. sp. (PI. XII. figs. 6-11, and
Pl. XIII. figs. 5, 6.)

Length, inch. Head united with first thoracic somite, which
is rounded off at base (Pl. XIT. fig. 6), the other somites of cephalo-
thorax being angular. Rostrum (fig. 7) composed of two small
projections with finely serrated edges. Anterior antenne
(fig. 8) 11-jointed, short, avd gradually tapering from base to
apex; second joint larger than the others, and from it proceed
a number of long sete ; from seventh joint proceeds a long, narrow
spine; there are several long sete at the apex. Posterior

154 MR. I. C. THOMPSON ON THE COPEPODA OF

antenne (fig. 9) two-branched, the first being 3-jointed and
terminated by two long lancet-shaped spines; the second has
one joint, and is terminated by a long sword-shaped spine.
First and second foot-jaws (figs. 10 & 11) each have a long apical
claw, the first is composed of two, the second of four joints.

First four pairs of swimming-feet nearly alike, both branches 3-
jointed (Pl. XIII. fig. 5); base and second joint of chief branch have
very fine serration on inner edge formed by minute hairs. Fifth
feet (Pl. XITI. fig. 6) are composed of two stout joints, the outer
bearing a number of long setx, some of them plumose ; the outer
edge of both joints has fine hairy serration. Caudal segments and
terminal sete are the same as in A. scwtatus.

Siphon very long and slender, reaching to the ends of caudal
segments.

One specimen only of this new species was taken at Funchal
Bay, Madeira. The angular shape of the posterior segments of
the cephalothorax readily distinguish it from A. scutatus, which
it resembles in general characters.

Arrotroeus Normant, Brady &§ Robertson.
Found very sparingly.

CYMBASOMA, 2D. gen.

Cephalothorax elegantly boat-shaped, 4- to 6-jointed; head
united with thorax. Anterior antenne 5- or 6-jointed, broad
and muscular. No posterior antenne or foot-jaws. First four
pairs of swimming-feet have a large basal joint, the two branches
being each 3-jointed’ and terminated by strong spinous sete.
Abdomen in female 2-jointed, the first much larger than the
second ; and from the centre of the latter are attached the ova on
long narrow branches.

CYMBASOMA RIGIDUM, n. sp. (Pl. XIII. figs. 1-4.)

Length § inch. First joint of cephalothorax (fig. 1) equal in
length to the succeeding five, the last being much smaller than
the others, and appearing as though part of the abdomen. Eyes
long and kidney-shaped, with conspicuous brain below. Ante-
rior antenne (fig. 2) about eight times as long as broadest part,
and terminated by two claw-like spines; the inner side of each
antenna has five hooked spines (fig. 2). From two raised circum-
ferences near the centre proceed long spinous sete, and several
smaller sete are situated near the apical portion of the antenne.

Mintern imp.
Sp

Linn. Soc. Jaunn. Zoot. Vor. XX. Pi.X.
_CANDACE BREVICORNIS

7-8

CANDACE NIGROCINCTAn sp.

Linn. Soc.Jdourn. Zoor,. Vor. XX. Ph. XI.

Tite cee ieee Loon. von Pe ap

7 Mintern imp.
11-5. MACHAIROPUS SANCTA-CRUCIS nsp
6-11. ACONTIOPHORUS ANGULATUS,n.sp,

Lum. Soc. Journ. POOE, VOL Dk, Peo

a3 eypppitlt

/ H
j Pe |
nm del. Mintern Bros. ump,
BASOMA fF iGIDUM.n. su. 5-6. ACONTIOPHORUS ANGULATUS.n sp.

:
:
.
:
;

MADEIRA AND THE CANARY JSLANDS. 155

First four pairs of swimming-feet (fig. 8) have strong muscular
basal joints, from which proceed two 8-jointed branches with thick
terminal sete. Fifth pair of feet each composed of a long
joint, at the apex of which are three strong spinous seta. Abdo-
men very muscular; the caudal sete are short, broad, and diver-
gent, and are each terminated by three strong sete.

One specimen only (a female) was taken in the tow-net at

Orotava, Teneriffe. The lower part of the body is of a deep

sepia colour. The animal is remarkably rigid and muscular.

Quite recently a second form, evidently belonging to this new
genus, has been found in the channel between Puffin Island and
Anglesea. It is probably a male, and I am describing it under
the name Cymbasoma Herdmani. It will be figured along with
other new species from Liverpool Bay in the second volume of
the ‘ Proceedings of the Liverpool Biological Society.’

e@
EXPLANATION OF THE PLATES,
PuatTe X.

. Candace nigrocincta, n. sp.,male. Xx 250.

. Posterior antenna of ditto. x 400.

. First foot-jaw of ditto. x 400.

. Fourth pair swimming-feet of ditto. x 400.
Fifth pair swimming-feet of ditto. x 400.

. Abdomen and caudal segments of ditto. x 400.
. Candace brevicornis, n. sp., female. x 250.

. Fourth swimming-foot of ditto. x 400.

Fig.

CIR TP ow de

Puate XI.

Fig. 1. Mecynocera Clausi,n. sp., female. Xx 250.
2. Eleventh, twelfth, and thirteenth joints of left anterior antenna of ditto,
showing saw-teeth. x 400.
3. Fourth swimming-foot of ditto. x 400.
4. Abdomen and caudal segments of male ditto. x 400.

Puate XII.
. Machairopus sancte-crucis, 0. sp. * X 250.
. Anterior antenna of ditto. x 400.
. Posterior antenna of ditto. x 400.
. Fourth pair swimming-feet of ditto. x 250.
Fifth swimming-foot of ditto. x 250.
. Acontiophorus angulatus, n. sp., female, x 250.
. Rostrum of ditto. x 400.
. Anterior antenna of ditto. x 400.

Fig.

CNW h Od

156 MR. J. S. BALY ON SOME GENERA

Fig. 9. Posterior antenna of ditto. x 400.
10. Anterior foot-jaw of ditto. x 400.
11. Posterior foot-jaw of ditto. x 400.

Puate XIII.

Fig. 1. Cymbasoma rigidum, n. sp., female. x 250.
2. Anterior antenna of ditto. x 400.
3. First swimming-foot of ditto. x 400.
4. Abdomen of ditto, with attached ova and caudal appendages.
5. First pair of swimming-feet of Acontiophorus angulatus. x 400,
6. Fifth foot of ditto. x 400.

Descriptions of some Genera and Species of Galerucine.

By Josreru 8S. Baty, F.LS.
[Read 2nd February, 1888.]}

THE examination of some extensive genera of Galerucine—Haplo-
sonyx, Cerotoma, and others—in which the anterior acetabula
are described as closed or entire, has convinced me that the views
expressed by myself in the Ent. Month. Mag. vol. xxiii. p. 268
are correct. In any long series of individuals belonging to these
various genera I have almost invariably found some species in
which, whilst certain specimens have the acetabula closed, others
have them open, in some instances these opposite states occurring
on the different sides of the same individual. This is strikingly
the case in Cerotoma and Monolepta. Under such circumstances
it must, I think, be conceded that the state of the acetabula
cannot be regarded as a primary, even if it can be retained as a
secondary character. It becomes therefore necessary to dis-
cover some other and more stable means by which to divide the
Galerucine into primary sections. This, | venture to suggest,
will be found in the form and structure of the hinder portion of
the prosternum. In a large number of genera, for instance
Haplosonyx, Gnidea, and numerous others, the prosternum, as in
the Chrysomelide and the earlier groups of the Phytophaga, is
produced just before its base into a distinct lobe or process, the
sides of the lobe (in cases where the acetabula are closed) being
connected with the apices of the epimera, whilst the hinder
margin of the lobe is free, and usually applied to the anterior
surface of the mesosternum. In a second large section of the
family, represented by Galeruca, Aulacophora, &c., this subbasal
lobe is entirely absent, the prosternum being connected without

AND SPECIES OF GALERUCINE. 157

break with the hinder margin of the antepectus, or true hinder
border of the under surface of the thorax.

In this group the acetabula are sometimes closed by the pro-
longation of the epimera across the base of the prosternum to
meet in the median line, as in Galeruca and Menippus. On the
presence or absence of the subbasal lobe, I would divide the
family into two primary groups, taking as secondary ones those
derived from the spines at the apices of the tibie, the bifid or
appendiculated claws, and from the relative length of the elytral
epipleure. My object in making these remarks is to justify the
use of characters drawn from the form of this basal lobe as a
diagnostic character in the new genera described in the present
paper.

Of the genera mentioned or described below, the first five have
the prosternum lobed ; in the rest the lobe is obsolete.

Genus CHARIDEA.

Corpus elongatum, modice convexum. Caput exsertum, oculis rotun-
datis ; encarpis elongatis, pyrifurmibus ; antennis filifurmibus ; palpis
maxillaribus articulo ultimo ovato. Thorax transversus, convexus,
disco calloso. Scutellum trigonatum. lytra parallela, modice trans-
versim convexa, punctato-striata, interspatiis interdum costatis;
epipleuris fere ad apicem extensis. Pedes robusti, simplices; tibiis
apice muticis ; metatursi articulo basali ad duos sequentes fere aequi-
longo. Prosternum inter coxas distincte visum, ante basin loba-
tum; acetabulis anticis integris aut paullo apertis.

Type Charidea punctato-striata, Motsch.

This genus may be known from Galerucella, in which the
typical species was placed by Motschulsky, by the lobed base of
the prosternum ; it ought to stand near Gidea, but is separated
from that genus by the punctate-striate and costate elytra, and
by the non-incrassate palpi.

Although the two species placed by me in this genus differ in
one having closed, the other open acetabula, they agree so entirely
in all other characters that I have no hesitation in placing them
under the same generic head.

1, CHanipEa Puncrato-striata, Motsch. Etud. Ent. ix. p. 25
(sub Galeruca).
Galeruca multicostata, Jacoby, Proc. Zool. Soc. November 1885, p. 146,
tab. xlvi. fig. 7.
Hab. Japan.

In this species the anterior acetabula are slightly open.

158 MR. J. S. BALY ON SOME GENERA

2. CHARIDEA ForruNeEt. Elongata, picea, nitida, antennis nigris,
supra subnitida, pube suberecta grisea sparse vestita; thorace elytris- ©
que sanguineis; thorace quam longo paullo latiore, fortiter punctato,
disci medio bicalloso, callis levibus; elytris parallelis, transversim
convexis, fortiter punctato-striatis, utrisque bicostatis. Long.
23 lin.

Hab. Northern China; collected by the late Mr. Fortune.

Head clothed with griseous hairs; vertex convex, separated
from the frout by a deep transverse groove; encarpe contiguous,
pyriform, longer than broad, extending downwards as far as the
insertion of the antennsx, interanteunal space excavated; lower
portion of clypeus transversely excavated ; antenne nearly as
long as the body, moderately robust, filiform, the second joint
short, the third nearly twice its length, the fourth and succeeding
ones each equal in length to the third. Thorax broader than
long; sides straight and nearly parallel from the base to beyond
the middle, thence slightly rounded and obliquely converging
towards the apex, the anterior angle obtuse, the hinder one acute ;
upper surface moderately convex, coarsely and strongly punc-
tured; on the disk are two large, slightly raised smooth callo-
sities, separated from each other by a narrow groove, but bounded
at the base, apex, and the outer margin by a deep irregular sulca-
tion; surface of disk more or less stained with piceous. LElytra
rather broader than the thorax, parallel on the sides, transversely
convex, not excavated below the basilar space ; each elytron with
twelve rows of large, deeply impressed punctures; the fourth
interspace at its base, the fifth and ninth for nearly their whole
length, confluent at their apices, strongly costate ; upper portions
of the tenth and eleventh interspaces subcostate.

This species may be known from C. punctato-striata by the
pubescence on the upper surface of the body, by the strongly
raised coste on the elytra, and by the piceous colour of the
head, under surface, and legs. The anterior acetabula in this
insect are entirely closed.

Genus Praryxantua, Baly.

1, PLaTyXANTHA CLYPEATA. Elongata, paraliela, dorso deplanata,
flava, nitida, thorace, mandibulis apice oculisque nigris ; thorace
transverso, lateribus e basi ad ultra medium rectis, paullo divergenti-
bus, hinc apicem versus convergentibus ; disco modice convexo, levi,
utrinque fovea parva, plus minusve distincta, impresso ; elytris paral-
lelis, ad latera convexis, dorso deplanatis, tenuiter confuse punctatis ;

AND SPECIES OF GALERUCINA. 159

nigris, plaga magna communi, subquadrata, prope medium posita,
flava. Long. 4 lin.

Var. A. Elytris flavis, basi et ad apicem piceo-tinctis.

Var. B. Elytris totis flavis.

Mas. Labro leviter incrassato, margine antico bilobato ; clypei dimidio
antico transversim excavato, margine antico medio bidentato ; abdo-
minis segmento anali trilobato, lobo intermedio fere plano, transversim
quadrato.

Fem. Labro non incrassato, margine antico leviter sinuato ; clypei disco
antico non excavato, margine antico imermi; abdominis segmento
anali apice extremo obtuso.

Hab. Andaman Islands.

Antenne equal to the body in length, filiform in both sexes,
rather more robust in the g; the basal joint slightly thickened,
subclavate, the second very short, moniliform, the third and fol-
lowing ones each equal in length to the basal joint; maxillary
palpi with the third joint obconic, not distinctly thickened in
the ¢, the apical joint conic, acute; lower disk of the clypeus
in the ¢ transversely concave, its anterior margin armed in the
middle with two small acute teeth; lower disk in the 9, its
auterior border excepted, thickened, its lower edge unarmed.
Thorax more than half as broad again as long; sides straight and
diverging from the base to beyond the middle, thence rounded and
converging towards the apex,the anterior and posterior angles each
armed with an obtuse tooth ; upper surface moderately convex, im-
punctate ; disk impressed on each side with a small, more or less
distinct fovea. Scutellum large, trigonate. Hlytra broader than
the thorax, parallel ; upper surface convex on the sides, flattened
along the suture, finely and irregularly punctured; interspaces
(when seen under a deep lens) closely covered with minute
punctures.

The above insect differs from the typical species in having the
antenne in the ¢ entirely filiform.

Genus GinrpEa, Baly.

1, GQENIDEA CHRULEIPENNIS. Oblongo-ovata, postice ampliata, rufo-
testacea, nitida, antennis (articulo basali excepto) flavis, elytris metal-
lico-ceruleis ; thorace profunde transversim excavato; elytris ob-
longis, infra basin transversim depressis, tenuiter punctatis. Long.
3 lin. .

Fem. Clypeo transversim sulcato.

Hab. Siam.

169 MR. J. S. BALY ON SOME GENERA

Antenne very slender, filiform, equal to the body in length,
the second joint very short, the third and fourth each equal in
length to the first ; front impressed just above the encarpe with
a deep fovea; encarpe thickened, trigonate; clypeus concave,
transversely srleate Thorax nearly twice as broad as long,
sides straight and slightly diverging from the base to beyond the
middle, thence obliquely rounded to the apex; anterior angles
slightly produced, obliquely truncate, the hinder ones acute ;
upper surface convex, very minutely punctured; disk impressed
with a broad transverse excavation, which terminates at some
distance from the Jateral margin, but is more deeply excavated at
each end. LElytra oblong, convex, transversely excavated below
the basilar space, and again obsoletely on the outer disk about
its middle, the humeral callus thickened; surface finely punc-
tured.

2. Ginipea JacoByr. Anguste oblonga, postice paullo ampliata,
convexa, nigra, nitida, labro antennisque flavis aut fulvis, pectore
abdomineque rufo-piceis ; thorace transversim sulcato, sulco utrinque
fortiter foveolato ; elytris oblongis, infra basin transversim excavatis,
fere impunctatis, rufis, apice plus minusve nigris. Long. 4—45 lin.

Mas. Clypeo transversim concavo ; labro ampliato, coneavo ; abdominis
segmento anali trilobato, lobo intermedio ad apicem leviter excavato.

Fem. Clypeo transversim sulcato; labro paullo incrassato.

Var. A. Labro nigro aut nigro-piceo ; elytris totis nigris.

Hab. Penang; Malacca; Borneo, Sarawak; Sumatra (Wal-
lace).

Antenne slender, equal to the body in length, and clothed with
erect hairs in the ¢, rather shorter in the 2; clypeus in the
3 transversely trigonate, concave; labrum in the same sex
dilated, rather deeply concave ; clypeus in the Q deeply sulcate
transversely, the labrum thickened. Thorax rather more than
one half broader than long ; sides straight and slightly diverging
from the base to beyond the middle, thence converging to the
apex; disk deeply excavated transversely, the sulcation termi-
nating some distance within the lateral margin, more deeply
excavated at each end. Jiytra oblong, convex, transversely
excavated below the basilar space, very minutely punctured, the
punctures only visible under a lens.

. .

AND SPECIES OF GALERUCINE. 161

3. CENIDEBA PALLIPES, Fabr. Syst. El. i. p. 479.—Subelongata, postice
vix ampliata, rufo-testacea, nitida, oculis, elytrorum apice, abdomine
pedibusque posticis (tarsis sordide fulvis exceptis) nigris; antennis
labroque flavis; thorace transverso, disco sat profunde transversim
suleato, sulco utrinque magis fortiter excavato; elytris oblongis,
infra basin leviter transversim depressis, tenuiter punctatis. Long.
34 lin.

Mas. Facie inferiore profunde excavata, clypeo utrinque spinis duabus
elongatis armato; labro ampliato, trilobato, lobis retrorsum spectan-
tibus, duobus externis compressis, trigonatis, intermedio lineariform1 ;
abdominis segmento anali utrinque emarginato.

Fem. Clypeo transversim concavo.

Hab. Sumatra (Sir Stamford Raffles).

Maxillary paipi in the ¢g with the third joint strongly
thickened and enlarged; the fourth short, nearly buried in the
third; clypeus in the ¢ deeply excavated, quadrispinose ; the
upper pair of spines acute, the lower two incurved, clothed with
coarse hairs; labrum in the same sex thickened, its hinder
margin trilobate, the outer lobes trigonate, acute, the middle one
narrow, obtuse; clypeus in the @ transversely sulcate; antenne
filiform in both sexes. Thorax twice as broad ag long; sides
straight and diverging from the base nearly to the apex; disk
finely but remotely punctured, transversely excavated, the sul-
cation more deeply impressed on each side, abbreviated before
reaching the lateral margin. LElytra finely punctured.

This insect in v. Harold’s Catalogue stands as a variety of
Haplosonyx sumatre.

Genus Crnormta, Baly.

1. CyNORTA OCELLATA. Anguste elongata, parallela, flava, subnitida,
oculis nigris; thorace quam longo paullo latiore, disco trifoveolato ;
elytris parallelis, longitudinaliter costatis, interspatiis subseriatim
punctatis. Long. 2}-23 lin.

Mas. Facie inferiore profunde excavata, quadrituberculata, tuberculis
duobus posticis conicis, duobus anticis compressis.

Hab. Malacca, Singapore ( Wallace).

Hyes large, black ; antenne very slender, filiform, slightly
exceeding the body in length; third joint equal in length to the
first, slightly longer than the feurth ; lower face deeply excavated
in the dg, armed with four tubercles—two, one on each side,
immediately below tle insertion of the antenna, conic, and two
compressed, placed above the anterior margin of the clypeus,

162 MR. J. S. BALY ON SOME GENERA

one on each side near the outer border. Thorax about one half
broader than long; sides straight and slightly diverging from
the base nearly to the apex; anterior angle slightly produced,
obtuse, hinder angle armed with an obtuse lateral tooth; upper
surface impressed with a large, shallow, trilobate fovea, which
covers nearly the whole central disk. Elytra parallel, convex ;
each elytron with seven or eight distinct longitudinal coste,
their interspaces finely punctured.

I possess two specimens of this species, both belonging to the
3 sex.

2. CYNORTA APICIPENNIS. Subfiliformis, parallela, flava, oculis, ely-
trisque apice nigris; thorace quam longo vix latiore, disco leviter
trifoveolato; elytris elevato-costatis, imterspatiis punctatis. Long.
3 lin. ;

Fem.? Antennis longitudine corporem paullo superantibus; clypeo
excavato, concavo.

Hab. Sarawak (Wallace).

Narrowly elongate, parallel. Head strongly exserted ; clypeus
concave, smooth; maxillary palpi robust, the upper two joints
conjointly ovate; antenne slender, filiform, longer than the body.
Thorax rather broader than long; sides slightly diverging and
sinuate from the base to beyond the middle, thence converging
towards the apex; upper surface impunctate, impressed with a
large shallow trifid excavation, which covers a considerable
portion of the surface. Hlytra parallel, subcylindrical; each
with eight or nine longitudinal costz, their interspaces distinctly
punctate. The abdomen in the only specimen before me is
shrivelled, so that I am unable to decide with certainty as to its
sex, but from the non dilated basal joint of the anterior tarsus I
believe it to be a female.

3. CYNORTA FACIALIS. Subfiliformis, parallela, flava, oculis elytrisque
apice nigris, abdominis apice nigro-piceo; thorace quam longo vix
latiore, disco leevi, pone medium bifoveolato; elytris parallelis, leviter
elevato-costatis, interspatis distincte punctatis. Long. 23 lin.

Mas. Clypei dimidio postico trigonato, flavo, medio fovea rotundata
impresso ; dimidio antico depresso, transversim concavo.

Hab. Sarawak: a single specimen (Wallace).

Clypeus with its upper half not depressed, trigonate, its

surface plane, impressed in the middle with a small round fovea ;
the lower half strongly depressed and separated from the upper

_—_

‘ft. es.

|
|

AND SPECIES OF GALERUCINE. 163

portion by a sutural line, its surface transversely concave ;
antenne filiform (the five upper joints in the solitary specimen
under examination broken off). Thorax scarcely broader than
long; sides slightly diverging and slightly sinuate from the base
to beyond the middle, thence rounded and converging towards
the apex; upper surface convex, smooth, impunctate, hinder
disk impressed on each side with a large shallow fovea. Elytra
parallel, subeylindrical ; each with eight or nine slightly elevated
cost, the interspaces distinctly punctured.

The peculiar formation of the clypeus in this species closely
resembles that of the same segment in Cheilovena. As the pecu-
larity in the present instance is, however, probably only sexual,
the removal of the insect from the genus in which I have placed
it will not be justified until the other sex is known.

Genus Monorepta, Hrichson.

This natural group, founded by Erichson in 1848 on an African
species, IZ. pauperata, has been subsequently divided by Chapuis,
myself, and others into smaller generic groups, on characters
derived chiefly from the open or closed state of the anterior
acetabula and on the length of the epipleure. I have already
stated my reasons for considering the first of these characters to
be in a great measure unsatisfactory, and at any rate one not to
be depended on by which to divide the Galerucine into primary
sections. In Monolepta (taken as a whole) the lateral angles of
the subbasal lobe of the prosternum are well developed, and in
the great majority of species join the apices of the epimera to
close the anterior acetabula; in’some instances, however (Lupe-
rodes alboplagiatus, &c.), the epimera are abbreviated before
reaching the sides of the lobe, and consequently the acetabula
remain distinctly open; in a third set (Ochralea) the acetabula
are found to be both closed and widely open in the same species, |
every intermediate stage occurring between the two extremes *.

The second, of great value when well defined and when really
terminating at a given point, as in Aulacophoraand other genera,

* In Luperodes preustus and L. discrepens, two insects which differ from the rest
of the genus in their oblong, not ovate form, the sides of the subbasal lobe are
not produced, the lobe itself being either narrowly wedge-shaped or noduliform ;
the epimera are also much abbreviated, leaving the acetabula broadly open.
These species should, I think, be retained in Luperodes.

164 MR. J. 8. BALY ON SOME GENERA

fails in the present group for the following reason. In many
species of MMonolepta (including the typical one) the elytral
epipleura gradually narrows towards the apex and extends back-
wards for nearly the whole length of the outer margin of the
elytron ; in others, the epipleura apparently ends at or about the
middle of the margin; on examination, however, it will be seen
that it does not terminate at that point, but although much
narrowed extends to a greater or less extent along the hinder
margin. The point at which the contraction takes place also
varies greatly, so that I have thought it well to place all the
species here described in Monolepta.

1. MONOLEPTA HAMORRHOIDALIS, Fabr. Syst. El. i. p. 490; Oliv.
Ent. vi. p. 629, t. 3. fig. 33.—Anguste ovata, postice paullo
amphata, convexa, flava aut rufo-flava, nitida, antennarum articulis
intermediis et apicalibus nigris aut nigro-piceis ; thorace quam longo
distincte latiore, convexo, lwvi; elytris convexis, nigris, apice plus
inmmusve sanguineis. Long. 23-3 lin.

as. Elytro utroque ante medium fovea magna subrotundata, margine
incrassata munito.

Hab. Australia; New Guinea, Mysol, Sulu Islands.

Antenne filiform, the third joint about one half longer than
the second ; the three lower joints flavous, the eighth, ninth, and
tenth, together with the basal portion of the eleventh, flavous; the
upper portion of this last, together with the fourth to the seventh,
black, or nigro-piceous. Thorax about one fourth broader than
long ; sides rounded, nearly straight and slightly diverging be-
hind the middle, anterior angle armed with an obtuse tubercle ;
disk convex, smooth. Llytra oval, convex, slightly excavated
below the basilar space in the 2, minutely punctured; black,
the apical portion to a greater or less extent bright sanguineous ;
each elytron in the ¢@ is impressed with a large rotundate-ovate
fovea placed on the anterior disk about halfway between the outer
margin and the suture, and extending downwards to a little below
the median line. Elytral epipleure extending below the middle.
Anterior acetabula closed.

I possess two specimens of this species (both males) from
Australia, the original locality given by Fabricius and Olivier,
one, if not both, having formed part of the collection made during
the voyage of the ‘ Coquille.’ In these specimens the sanguineous
coloration of the elytra is confined to the extreme apex, but in
those from the Malay Archipelago, collected by Mr. Wallace,

AND SPECIES OF GALERUCINE. 165

the red colour covers xearly the whole hinder disk, in some
individuals even extending upwards beyond the median line.

2. Mono.tepra CuminGiI. Anguste ovata, postice paullo ampliata,
convexa, rufo flava, nitida, antennarum articulis intermediis ultimique
apice, tibiis tarsisque nigris ; thorace quam longo vix dimidio latiore,
subcylindrico, disco paullo deplanato; elytris minute punctatis, nigris,
utrisque macula parva basali dimidioque postico sanguineis. Long.
3 lin.

Mas. Elytro utroque callo subhumerali, hoc medio fovea rotundata,
profunde impresso, instructo.

Hab. Philippine Islands, Manilla.

Encarpe and lower face pale flavous. Antenne filiform, the
third joint twice the length of the second; the three lower ones
rufo-flavous, the fourth to the seventh black; the four outer
joints, the black apex of the apical one excepted, white. Thorax
nearly one half broader than long; sides straight and very
slightly diverging from the base to beyond the middle, thence
slightly converging to the apex, the hinder angle acute, the
anterior one slightly produced into an obtuse tubercle; disk
subcylindrical, very minutely punctured, the puncturing only
visible under a strong lens. Elytra convex, not excavated below
the basilar space, finely and rather closely punctured. Hach
elytron in the ¢ with a large subhumeral callus, in the centre of
which is a round fovea.

The narrow subcylindrical thorax in both sexes will at once
separate the present species from its allies; the ¢ also may be at
once known by the small size and different shape of the elytral

fovea.

3. MoNnoLEPTA FOVEICOLLIS. Ovata, postice ampliata, convexa,
flava, nitida, pedibus rufo-testaceis, antennarum articulis intermediis
nigris ; thorace transverso, disci medio transversim excavato ; elytris
tenuiter punctatis, nigris, apice sanguineis. Long. 3} lin.

Mas. Elytro utroque fovea magna ovata, disco externo ante medium
posita, profunde impresso.

Hab. Batchian (Wallace) : a single specimen.

_ Antenne filiform, the third joint nearly one half longer than the

_ second ; the fourth to the seventh black, the rest flavous. Thorax

_ nearly twice as broad as long; sides obliquely diverging from

__ the base to the middle, thence rounded and converging to the apex,

_ apical angle produced into a flattened obtuse tubercle; disk

LINN. JOURN.—ZOOLOGY, VOL. xx. 14.

166 MR. J. 8S. BALY ON SOME GENERA

transversely convex, very minutely punctured, its middle third
with a broad but shallow transverse depression, which is rather
more deeply excavated on either side of the median line. Elytra
convex, not excavated below the basilar space, minutely but not
closely punctured ; outer disk of each elytron in the g with a
large oval fovea, which commences at some little distance below
the humeral callus and extends downwards as far as the middle
line. Elytral epipleure extending for a short distance below the
middle. Anterior acetabula closed.

4. MonoLEpTA TERMINATA, Guér. Voy. de la Coquille, ii. Zool. p. 149.
—Anguste ovata, postice paullo ampliata, convexa, rufo-testacea, nitida,
tibiis tarsisque nigris aut nigro-piceis, antennis pallide flavis, articulo
ultimo nigro; thorace quam longo latiore, convexo, levi; elytris
convexis, tenuiter punctatis; nigris, apice sanguineis. Long. 23-
3 lin.

Mas. Elytro utroque ante mediun fovea magna subovata male definita,
extus late sed leviter elevato-marginata, fundo leviter excavata, im-
presso.

Hab. Java (Guérin), Borneo, Sumatra ( Wallace, Raffles).

Antenne filiform, the third jomt one half longer than the
second; pale flavous; the terminal and occasionally the upper
half of the penultimate joints more or less stained with black.
Thorax about one fourth broader than long ; sides nearly straight
and diverging from the base to the middle, thence rounded and
converging towards the apex, the anterior angles thickened and
produced, obtuse, hinder angles acute; hinder margin obtusely
rounded; disk convex, smooth, impunctate. Elytra oval, convex,
finely but not very closely punctured ; on the anterior disk of each
elytron in the ¢ (the only sex known to me) is a large, very
shallow and ill-defined excavation, the outer and hinder edges of
which are broadly but slightly thickened.

5. Monotepra Wa.uacer. Late ovata, postice paullo ampliata
convexa, sanguinea, nitida, tibiis tarsisque piceo tinctis, antennis
pallide flavis, basi flavo-testaceis, articulis duobus ultimis nigris,
abdomine flavo-rufo ; thorace quam longo latiore, convexo, levi;
elytris convexis, minute punctatis, nigris, apice sanguineis. Long.
3-33 lin.

Mas. Elytro utroque fovea magna subhumerali profunde excavata,
limbo incrassata impresso.

Hab. Celebes (Wallace).

————

(i 2

Se = eo ee |

AND SPECIES OF GALERUCIN®. 167

Antenne filiform, nearly equal to the body in length in the ¢,
the third joint nearly twice as long as the second; pale flavous,
the two lower joints flavo-testaceous, the two upper ones black.
Thorax about one third broader than long; sides obliquely di-
verging and obsoletely sinuate from the base to the middle,
thence rounded and converging to the apex, anterior angle pro-
duced into an obtuse tubercle, the hinder angle slightly produced,
acute ; disk convex, smooth, impunctate. LElytra convex, very
minutely punctured ; each elytron in the ¢ with a large ovate,
deeply excavated fovea, placed on the outer disk immediately below
the humeral callus and extending nearly to the middle, its entire
margin strongly thickened.

I possess five specimens of the above insect collected by
Mr. Wallace in Celebes: the broader form, together with the
position of the elytral fovea, will at once separate it from the
allied species; the elytral epipleure are also less abruptly nar-
rowed below the middle than in the other similarly-coloured
specific forms.

The five species described above are usually confounded in
collections under the name of hemorrhoidalis or terminata: the
structural differences, however, at once separate them.

The following short Table will assist in distinguishing the
species described above :—

1. Thorax impressed in the centre with a transverse fovea.
fovercollis.
2. Thorax not transversely sulcate.
a. Thorax only slightly broader than long .. Cwmingiz.
aa. Thorax distinctly broader than long.

b. Thorax one half broader than long .... hemorrhoidalis.
bb. Thorax one third broader than long.

o, Miytra oblong-ovate .............. terminata.

ec. Elytra broadly ovate ............ Wallace.

6. MoNOLEPTA VERTICALIS. Anguste ovata, postice paullo ampliata,
convexa, nigro-picea, nitida, capite, antennis basi, thorace pedibusque
anticis pallide flavis, pedibus posticis quatuor piceo-fulvis ; capitis
vertice fovea rotundata magna impresso ; thorace transverso, convexo,
levi; elytris tenuiter punctatis, utrisque maculis duabus, una infra
basin, altera prope apicem, flavis, ornatis. Long. 2 lin.

14*

168 MR. J.S. BALY ON SOME GENERA

Var. A. Elytris pallide flavis, utrisque limbo fasciaque prope medium

nigro-piceis, pedibus fulvis.

Var. B. Elytris fere totis flavis.

Hab. Malay Archipelago, Batchian, Aru and Sulu Islands
(Wallace).

Vertex impunctate, impressed with a large round fovea; antennz
three fourths the length of the body, second joint short, the third
slightly longer, four or five lower joints flavous or fulvous, the
rest nigro-piceous. Thorax nearly twice as broad as long; sides
rounded, the anterior and posterior angles produced, subacute ;
disk transversely convex, smooth, impunctate. Elytra oblong-
ovate, very slightly dilated posteriorly, convex, not depressed
below the basilar space, remotely impressed with fine punctures,
their interspaces very minutely punctured. Abdominal segments
in some specimens narrowly edged with flavous.

The outer margins of the flavous spots on the elytra are ill-
defined, being more or less stained with piceous.

Genus PARAULACA.

Corpus oblongum, convexum, postice paullo ampliatum. Caput exser-
tum; antennis filiformibus. Thorax transversus, dorso transversim
sulcatus. Elytra convexa, confuse vel subseriato-punctata; epi-
pleuris fere ad apicem extensis. Pedes simplices; ¢ibiis apice spina
brevi armatis; ungutculis appendiculatis. Prosternum inter coxas
elevatum, basi non lobatum ; acetabulis anticis apertis.

Type Paraulaca (Aulacophora) angulicollis, Motsch.

The above genus differs solely from Pseudocophora in having
appendiculate instead of bifid claws; occupying the samefrela-
tive position to that genus that Diacantha does to Aulacophora.
In both genera the prosternum is not lobed at its base, and the
apices of the epimera are free, not closing the acetabula.

Genus PsEupocopnora, Jacoby, Notes Leyd. Mus. vi. p. 69.

The males of the present genus (so far as they are yet known)
have a deep sutural fovea placed immediately below the basilar
space on the elytra ; this fovea, the edges of which are thickened,
is armed with two or four raised tubercles or teeth ; the number
and shape of these teeth, taken in conjunction with the form and

AND SPECIES OF GALERUCIN®. 169

sculpture of the median lobe of the anal segment of the abdomen,
afford good diagnostic characters for the separation of the males
of the various species. |

Specimens of all of the species described in the present paper
are contained in my collection. P. bicolor, Jacoby, Proc. Zool.
Soc. 1887, p. 111, trom Ceylon, is unknown to me.

The following Table will assist the student in working out the
species described below.

TaBE I. (Males).

A. Elytral fovea quadridentate.
a. Median lobe of the trilobate anal segment of the abdomen

plane.
b. Hinder half of the elytra (the outer margin excepted)
REI en et ee 1. Buquetic.
6b. Hinder half of elytra entirely black .... 2. distincta.
aa. Median lobe of anal segment of abdomen more or less
concave.
b. Median lobe slightly concave .......... 3. Hrichsoni.

bb. Median lobe deeply excavated.
e. Elytral fovea with the anterior pair of teeth bifid.
4. ambusta.
ec. Elytral fovea with the anterior pair of teeth simple.
5. Wallacet.

B. Elytral fovea bidentate.
a. Median lobe of anal segment of abdomen plane.

b. Breast and abdomen black ............ 6. uniplagiata.
bb. Breast and abdomen flavous .......... 7. flaveola.
aa. Median lobe of anal segment concave.... 8. brunnea.

TasBxeE II. (Females).

A. Pygidium entire.
a. Apex of anal segment of abdomen deeply and narrowly

METI 2 le Woa'e woolen os eels cease oe 9. pectoralis.
aa. Apex of anal segment narrowly angulate-emarginate.
1. Buqueti.

aaa. Apex of anal segment broadly subangulate-emarginate.
3. Hrichsone.

170 MR. J. S. BALY ON SOME GENERA

B. Apical margin of pygidium more or less deeply emarginate.
a. Apex of pygidium faintly emarginate .... 10. perplezxa.
aa. Apex of pygidium trifid, the middle lobe elongate, its

apical surface tuberculate_............ 8. brunnea.
aaa, Apex of pygidium trifid, the middle lobe only slightly
produced. | ..0). «ssh Ee 56a be 4. ambusta.

1. Psevpocopnora Buaquetii, Guér.

Galeruca Buquetii, Guér. Voy. Coquille, 1830, p. 143.

Oblongo-ovata, postice paullo ampliata, convexa, rufo-fulva aut fulva,
pectore, abdomine pedibusque posticis quatuor nigris, antennis pal-
lide flavis; thorace profunde transversim sulcato; elytris postice
ampliatis, convexis, infra basin in ? leviter transversim depressis,
distincte subseriato-punctatis, punctis pone medium minus fortiter
impressis, confuse dispositis; rufis, basi late limboque externo
nigris. Long. 23-3 lin.

Mas. Elytris infra basin fovea magna communi, fundo quadridentata

instructis ; abdominis segmento anali trilobato, lobo intermedio
piano.

Fem. Abdominis segmento anali apice rotundato, medio abrupte angu-
lato-emarginato.

Hab. Java.

Antenne pale flavous; eyes prominent, black. Thorax nearly
twice as broad as long; sides nearly straight and diverging from
the base to beyond the middle, thence obliquely converging to the
apex, anterior angle with an obtuse oblique tubercle ; disk deeply
suleate transversely, the sulcation usually entire, rarely inter-
rupted in the middle. Elytra oblong, dilated posteriorly, convex,
slightly depressed in the 2 below the basilar space, distinctly
subseriate-punctate, the spaces between the rows of punctures
more or less distinctly thickened ; the punctures on the hinder
disk finer and arranged without order. In the ¢ below the
basilar space is a large common fovea, surrounded by a thickened
margin, its surface armed with two pairs of teeth, the anterior
pair thickened at the base, their apices deflexed and poste-
riorly produced, the hinder one flattened, truncate, and abruptly
deflexed towards the suture; these Berne in some specimens are
concolorous with the surface of the fovea, in others they are

more or less fulvous; the anterior pair in some instances are
obsoletely bifid.

4
.

AND SPECIES OF GALERUCINE. 171

2. PSEUDOCOPHORA DISTINCTA. Late ovata, postice paullo ampliata,
convexa, nitida, subtus nigra, pedibus anticis piceo-fulvis; supra
piceo-fulva, antennis (basi exceptis) pallide flavis, elytrorum dimidio
postico et (in mare) fovea subbasali nigris ; thorace profunde trans-
versim sulcato ; elytris subseriatim punctatis. Long. 3 lin.

Mas. Elytris fovea magna subbasali, fundo quadridentata imstructis ;
abdominis segmento anali trilobato, lobo intermedio plano.

Hab. Borneo (Wallace) : a single specimen.

Antenne slender, filiform, three fourths the length of the body,
pale yellow, the two lower joints piceo-rufous. Thorax nearly
twice as broad as long; sides obliquely diverging and slightly
sinuate from the base to beyond the middle, thence obliquely
converging to the apex; disk deeply excavated transversely just
bebind the middle, the sulcation interrupted on the median line.
Elytra dilated from the base towards the apex, the latter broadly
rounded ; above convex, substriate-punctate on the anterior disk,
irregularly punctured behind the middle ; immediately below the
basilar space in the male (the only sex known to me) is a
large common fovea, surrounded by a thickened margin, its sur-
face armed with four teeth, the anterior pair acute, their apices.
slightly deflexed, the hinder pair flattened, truncate and abruptly
deflexed towards the suture.

This species is closely allied to P. Buquetit; in addition,
however, to its different coloration, it is much broader than that
insect.

3. PsEUDOCoPpHORA EricHsonI. |

Pseudocophora Buquetii, Jacoby, Notes Leyd. Mus. vi. p. 69 (pars).

Ovata, postice ampliata, convexa, nigra, nitida, capite, thorace scutello-
que fulvis aut rufo-fulvis, antennis pallide flavis, pedibus anticis.
piceis ; thorace quam longo plus dimidio latiore, sat profunde trans-
versim sulcato; elytris distincte subseriatim punctatis, punctis apicem
versus confusis, fere obsoletis ; flavo-fulvis, dimidio antico apiceque
extremo nigris. Long. 3-3 lin.

Mas. Elytris infra basin fovea magna communi quadridentata, den-
tibus duobus anticis obtusis, duobusque posticis conicis ; abdominis-
segmento anali trilobato, lobo intermedio leviter concavo, apice
rotundato.

Fem. Abdominis segmento anali apice emarginato.

Hab. Sumatra.

Antenne filiform, the third and fourth joints nearly equal in

length in both sexes, pale flavous, the basal joint fulvous, the

172 MR. J. S. BALY ON SOME GENERA

three apical ones stained with piceous. Thorax more than one
half broader than long; sides very slightly diverging and sinuate
from the base to beyond the middle, thence converging towards
the apex, the hinder angles subacute, the anterior ones slightly
produced, obtuse; disk transversely sulcate immediately behind
the middle, the suleation strongly impressed during its whole
length. Elytra oblong, dilated posteriorly, convex, distinctly and
rather strongly punctured, the puncturing nearly obsolete towards
the apex; the punctures on the anterior half of the inner disk
irregularly arranged in double rows.

The above species was confounded by Jacoby with Buquetiz,
Guér., on which he founded the genus; although similar in colo-
ration to that insect, it is well separated by the structural
characters in both sexes.

4, PseEuDocoPHoRA AmBusTA, Erichs. Nov. Act. Leopol. Carol. xv.
1834, Suppl. i. p. 272.—Oblongo-ovata, postice paullo ampliata,
convexa, fulva aut flava, antennis pallide flavis; pectore, abdomine,
pedibus posticis quatuor elytrorumque limbo externo nigris ; thorace
transversim sulcato; elytris subseriato-punctatis, nigro-limbatis.
Long. 23 lin.

Mas. Elytris infra basin fovea magna, nigra, margine elevata, fundo
quadridentata, dentibus duobus anticis bifidis; abdominis segmento
anali trilobato, lobo intermedio concavo.

‘em. Abdominis pygidio apice trifido, lobo imtermedio elongato
tuberculo conico instructo; segmento anali medio concavo-emar-
ginato.

Var. A, ¢. Elytrorum limbo externo nigro obsoleto.

Hab. Type, Luzon (Erichson) ; var. A, Celebes (Wallace).

Antenne pale flavous, the third joint twice the length of the

second, rather longer than the fourth. Thorax twice as broad as
long; sides slightly diverging and rather strongly sinuate from
the base to beyond the middle, thence obliquely converging to
the apex, the anterior angles slightly excurved; disk trans-
versely sulcate, the sulcation interrupted in the middle part of
its course. LElytra similar in form to those of P. Buquetit;
their surface less strongly punctured, the subcostate interspaces
being absent. The males may be known by the anterior pair of
teeth on the subbasal fovea being distinctly bifid, not single as in
the other species of the genus; the hinder pair are formed as in
P. Buquetii.

ta ee, ees. ee

‘al

yee

rate

AND SPECIES OF GALERUCIN &. 173

5. PsrupocopHoraA Wauuacel. Oblongo-ovata, postice paullo
ampliata, convexa, fulva, nitida, pectore pedibusque nigris, antennis
nigro-piceis; thorace transversim sulcato; elytris oblongis, postice
paullo ampliatis, convexis, subseriatim punctatis, punctis apicem
versus confusis. Long. 27 lin.

Mas. Elytris infra basin fovea magna, elevato-marginata, fundo quadri-
dentata, dentibus duobus anticis simplicibus; abdominis segmento
anali trilobato, lobo mtermedio basi concavo-excavato.

Hab. Bouru (Wallace).

Antenne nigro-piceous. Thorax nearly twice as broad as long ;
sides slightly converging and slightly sinuate from the base to
far beyond the middle, thence slightly converging to the apex,
the anterior angle acute; disk deeply sulcate transversely, the
sulcation less deeply excavated on the median line. LElytra
convex, subseriate-punctate anteriorly, irregularly punctured
behind the middle ; elytral fovea in the ¢ (the only sex known to
me) quadridentate ; the anterior pair of teeth simple, the posterior
pair compressed, incurved.

6. PSEUDOCOPHORA UNIPLAGIATA, Jacoby, Notes Leyden Mus. vi.
p- 214.—Oblongo-ovata, postice paullo ampliata, convexa, fulva,
nitida, pectore, abdomine pedibusque posticis quatuor elytrorumque
fovea subbasali nigris, antennis pallide flavis; thorace profunde trans-
versim sulcato; elytris distincte subseriato, pone medium confuse
punctatis. Long. 23-3 lin.

Mas. Elytris infra basin fovea magna communi nigra, elevato-marginata,
antice bituberculata instructis; abdominis segmento anali trilobato,
lobo intermedio oblongo, profunde excavato.

Hab. Sumatra; Siam.

Similar to P. brunnea, the elytra more strongly punctured, the

anal segment of abdomen of an entirely different form.

7. PSEUDOCOFHORA FLAVEOLA. Ovata, postice ampliata, flava,
nitida, thorace transversim sulcato; elytris tenuiter punctatis. Long.
23 lin. |

Mas. Elytris infra basin fovea magna communi antice bituberculata,
tuberculis obtusis, contiguis; abdominis segmento anali trilobato,
lobo intermedio plano, apice late rotundato.

Hab. Andaman Islands.

Antenne filiform ; eyes black. Thorax transverse ; sides from

the base to far beyond the middle nearly straight, diverging,

’ then obliquely converging towards the apex; disk transversely

sulcate, nearly impunctate. Elytra broadly oblong, dilated

174 MR. J. 8S. BALY ON SOME GENERA

posteriorly, convex, impressed below the base in the ¢ witha
large common fovea, the anterior margin of which is furnished
with two obtuse tubercles, placed one on each sutural margin
immediately below the scutellum; general surface of elytra
minutely and remotely punctured.

The 2 of this species is unknown to me.

8. PSEUDOCOPHORA BRUNNEA, Baly, Journ. Linn. Soe. vol. xx. p. 26.
Oblongo-ovata, postice paullo ampliata, convexa, fulva, nitida, pec-
tore, abdomine pedibusque posticis quatuor nigris ; elytrorum limbo
externo et (tn mare) fovea magna subbasali nigris ; thorace profunde
transversim sulcato; elytris substriato pone medium confuse punc-
tatis. Long. 3 ln.

Mas. Elytris infra basin fovea magna communi, elevato-marginata, antice
bituberculata instructis ; abdominis apice trilobato, lobo intermedio
plano.

Fem. Pygidii apice trifido, lobo intermedio elongato, tuberculo acuto
instructo ; segmento anali late emarginato.

Hab. Malacca; Celebes. .

Thorax nearly twice as broad as long; sides nearly straight and
very slightly diverging from the base to beyond the middle, thence
slightly converging to the apex ; disk deeply sulcate transversely,
the sulcation rather less deeply impressed on the median line.
Elytra sculptured as in P. Wallacei, differmg in the number
of teeth on the subbasal fovea; in the present species, as in the
preceding two, the anterior margin of the fovea is armed with
two obtuse tubercles, the hinder pair being obsolete.

The male of the present species differs from the same sex of
P. uniplagiata, Jac., in the form and sculpturing of the anal
segment of the abdomen; in P. brunnea the median lobe is
subquadrate and plane, in P. wniplagiata it is oblong and deeply
concave. The coloration of the insect closely resembles that of the
typical form of P. ambusta, Erichs.

Females of which the Males are unknown to me.

9. PSEUDOCOPHORA PECTORALIS. Ovata, postice ampliata, convexa,
flava, nitida, pectore abdomineque (hujus apice excepto) nigris;
thorace transverso, disco transversim sulcato; elytris convexis, infra
basin von excavatis, distincte punctatis, punctis hic illic subseriatim
dispositis. Long. 2% lin.

Fem, Abdominis segmento anali apice profunde emarginato.

a ee ee SS eee errr

eS ee ee eh Pe oa a a.

AND SPECIES OF GALERUCIN2. 175

Hab. Assam: a single specimen.

Antenne filiform ; eyes black. Thorax nearly twice as broad
as long; sides nearly straight, diverging from the base to
beyond the middle, thence obliquely converging towards the
apex; disk smooth, nearly impunctate, impressed just behind
the middle with a transverse groove. LElytra rather strongly
punctured.

10. PSEUDOCOPHORA PERPLEXA. Ovata, postice paullo ampliata,
convexa, piceo-fulva, nitida, metapectore, abdomine pedibusque
posticis quatuor nigris, antennis, tibiis tarsisque anticis pallide flavis ;
thorace sat profunde transversim sulcato; elytris tenuiter punctatis.
Long. 37 lin.

Fem. Abdominis pygidio apice leviter angulato-emarginato ; segmento
anali apice obtuso, obsolete sinuato.

Hab. Philippine Islands.

Antenne yellowish white. Thorax nearly twice as broad as
long; sides slightly diverging and sinuate from the base to
beyond the middle, thence converging towards the apex, the
hinder angle acute, the anterior slightly produced, obtuse ; upper
surface transversely convex, deeply sulcate transversely im-
mediately behind the middle, the sulcation nearly as deeply
impressed in its median portion as on its sides; disk smooth,
impressed with a few distant punctures on the sides. Elytra
oblong, slightly dilated posteriorly, convex, finely but distinctly
punctured.

The present insect closely resembles the same sex of P. ambusta,
var. A; but the entirely different form of the pygidium in the
two species at once separates them.

Genus AuLacopHora, Chevr.

The first six species here described have simple antenne in
both sexes; in the last eight the antenne in the males have the
third to the sixth joints compressed, and more or less thickened
and dilated ; the females of this section appear to be much rarer in
collections than the other sex, and only in a few cases are known
tome. ‘The following Table gives the differential characters of
the males of this latter group :—

176 MR. J. S. BALY ON SOME GENERA

TABLE.

I. Vertex and front on each side with an elevated ridge.

1. Frontal ridges transverse-............. 8. palliata.
2. Frontal ridges oblique.
a. Hiytra black |... se osc 7. orientalis.

aa. Klytra flavous, with black markings.. 11. bipartita.

3. Frontal ridges perpendicular.
b. Bbytra black’... eee ee 9. frontalis.
bb. Elytra flavous, with black markings... 10. fraudulenta.

II. Vertex and front without elevated ridges.
1. Elytra flavous, with black markings.
a. Antenne with their outer half

black... 6W0025 9 Ose 12. Olivierc.
aa, Antenne flayous) .....:. 5) snes 13. occipitalis.
2s, Blytra black i. .0.6% Ss... 236 eee 14. levifrons.

1. AULACOPHORA ANALIS, Weber.

Galeruca analis, Weber, Obs. Ent. p. 55, 1810; Fabr. Syst. El. i.
p. 482.

Subelongata, postice paullo ampliata, flava, nitida, thorace transverso,
vix pone medium transversim suleato; elytris fra basin obsolete
depressis, tenuissime punctatis, opacis, utrisque plaga magna humerali,
ad basin et ad marginem exteriorem adfixa, alterague pone medium,
subrotundata, seepe ad marginem extensa, nigris aut ceruleo-nigris,
Long. 33-4 lin.

A. ‘ Abdominis ano et margine, tibiis tarsisque nigrescentibus.”

B. Pygidio, abdomine, tibiis tarsisque nigris.

C. Abdomine pygidioque flavis, hoc szpe nigro-piceo tincto, tibiis
tarsisque ut in B.

Mas. Abdominis segmento anali trilobato, lobo intermedio quadrato-
oblongo, disco leviter concavo, medio elevato-vittato.

Fem. Abdominis segmento anali apice obtuso, medio obsolete emar-
ginato.

Hab. Sumatra, Celebes, Sulu Islands, Philippines.

Antenne slender, filiform in both sexes; the third joint rather
longer than any of the following ones. Thorax rather more than
one half broader than long; sides nearly straight and parallel,
sometimes slightly diverging from the base to the middle, very
slightly converging towards the apex anteriorly ; disk smooth,
impuuctate, impressed immediately behind the middle with a

‘
’

AND SPECIES OF GALERUCINE. 177

deep transverse groove. Elytra narrowly oblong, slightly dilated
posteriorly ; convex, very faintly excavated on the suture below
the basilar space, opaque, very minutely punctured.

A. analis, Weber, agrees very closely in general form with
A. bicolor of the same author; the structural differences, how-
ever, in the anal segments of the abdomen in both sexes, although
slight, are apparently constant, and sufficient to separate the two
insects ; 4. analis is also smaller than the other species.

2. AULACOPHORA WILsont. Anguste oblonga, postice ampliata,
convexa, nitida, subtus nigra, prothorace pedibusque anticis flavis, his
plus minusve piceo tinctis; supra flava, antennis (basi exceptis) scu-
telloque nigris ; thorace transversim sulcato, suleo fere recto, medio
magis fortiter excavato; elytris tenuiter punctatis. Long. 35 lin.

Mas. Abdominis segmento anali trilobato, lobo intermedio profunde
concavo; pygidii apice obtuso.

Fem. Abdominis segmento anali apice concavo-emarginato, incissure
margine leviter deflexo.

Hab. Australia, Melbourne.

Antenne moderately robust, filiform, the first joint, together
with the basal portions of the second and third, fulvous ; labrum
nigro-piceous ; eyes black. Thorax with its sides nearly straig)t
aud only very slightly diverging from the base to beyond the
middle, thence obliquely converging towards the apex; disk
transversely sulcate, the sulcation straight, rather more deeply
excavated in the middle. LElytra oblong, dilated posteriorly,
convex, obsoletely depressed below the basilar space, the latter
sometimes obsoletely thickened; the surface minutely punc-
tured.

In addition to other structural differences, the sculpturing of
the thorax will distinguish the above insect from A. nigroscutata,
which species it closely resembles in coloration.

3. AULACOPHORA PErRRouDI. Elongato-oblonga, postice paullo am-
pliata, convexa, flava, nitida, antennis (basi exceptis), pectore abdomi-
neque nigris, tibiis tarsisque nigro-piceis ; thorace transversim sulcato,
suleco medio fere obsoleto; elytris infra ‘basin leviter excavatis,
tenuiter sed distincte punctatis. Long. 4 lin.

Fem. Abdominis segmento anali rotundato, utrinque sinuato.

Hab. New Caledonia.
Eyes and antenne black, the latter with the two lower joints

178 MR.J. 8S. BALY ON SOME GENERA

flavous. Thorax twice as broad as long; sides diverging and

slightly sinuate from the base to beyond the middle, thence ~

rounded and converging to the apex; lateral margin strongly
reflexed; disk transversely sulcate, the sulcation strongly
excavated on each side, nearly obsolete on the middle disk.
Elytra narrowly oblong, slightly dilated posteriorly, slightly
but distinctly excavated below the basilar space, finely punc-
tured.

I only know a single specimen (a 9); it is at once to be
known from its allies by the interrupted suleation on the thorax.
This species is the one to which, in a note to the second Table in
my former paper on Awlacophora, I have alluded to as palustris,
Perroud ; that species belongs (as I have subsequently ascertained)
to the genus Monolepta.

4. AULACOPHORA ARGYROGASTER, Montrouz. Ann. Soc. Ent. France,
1861, p. 299.—Anguste oblonga, postice paullo ampliata, convexa,
sordide flava, subnitida; ore, antennis (basi exceptis) corporeque infe-
riore (pedibus anticis anoque exceptis) nigris aut nigro-piceis, abdo-
mine sparse argenteo-sericeo; thorace sat profunde transversim
suleato; elytris subopacis, tenuiter punctatis. Long. 33 lin.

Mas. Abdominis segmento anali trilobato, lobo intermedio profunde
concavo ; pygidii apice extremo obtuso.

Fem. Abdominis segmento anali late concavo-emarginato ; pygidii apice
acuto.

Hab. New Caledonia.

Antenne filiform, basal joint in the ¢ slightly thickened and
compressed, clavate ; the third, fourth, and fifth nearly equal in
length in both sexes. Thorax nearly twice as broad as long;
sides nearly parallel and slightly sinuate from the base to beyond
the middle, thence rounded and converging towards the apex ;
upper surface very minutely punctured, the puncturing coarser
on the sides; disk impressed across the middle with a deep
transverse sulcation, which in the ¢ is broader and more deeply
excavated in the middle, the anterior margin of the dilated por-
tion being thickened and obsoletely bituberculate; in the ?
the anterior margin of the sulcation (which is of nearly equal
depth and breadth throughout) is not thickened nor tuberculate.
Elytra oblong, not distinctly excavated below the basilar space,
minutely punctured.

The peculiar sculpture of the thorax in the ¢ and the acute

AND SPECIES OF GALERUCIN 2. 179

apex of the pygidium and sculpture of the anal segment of the
abdomen in the 2, conjointly with the coloration of the under
surface, will separate this species from its congeners.

5. AULACOPHORA AUSTRO-CALEDONICA, Montrouz. Ann. Soc. Ent.
France, 1861, p. 299.—Subelongata, postice paullo ampliata, con-
vexa, flava, nitida, verticis maculis duabus, pectore abdomineque
nigris, hoc basi et ad latera pallide piceo; thorace transversim sul-
cato, sulco fere recto; elytris nitidis, utrisque plagis magnis duabus,
una basali ad suturam abbreviata, altera pone medium _positis,
nigris. Long. 24-3 lin.

Mas. Antennis filiformibus, robustis; abdominis segmento anali tri-
lobato, lobo intermedio oblongo, profunde concavo.

Fem. Abdominis segmento anali lateribus rectis, oblique convergenti-
bus, apice extremo obtuso.

Hab. New Caledonia, Balade.

Vertex with two large black patches, separated in the middle
by a narrow flavous line; antenne filiform in both sexes, more
robust in the ¢ thanin the other sex, the joints slightly stained
with piceous. Thorax more than one half broader than long ; its
sides sinuate and slightly diverging from the base to the middle,
slightly produced immediately before the latter, thence rounded
and converging to the apex; disk impressed across the middle
with a deep transverse groove. LElytra oblong, very slightly
dilated posteriorly, minutely punctured.

This species closely resembles A. occipitalis in the coloration
of its head and underside; the antenne,’ however, of the ¢ of
austro-caledonica are simple, not dilated as described by Mr.
Jacoby (Annal. del Mus. Civic. Stor. Nat. Gen. 1886, vol. iv.
p- 53); the male assigned by him to the present insect belongs
to another species nearly allied to, if not identical with,
A. occipitalis. I possess both sexes of austro-caledonica from
New Caledonia.

6. Autacopuora Lewisi1, Baly, Journ. Linn. Soc. vol. xx. p. 24.—
Mas. Antennis robustis, ad apicem paullo attenuatis, abdomine nigro
avt piceo-nigro, apice extremo piceo-fulvo ; segmento anali trilobato,
lobo intermedio longitudinaliter sulcato.

Fem. Abdominis segmento anali apice utrinque leviter sinuato.

Var. A. Corpore subtus toto glabro.

Hab. India and China (type and var. A}; Malay Archipelago
(var. A).

180 MR. J. 8S. BALY ON SOME GENERA

The specimen from which I originally drew up the diagnosis
of this species (a 2) had been carded, and the apex of the
abdomen was so obscured by gum that I described it as
entire ; since then I have had the opportunity of examining many
specimens of both sexes, and find that the apex of the female
anal segment is distinctly, although slightly, sinuate on each
side.

7. AULACOPHORA ORIENTALIS, Hornst. Schrift. Berl. Ges. viii.
AVB8,4p.oy ts Lone, 3.—Anguste oblonga, postice paullo ampliata,
convexa, rufo-fulva, nitida, antennis flavis, oculis elytrisque nigris ;
thorace transverso, disco transversim suleato, elytris convexis, infra
basin leviter transversim depressis, tenuiter punctatis. Long.
3 lin.

Mas. Capitis vertice utrinque creta elevata, oblique posita, instructo ;
antennarum articulis tertio, quarto quintoque incrassatis, difformi-
bus ; abdominis segmento anali lobo intermedio oblongo-quadrato,
plano aut leviter concavo.

Var. A, 3. Antennis, tibiis tarsisque nigro-piceis.

Hab. Java, New Guinea, Dorey, Ceram, Gilolo; var. A, New

Guinea ( Wallace).

Vertex in the ¢ with an elevated ridge, placed obliquely on
each side just above the eye and running inwards to nearly
meet its fellow in the median line; antennz in the g with the
basal joint thickened, slightly curved, clavate, its outer edge sub-
carinate; second joint small, the third thickened and subclavate,
its apex truncate; the fourth and fifth also thickened, dilated
laterally, subtrigonate, each shorter than the third; the apex of
the fourth obliquely truncate, its anterior and outer angle
produced, acute or subacute; the fifth with its outer sur-
face deeply concave, the anterior angles of the concavity
acute. ‘Thorax nearly twice as broad as long; sides nearly
straight and parallel from the base to beyond the middle;
upper surface transversely sulcate immediately behind the
middle, rather coarsely punctured on the sides in front. Elytra
slightly convex, faintly depressed below the basilar space, finely
punctured.

8. AULACOPHORA PALLIATA, Schaller, Abhandl. Hall. Ges. 1. 1783,
p- 279; Fabr. Mant. 1. 1787, p. 87; Oliv. Ent. vi. p. 625, £. 2.
fig. 25, a-b.—Anguste oblonga, postice ampliata, convexa, fulvo-rufa,

AND SPECIES OF GALERUCINA. 181

nitida, elytris nigris ; thorace transversim sulcato, ad latera punctato.
Long. 3} lin.

Mas. Capitis vertice utrinque creta crassa transversa instructo; anten-
narum articulis tertio, quarto quintoque ampliatis, difformibus ; abdo-
minis segmento anali lobo intermedio quadrato-oblongo, leviter
concavo.

Hab. India; Malay Archipelago.

Front on each side in the ¢ with a strongly thickened trans-
verse ridge; antenne in the same sex with the basal joint
thickened, slightly curved, the second very small, the third and
two following ones thickened and laterally dilated, the third
slightly longer than broad, trigonate, its outer and upper angle
slightly produced, acute, the fourth very slightly shorter than
the third, quadrangular, its upper and outer angle broadly and
obliquely truncate; the fifth shorter than either of the preceding
two, subtrigonate, its outer surface longitudinally concave, the
outer and upper angle of the concavity bidentate. Thorax nearly
twice as broad as long; sides nearly straight and parallel from
the base to beyond the middle, thence obliquely converging
towards the apex; disk transversely sulcate just below the
middle, coarsely punctured on the sidesin front. Elytra oblong,
slightly dilated posteriorly ; convex, slightly impressed below
the basilar space, finely punctured.

9. AULACOPHORA FRONTALIS. Anguste oblonga, postice paullo am-
pliata, flavo-fulva, nitida, antennis pallide flavis, elytris nigris; tho-
race transversim sulcato, remote punctato, punctis ad latera subcrebre
dispositis ; elytris infra basin transversim excavatis, tenuiter punc-
tatis. Long. 23 lin.

Mas. Capitis fronte bisuleato, utrinque inter sulcum et oculum tuber-
culo oblongo, dorso compresso et intus curvato, instructo; antenna-
rum articulis tertio, quarto quintoque ampliatis; abdominis seg-
mento anali trilobato, lobo intermedio quadrato-oblongo, leviter
concavo.

Hab. Borneo, Sarawak (Wallace).

Front and vertex longitudinally bisulcate; placed longitudi-
nally on each side between the sulcation and the eye is an
oblong protuberance, the upper edge of which is compressed and
incurved ; front separated from the encarpe by a deep transverse
groove ; antennein the ¢ with the basal joint thickened, slightly
curved, its lower surface deeply emarginate, the third and two

LINN. JOURN.—ZOOLOGY, VOL. Xx. 15

182 MR. J. 8S. BALY ON SOME GENERA

following joints thickened and dilated, trigonate, the third
nearly twice as broad as long, its outer edge strongly com-
pressed, carinate, the fourth and fifth nearly equal in length,
each rather shorter than the third, the upper and outer angle
of the fifth emarginate, bidentate. Thorax rather more than
one half broader than long; sides nearly straight from the base
to the middle, thence slightly rounded and converging towards
the apex; disk transversely sulcate immediately behind the
middle, very distantly and minutely punctured, the puncturing
rather closer on the sides. Elytra oblong, dilated posteriorly,
convex, transversely excavated below the basilar space, finely
punctured.

The three similarly coloured species described above are readily
separated in the male sex by the position and form of the frontal
tubercles. The females are not known to me.

10. AULACOPHORA FRAUDULENTA, Jacoby, Annal. del Museo Civ.
di Storia Natur. di Genova, vol. iv. p. 52 (1886).—Anguste oblonga,
postice ampliata, convexa, flava, nitida, antennis extrorsum infuscatis ;
oculis, metasterno elytrisque nigris, his fascia lata prope medium
flava. Long. 3 lin.

Mas. Capitis fronte, utrinque prope oculum tuberculo oblongo magno,
longitudinaliter posito, instructo ; antennarum articulis tertio, quarto
quintoque ampliatis, trigonatis; abdominis segmento anali trilobato,

lobo intermedio oblongo-quadrato, apice bilobato, disco leviter longi-
tudinaliter concavo.

Hab. New Guinea.

Front on each side in the ¢ with a strongly raised oblong
tubercle, placed longitudinally close to the inner margin of the
eye, apex of the tuberosity torulose, slightly incurved ; antenne
in the same sex with the basal joint thickened, its outer edge
concave-emarginate ; the third and two following joints dilated,
trigonate, the third and fifth equal in length and breadth, the
fourth longer than either of the two others, but less broadly
dilated; in the specimen before me, sent by Mr. Jacoby, the
outer half of the antenne is stained with fuscous (the author
describes these organs as entirely flavous)*. Thorax nearly

* The 9 of this insect is unknown to me, the specimen sent by Mr. Jacoby

as belonging to that sex proving to be the ¢ of a similarly coloured species
having simple antenne.

AND SPECIES OF GALERUCINA. 1838

twice as broad as long; sides nearly straight and slightly di-
verging from the base to beyond the middle, thence rounded and
converging towards the apex; disk impressed just behind the
middle with a transverse sulcation ; surface finely and remotely
punctured, the punctures more crowded on the sides in front.
Elytra convex, faintly excavated on the suture below the basilar
space, finely punctured.

1]. AULACOPHORA BIPARTITA. Anguste oblongo-ovata, postice
paullo ampliata, convexa, flava, subnitida, vertice, oculis, thoracis
dimidio antico, metapectore abdomineque nigris; thorace transversim
sulcato, suleo medio minus fortiter impresso, disco interdum ante
basin maculis parvis male definitis nigro-piceis, instructo; elytris
oblongis, postice paullo ampliatis, subcrebre punctatis, obsolete longi-
tudinaliter sulcatis; utrisque plagis duabus magnis, una basali, unaque
vix pone medium, nigris. Long. 3 lin.

Mas. Capitis vertice utrinque creta oblique posita instructo; an-
tennarum articulis tertio, quarto quintoque compressis, triangulariter
dilatatis; abdominis segmento anali trilobato, lobo intermedio con-
cavo.

Fem. Antennis simplicibus; abdominis segmento anali apice obtuso.

Var. A, Q. Capite toto flavo, thorace utrinque ante medium
plaga magna nigra instructo; elytrorum plaga infra medium ob-
soleta.

Hab. Flores; Sarawak (Wallace). Var. A, Java.

Clypeus in the g with a narrow, rather strongly raised longi-
tudinal ridge; front just above the encarpx with a deeply im-
pressed fovea; obliquely placed on each side just within the
eye is a broad slightly raised elongated ridge; eyes in the ¢
large, prominent. Antenne with the third and following two
joints compressed, triangularly dilated, the third less broadly so
than the two others; third and fourth equal in length, the
fifth rather shorter. Thorax transverse; sides slightly diverging
and sinuate from the base to beyond the middle, thence ob-
liquely rounded and converging to the apex; upper surface
minutely punctured, impressed immediately behind the middle
with a deep transverse groove. LElytra oblong, slightly dilated
towards the apex ; convex, distinctly and rather closely punc-
tured ;“disk of each elytron with five or six faint longitudinal
sulcations.

184 MR. J. S. BALY ON SOME GENERA

12. AuLacoPHora OLIVIERI.

Galeruca analis, Oliv. Ent. vi.

Oblongo-ovata, convexa, flava, nitida, labro, antennis (basi exceptis),
mesosterno, abdominis apice, tibiis tarsisque nigro-piceis aut nigris ;
thorace quam longo fere duplo latiore, transversim sulcato, sulco
medio minus fortiter impresso; elytris tenuiter punctatis, utrisque
plaga magna basali, ad suturam abbreviata, altera pone medium, ad
marginem plerumque adfixa, apiceque nigris. Long. 3—3% lin.

Mas. Antennarum articulis tertio, quarto quintoque incrassatis ; abdo-
minis segmento anali trilobato, lobo imtermedio longitudinaliter
suleato.

Fem. Antennis filiformibus; abdominis segmento anali apice ro-
tundato.

Hab. Australia, Hunter’s River, Rockhampton, Paroo River.

Front impressed immediately above the encarpe with a small
fovea; antenne robust, the third, fourth, and fifth joints in the
3 thickened, the third nearly twice as long as broad, obconic, the
fourth and fifth each shorter than the third, laterally dilated, tri-
gonate; antenne in the @ simple, filiform; the five lower
joints in the 3, the three or four basal ones in the Q, flavous or
fulvous, more or less stained with piceous, the remaining joints
in each sex nigro-piceous or black. Thorax nearly twice as
broad as long ; sides slightly diverging and slightly sinuate from
the base to just beyond the middle, thence rounded and con-
verging to the apex ; upper surface remotely and minutely punc-
tured, transversely sulcate immediately behind the middle, the
sulcation rather less deeply impressed on the middle disk.
Elytra oblong, slightly dilated posteriorly, convex, minutely
punctured, the basilar space obsoletely thickened. Tibize and
tarsi black, the basal portion of the former sometimes flavous.

This species usually stands in collections as A. analis, Weber ;
it is very constant in pattern and coloration, and differs from the
author’s description of analis in the dark antenne, in the black
apices of the elytra, and in the similarly coloured mesosternum.
lts locality is also entirely different, all the specimens that I have
seen having been brought from Australia; whereas Weber gives
Sumatra as the habitat of his insect.

13. AULACOPHORA OCCIPITALIS. Subelongata, postice paullo am-
pliata, convexa, flava, nitida, oculis, verticis plagis duabus, pectore
abdomineque nigris; thorace prope medium transversim sulcato ;

AND SPECIES OF GALERUCINA. 185

elytris infra basin leviter transversim excavatis, utrisque plaga basali
subquadrata, ad suturam et ad marginem abbreviata alteraque pone
medium subrotundata, nigris. Long. 3 lin.

Mas. Antennarum articulis tertio, quarto quintoque compressis et dila-
tatis; abdominis segmento anali trilobato, lobo mtermedio oblongo,
profunde concavo.

Hab. New Guinea, Dorey; Bouru (Wallace).

Vertex with two large black patches, divided in the median line
by a flavous line ; antenne with the third, fourth, and fifth joints
compressed and dilated in the 3; the third twice as long as
broad, narrowly cuneiform, the fourth shorter and more strongly
dilated than the third, trigonate, its antero-external angle pro-
duced, very acute; the fifth still shorter than the fourth, subtri-
gonate. Thorax about one half as broad again as long; sides
sinuate, subparallel, slightly produced just beyond the middle,
thence obliquely converging towards the apex; disk impressed
across the middle with adeep suleation. Hlytra faintly excavated
below the basilar space, nitidous.

The ¢ of A. occipitalis differs from the same sex of A. frau-
dulenta, Jac., in the absence of the frontal tubercles, and in
the different length and shape of the dilated joints of the
antenne, also in the black patches on the vertex, and in the
black breast and abdomen. The females of both species are
unknown to me.

14. AULACOPHORA L&VIFRONS. Anguste oblonga, postice ampliata,
rufo-fulva, nitida, antennis flavis, tibiis (basi exceptis) tarsisque piceis
aut nigro-piceis, elytris tenuiter punctatis, nigris; thorace vix pone
medium transversim sulcato, sulco profunde impresso, paullo infra
marginem lateralem abbreviato. Long. 3 hn.

Mas, Capitis vertice non tuberculato ; antennarum articulis tertio, quarto
quintoque incrassatis et ampliatis, difformibus.

Fem. Antennis filiformibus, abdominis segmento anali apice obtuse
truncato.

Var. A, 9. Tibiis tarsisque rufo-fulvis.

Hab. Java; Singapore (Wallace).

Vertex and front smooth, not tuberculate, the latter impressed
just above the encarp with a deep fovea; antennw in the ¢ with
the basal joint slightly thickened, subclavate, the third joint slightly
thickened, obconic ; the fourth equal in length to the preceding
one, rather strongly thickened, slightly compressed, noduliform,

186 MR. J. S. BALY ON SOME GENERA

the fifth shorter than either of the preceding two, broader than °
long, trigonate, its antero-external angle slightly produced, —
acute. Thorax more than one half as broad again as long; sides
nearly straight and diverging from the base to beyond the
middle, thence obliquely converging towards the apex; disk
deeply transversely sulcate immediately behind the middle, the
sulcation more deeply impressed on each side, abbreviated just
within the lateral margin; surface very finely and remotely
punctured, the punctures more crowded on the sides in front.
Elytra oblong, dilated posteriorly ; convex, transversely excavated
below the basilar space, finely punctured.

The abdomen of the ¢ in the only specimen known to me is
not fully developed; I am therefore unable to give the form of
the median lobe of the anal segment of the abdomen.

Genus LEPTOXENA.

_ Corpus angustum, elongatum, convexum. Caput exsertum ; antennis
filiformibus, extrorsum leviter incrassatis et paullo compressis ;
oculis integris; palpis maczillaribus ovatis, articulo ultimo acuto.
Thorax transversus, transversim convexus. lytra subcylindrica;
epipleuris a basi ad angulum posticum extensis. Pedes robusti;
tibiis inermibus ; wnguiculis bifidis, dente interno breviore ; femoribus
tibiisque posticis in ¢ incrassatis; tarsis posticis articulo basali ad
duos sequentes fere zquilongo. Prosternum basi non lobatum ;
acetabulis anticis fere clausis, epimeris anticis cum antepectoris mar-
gine postico intime connexis.

Leptoxena is allied to Galerucella, but may be separated from
that genus by its narrower form, thickened hinder legs in
the ¢, and by the anterior epimera being continuous with the
hinder border of the antepectus for their whole length, whereas
in Galerucella their apices are free.

LEPTOXENA EXIMEA. Anguste elongata, parallela, transversim con-
vexa, fusco-fulva, subnitida, thoracis punctis duobus, antennis (basi
exceptis) abdomineque nigris ; thorace transverso, lateribus late rotun-
datis, antice sinuatis; disco nitido, leviter transversim convexo,
utrinque leviter excavato ; elytris fere parallelis, subcylindricis, infra
basin non excavatis, fortiter et irregulariter punctatis, vittis non-
nullis, leviter elevatis instructis, interstitiis reticulatis. Long.
5 lin.

AND SPECIES OF GALERUCINA. 187

Mas. Abdominis segmento anali apice angulatim inciso; femoribus

tibiisque posticis incrassatis, his extrorsum ante apicem sinuatis.

Fem. Abdominis segmento anali integro; pedibus posticis non in-

crassatis.

Hab. Andaman Islands (Calcutta Museum and my own
collection).

Head strongly exserted, vertex swollen ; antenne with the first
joint curved, clavate, the second and third short, nearly equal,
the fourth as long as the preceding two united, these three
joints cylindrical, the fifth and following ones compressed, the five
upper ones being thickened and moderately dilated; the six
upper joints black. Thorax nearly three times as broad as
long; sides broadly rounded, sinuate before the middle, the
hinder angles obsolete; disk transversely convex, slightly but
broadly depressed and excavated on each side; nitidous,
the central third, the median line excepted, rather closely
punctured, sides iapunctate; on each side, but at some dis-
tance from the outer margin, is a round: black or nigro-
piceous spot. Scutellum clothed with sericeous hairs. Elytra
rather broader than the thorax, parallel on the sides; upper
surface transversely convex, not depressed or excavated below
the basilar space ; closely and strongly punctured, the interspaces
subelevate-reticulate ; each elytron with seven or eight slightly
raised longitudinal cost, those on the outer disk more distant
and less defined.

I possess both sexes of this species from the Andaman
Islands; I have also received it, without locality, from the
Calcutta Museum.

List of Species.
Charidea (n. g.) punctato-striata (Motsch.) Japan.

MILAN ate iV ccehpussieviae'ds ceseoseees Northern China.
Platywantha clypeata ......escrcosccecssevees Andaman Islands.
CEnidea ceruleipennis .......2...- Peet ues Siam.

SMEs da t00's's cv ovens svscednoesecsss Borneo.

Oe Sumatra.

PPPOD CORNED F oiisc ces. cucucescscoccescsvene Malacca.

MME ios a ays sine ovary cevicenvenecave Sarawak,

I Sica Sosiv asm cvedayacdvavees uf:

Monolepta kemorrhoidalis (Fabr.) ......... Australia; New Guinea.

ERT cotiew ys cased ears casks ve vnessess Manilla.

188

ON SOME GENERA AND SPECIES OF GALERUCINE.

Monolepta foveicollis .........cecseececsceees Batchian.
terminata; Guébe.1~ tek. 8t.k ee ae Java.
a Wallaget sci wcsetns Swed Poe ch aebete Celebes.
VETTICALES 0x 3 hinSastca iach encene eee eee Batchian.
Paraulaca (n. g.) angulicollis (Motsch.).
Pseudocophora Buquetit (Guér.) ......... Java.
GESCUNICUG.. «dan 5 upewgnet Se tee ee ae Borneo.
Les FUMCMBONE SA Linn te baeelN beer ce da tweak Sumatra.
ambusta (Hirichs.)* &. ....c0....ceativet Luzon.
Ss MOMMA. 556 vi gnie.e capes mee Ree! Bouru.
uniplagiata, Jacoby .........0e00. 60 Sumatra.
JLODEONG i cctee castes OoteeN cetera Andaman Islands.
SS GUILT Sohseway os ners owes Seton eaneee Malacca.
DEOUOT QUES. \aninianais sive ax omu tee he oe Assam.
WET DUOMO LS a ostsdas SRE dene ee Manilla.
Aulacophora analis (Weber)..........0.+4+ Sumatra.
WESONG «cies crieeo se neghe Vee eE Ce TEER Melbourne.
SS PEPTOUME » 2 vona cues npecota ner see N. Caledonia.
argyrogaster (Montrouzier)......... 4)
austro-caledonica (Montrouzier) ... 7
Yh) Se ee eae mee AS China, &e.
orientalis (Hornst. ii .5.i:..a0. teak Java.
pabliiata (Schaller). sa... <sieeevne sh India.
NDOT REDS » lc Cui wscionsuh ini vecs caet Borneo.
Sraudulenia, Jacoby......... +++. N. Guinea.
ULBOIIOL a nsnbitaes <tinin Pocus rains site Flores.
OUI x akin oct canigek womans aceon coated Australia.
QUCIDURM IR cds 25h 0, Med. donk. s eae N. Guinea.
me WOUIIT ONS, loves oictvs viecsSaneosse + vcvlepire Java.

LCPOLONAA(D... F.)'CLUNED nun oe ccedencnes Andaman Islands.

EVOLUTION THROUGH CUMULATIVE SEGREGATION. 189

Divergent Evolution through Cumulative Segregation. By Rev.
Joun Toomas Guticx. (Communicated by Atrrep Russe
Wattacz, F.L.S.)

[Read 15th December, 1887. |
INTRODUCTION.

In my study of Sandwich-Island terrestrial mollusks my atten-
tion was early arrested by the fact that wide diversity of allied
species occurs within the limits of a single island, and in
districts which present essentially the same environment. As
my observations extended, I became more and more impressed
with the improbability that these divergences had been caused
by differences in the environment. It was not easy to prove that
sexual selection had no influence; but, owing to the very low
grade of intelligence possessed by the creatures, it seemed im-
possible that the form and colouring of the shells should be the
result of any such process. I was therefore led to search for
some other cause of divergent transformation, the diversity of
whose action is not dependent on differences in nature external
to the organism.

I found strong proof that there must be some such principle, not
only in the many examples of divergence under uniform activities
in the environment, but in the fact that the degrees of diver-
gence between nearly allied forms are roughly measured by the
number of miles by which they are separated, and in the fact
that this correspondence between the ratios of distance and
the ratios of divergence is not perceptibly disturbed by passing
over the crest of the island into a region where the rainfall is
much heavier, and still further in the fact that the average
size of the areas occupied by the species of any group varies,
as we pass from group to group, according as the habits of the
group are more or less favourable to migration. I perceived that
these facts could all be harmonized by assuming that there is some
cause of divergence more constant and potent than differences in
nature external to the organism; and that the influence of this
cause was roughly measured by the time and degree of separation.

During the summer of 1872 I prepared two papers in which
these facts and opinions were presented. One of these, entitled
“The Variation of Species as related to their Geographical Dis-
tribution, illustrated by the Achatinelline,”’ was published in

LINN. JOURN.— ZOOLOGY, VOL. XxX. 16

190 _ REV. J. T. GULICK ON DIVERGENT EVOLUTION

‘Nature’ for July 18, 1872; the other, entitled “ Diversity of
Evolution under one Set of External Conditions,” after being read
before the British Association for the Advancement of Science in
August 1872, was, through the kindness of Mr. Alfred Wallace,
brought before the Linnean Society, and was finally published in
the Linnean Society’s Journal, Zoology, vol. xi. pp. 496-505.

In the former paper I used the following words in calling
attention to the impossibility of explaining the origin and distri-
bution of these forms by Natural Selection. ‘“ Whether we call
the different forms species or varieties, the same questions are
suggested as to how they have arisen and as to how they have
been distributed in their several localities. In answering these
questions, we find it difficult to point to any of those active
causes of accumulated variation, classed by Darwin as Natural
Selection. .... There is no reason to doubt that some varieties
less fitted to survive have disappeared; but it does not follow
that the ‘Survival of the Fittest’ (those best fitted when com-
pared with those dying prematurely, but equally fitted when
compared with each other) is the determining cause which has
led to these three species being separated from each other in
adjoining valleys. Zhe ‘Survival of the FHittest’ still leaves a
problem concerning the distribution of those equally fitted. It
cannot be shown that the ‘ Survival of the Fittest’ is at variance
with the survival, under one set of external circumstances,
of varieties differing more and more widely from each other
in each successive generation. The case of the species under
consideration does not seem to be one in which difference of
environment has been the occasion of different forms being
preserved in the different localities. It is rather one in which
varieties resulting from some other cause, though equally fitted
to survive in each of the localities, have been distributed accord-
ing to their affinities in separate localities.”

In the latter paper I raised the following questions con-
cerning Natural Selection. ‘The terms ‘ Natural Selection ’ and
‘Survival of the Fittest’ ....imply that there are variations
that may be accumulated according to the differing demands of
external conditions. What, then, is the effect of these variations
when the external conditions remain the same? Or, can it be
shown that there is no change in organisms that is not the result
of change in external conditions? Again, if the initiation of
change in the organism is through change in the environment, ...

THROUGH CUMULATIVE SEGREGATION. 191

does the change expend itself in producing from each species just
one new species completely fitted to the conditions, or may tt pro-
duce from one stock many that are equally fitted?” (p. 497). In
answering these questions I called “attention to the variation
and distribution of terrestrial mollusks, more especially those
found on the Sandwich Islands,” and gave what seemed to me
strong reasons for believing that “The evolution of these dif-
ferent forms cannot be attributed to difference in their external
conditions. ....I£ we would account for the difference and the
limited distribution of these allied forms on the hypothesis of
evolution from one original species, zt seems to me necessary to
suppose two conditions, Separation and Variation. I regard
Separation as a condition of the species, and not of surrounding
nature, because it is a state of division in the stock which does
not necessarily imply any external barriers, or even the occupation
of separate districts. This may be illustrated by the separation
between the castes of India, or between different genera occupying
the same locality. ... We must suppose that they [the diverging
forms| must possess an inherent tendency to variation so strong
that all that is necessary to secure a divergence of types in the
descendants of one stock is to prevent, through a series of gener-
ations, their intermingling with each other to any great degree”
(pp. 498-499). Lalso called attention to the fact that some forms
of Natural Selection must “ prevent variation and give a wider
diffusion to forms that would otherwise be limited in their range
and variable in their type. Natural Selection is as efficient in
producing permanence of type in some cases as in accelerating
variation in other cases’ (p. 504). On page 499 I pointed out
the law that “ The area occupied by any species must vary directly
as its power and opportunity for migration, and inversely as
its power of [divergent] variation.”” And on page 505 I gave a
brief summary of my reasons for believing that “Separation
without a difference of external circumstances is a condition suffi-
cient to ensure .. . divergence in type.”

Subsequent investigation has led to the development of my
theory, with a fuller discussion of the causes and laws that are
revealed in these phenomena. In an article published in ‘ The
Chrysanthemum’ (Yokohama and London, Triibner & Co.),
January 1883, I state my belief “ that the quality, the diversity,
and the rapidity of the variation depend chiefly upon the nature
of the organism; and that while the nature of the external

16*

192 REV. J. T. GULICK ON DIVERGENT EVOLUTION

conditions has power to winnow out whatever forms are least
fitted to survive, there will usually remain a number of varieties
equally fitted to survive; and that through the law of segregation
constantly operating in species distributed over considerable
areas, these varieties continue to diverge both in form and in
habits till separate species are fully established, though the con-
ditions are the same throughout the whole area occupied by the
diverging forms.”’ The conclusion reached was, that “ The theory
that diversity of Natural Selection is, like variation, an essential
factor in producing diversity of species, is untenable. On the
contrary, we find that diversity of Natural Selection is not
necessary to diversity of evolution, nor uniformity of Natural
Selection to uniformity of evolution; but while variation and
separation are the essential factors in diversity, and intercrossing
and unity of descent the essential agents in uniformity of evolu-
tion, Natural Selection may be an important ally on either side.”

In an article on “ Evolution in the Organic World,” published
in ‘The Chinese Recorder’ (Shanghai), July 1885, I use the fol-
lowing language :—“ We see what Natural Selection cannot
explain by considering the nature of the process. The sur-
vival of the fittest results in the separate breeding of the fittest,
and therefore in the increasing fitness of successive generations
of survivors ; but how can it account for the division of the survi-
vors of one stock, occupying one country, into forms differing more
and more widely from each other? To explain such a result we
must find some other law. Iam prepared to show that there is such
a law rising out of the very nature of organic activities, a law of
Segregation, bringing together those similarly endowed, and sepa-
rating them from those differently endowed.”

Without Variation there can be no Segregate Breeding; and
without Segregate Breeding and Heredity there can be no accu-
mulation of divergent variations resulting in the formation of
races and species. In producing divergent evolution, the causes
of Variation and Heredity are therefore as important as the
causes of Segregate Breeding ; and though I pass them by in my
present discussion, I trust it will not be attributed to an under-
estimate of their importance. Though I do not stop to discuss
the causes of variation, my reasoning rests on the observed
fact that in every department of the organic world variation is
found, and that in the vast majority of cases, if not absolutely in
all, the diversities to which any freely intergenerating group of

OP Ha Ka. s.

THROUGH CUMULATIVE SEGREGATION. 1938

organisms is subject follow the general law of “ Frequency of
Deviation from an Average.” As this isa law according to which
half of the members of the intergenerating group are above and
half below the average in relation to any character, there must
often occur simultaneous variation of several individuals in some >
character which tends to produce Segregate Breeding. The
reality and importance of this law is not at all dependent on the
reality of any of the theories of heredity and variation that are
now being discussed. Whatever may be the causes that produce
variation, whether they depend entirely upon changes in external
conditions, or are chiefly due to changing activities in the
organism and the hereditary effects of acquired characters, or are,
as Weismann maintains, the direct result of sexual reproduction
which never transmits acquired characters,—in any and every
case this law of Deviation from an Average remains undisturbed,
and is recognized as an important factor in the present paper.
It therefore cannot be urged that the theory here advanced
assumes simultaneous variation without any ground for making
such an assumption ; nor can it be said that it rests on the in-
credible assumption that chance variation of very rare kinds will
be duplicated at one time and place, and will represent both
Sexes.

Moritz Wagner first discussed what he calls “ The law of the
migration of organisms” in a paper read before the Royal
Academy of Sciences at Munich, in March 1868; but my attention
was not called to it till after the reading of my paper before the
British Association in August 1872. In a fuller paper entitled
“The Darwinian Theory and the Law of the Migration of
Organisms,” an English translation of which was published by
Edward Stanford (London, 1873), the same author maintains that
“the constant tendency of individuals to wander from the
station of their species is absolutely necessary for the formation
of races and species”’ (p. 4). ‘‘The migration of organisms and
their colonization are, according to my conviction, a necessary
condition of natural selection” (p. 5). On pp. 66 and 67 he
expands the same statement, and objects to Darwin’s view “ that
on many large tracts all individuals of the same species have
become gradually changed.” Again, he contends that “ Trans-
formation is everywhere and always dependent on isolation in
order to have lasting effect. Without separation from the home
of the species, this wonderful capacity would be completely

194: REV. J. T. GULICK ON DIVERGENT EVOLUTION

neutralized” (p. 74). “ Natural Selection is not in itself an
unconditional necessity, but is dependent on migration and
geographical isolation during a long period, together with altered
conditions of life” (p. 57). ‘ Where there is no migration, that
is where no isolated colony is founded, natural selection cannot
take place ’’ (p. 59).

A comparison of his paper with my two papers published in
1872, already referred to, will show several fundamental dif-
ferences in the two theories. He maintains that :—

(1) The separation of a few individuals from the rest of the
species is absolutely necessary for the operation of Natural
Selection, and therefore for any transformation of the species,
no matter how great the change of conditions may be in the
original home of the species.

(2) Migration and geographical barriers are the only effectual
causes, independent of human action, by which a few individuals
can be separated from the rest of the species, and are, therefore,
necessary to the transformation of species.

(3) Exposure to a new form of Natural Selection is a necessary
condition for any transformation of a species.

(4) Difference of external conditions is necessary to difference
of Natural Selection, and therefore necessary to any transforma-
tion of species.

(5) Geographical isolation and altered conditions of life are
necessary conditions for Natural Selection, as that is for the
modification of species.

(6) The separation of which he speaks is the entering of a few
individuals into a new territory, where the conditions are dif-
ferent from those in the old habitat, and where the body of the
species fail of reaching them.

My chief positions were the following, in strong contrast with
the foregoing :—

(1) Separate generation is a necessary condition for divergent
evolution; but not for the transformation of all the survivors of
a species in one way.

(2) “Separation does not necessarily imply any external
barriers, or even the occupation of separate districts.”

(3) Diversity of Natural Selection is not necessary to diversity
of evolution.

(4) Difference of external conditions is not necessary to di-
versity of evolution. |

THROUGH CUMULATIVE SEGREGATION, 195

(5) “Separation and Variation,” that is, Variation not over-
whelmed by crossing, “is all that is necessary to secure a di-
vergence of types in the descendants of one stock,’ though
external conditions remain the same, and though the Separation
is other than geographical.

(6) The Separation of which I speak is anything, in the
species or in the environment, that divides the species into two
or more sections that do not freely intercross, whether the dif-
ferent sections remain in the original home or enter new and
dissimilar environments.

Though these propositions were very briefly and imperfectly
presented, I am not aware that any better statement of the facts
of Segregation had been previously published.

The present paper is the result of a long continued endeavour
to understand the relations in which this factor stands to Natural
Selection and the other causes that co-operate in producing
divergent evolution; and though my work has been done under
the great disadvantage of entire separation from libraries, and
from other workers in similar lines, I trust it may contribute
something towards the elucidation of the subject. In expanding
my theory I have been unable to make any use of the positions
taken in Moritz Wagner’s paper, as they seem to me very
extreme and far removed from the facts of nature. The two
theories correspond chiefly in that they discuss the relation of
Separation to the transformation of species; while the explana-
tions given of the nature, causes, and effects of Separation widely
differ. I am informed that my paper on “ Diversity of Evolution
under One Set of External Conditions ” was translated and circu-
lated in Germany ; but whether it had any effect in modifying
Wagener’s theory, I have not the means of knowing.

I have recently discovered that the principle of’ Segregate
Breeding, which I have found to be of such importance in the
evolution of species, is allied to the law of Segregation pro-
pounded by Spencer in his ‘ First Principles.’ By direct con-
sideration of the conditions that have been found necessary for
the development of divergent races of domestic plants and animals,
T have discovered Segregate Breeding as a necessary condition for
divergent evolution ; and by direct observation on the propaga-
tion of plants and animals under natural conditions, I have
discovered Cumulative Segregation as a constant result fron-
certain forms of activity in the organism when dealing with a

196 REV. J. T. GULICK ON DIVERGENT EVOLUTION

complex environment; it is therefore with special pleasure that
I observe that a law of very similar import may be derived by a
wholly different method from the general laws of action and
reaction in the physical world. It should, however, be noticed
that in the brief references made to the subject in Spencer’s
‘Principles of Biology’* it is assumed that “ Increasingly-
definite distinctions among variations are produced wherever
there occur definitely-distinguished sets of conditions to which
the varieties are respectively subject,’ and only where these
occur ; for “ Vital actions remain constant so long as the external
actions to which they correspond remain constant;” and no
reference is anywhere made to the principle that whatever causes
sexual separation between dissimilar members of one family, race,
or species tends not only to perpetuate, but to increase their
dissimilarity in the succeeding generations. The view maintained
in the following paper is, I believe, in better accord with the
fundamental principle that “ Unlike units of an aggregate are
sorted into their kinds and parted when uniformly subject to the
same incident forces,” t as is also the teaching of Spencer’s
‘Principles of Biology’ in one passage ; for I have recently dis-
covered that in a single paragraph of this work it is maintained
that, while exposed to the same external conditions, the members
of the same species may be increasingly differentiated, “ until at
length the divergence of constitutions and modes of life become
great enough to lead to segregation of the varieties.’ If
the segregation had been introduced as a necessary condition
without which the divergence of families and races could not
take place, the position taken in this paragraph would have been
essentially the same as the one I have adopted. In the next
section, however, he abandons the position, using the following
words :—“ Through the process of differentiation and integration
which of necessity brings together, or keeps together, like indi-
viduals, and separates unlike ones from them, there must never-
theless be maintained a tolerably uniform species, so long as there
continues a tolerably uniform set of conditions in which it may
exist.’ [The italics are mine. |

I trust my endeavour to contribute something toward the
development of the theory of divergent evolution will not be

* Compare §§ 91, 156, 169, 170.

t See Spencer’s ‘ First Principles,’ § 166, near the end; also a fuller state-
ment in § 169. t See § 90.

Bh Phe pin tea

THROUGH CUMULATIVE SEGREGATION. 197

attributed to any lack of appreciation of what has already been
accomplished. The propounders of a doctrine which has pro-
foundly influenced every department of modern thought need no
praise from me; but as their theory is confessedly incomplete,
and as one of the leaders in the movement has called attention to
the need of a rediscussion of the fundamental factors of evolution,
I offer my suggestions and amendments after prolonged and
careful study.

Physiological Selection and Segregate Fecunditg.

The abstract of Mr. Romanes’s paper on “ Physiological Selec-
tion,” given in ‘ Nature’ August 5th, 12th, and 19th, 1886, did
not come into my hands till the following January, when my
theory of Divergent Evolution through Cumulative Segregation,
which had been gradually developing since the publication of my
paper on “ Diversity of Evolution under One Set of External
Conditions,” was for the most part written out in its present
form. Since then, and with reference to the discussion on
Physiological Selection, I have worked out the algebraic formulas
given in the last chapter, and have introduced explanations of
the same; but at the same time I have removed several chapters
in which the principle of selection was discussed at length, and
have endeavoured to bring the whole within a compass that
would allow of its being published by some scientific society.
In order to attain this end, I reserve for another occasion a dis-
cussion of the principles of Intensive Segregation, under which
name IJ class the different ways in which other principles com-
bine with Segregation in producing Divergent Evolution.

It was my intention to bring together examples of the differ-
ent forms of Segregation discussed, that they might be pub-
lished with the theoretical part; but the large number of pages
found necessary for even the briefest presentation of the prin-
ciples involved, and the fact that Mr. Romanes’s paper has ap-
peared relating to some of the same problems, leads me to present
the results of my studies without further delay. The facts on
which large portions of my theory rest are of the most familiar
kind, and no additional light would be gained though their
numbers were multiplied a hundredfold. Indeed one of the
marked features of my theory is that in its chief outlines it rests
on facts that are universally acknowledged. The aim of the

198 REV. J. T. GULICK ON DIVERGENT EVOLUTION

theory is to show the connection of these facts with divergent
evolution.

Though many divergences appear in our method of treating
the subject, the fundamental theory underlying my Segregate
Fecundity and Mr. Romanes’s Physiological Selection seems to be
very similar, if not the same. The most important differences I
have noticed are, (1) that he seems to regard mutual sterility as
sufficient to account for the separate propagation of species and
varieties thus characterized, without calling in the aid of any

other form of segregation, while I regard it as a Negative form of |

Segregation that would result in the general destruction of all
life if not associated with what I call Positive forms of Segre-
gation ; and (2) that he maintains that ‘ Physiological selection
is almost exclusively a theory of the origin of species, seeing that
it can but very rarely have had anything to do with the formation
of genera, and can never have had anything at all to do with the
formation of families, order, or classes. Hence, the evidence which
we have of the evolutionary influence of physiological selection,
unlike that which we have of the evolutionary influence of natural
selection, is confined within the limits of specific distinctions,” *
while I maintain that Segregation of some form is a necessary
condition for all divergent evolution, and that in fact Segregate
Fecundity in many cases prevents the intercrossing of divergent
forms that, though descended from a common stock, now belong
to different families and orders.

The first of these differences, though of considerable importance,
is, I think, due to the method of presentation, rather than to any
fundamental discrepancy in the theories. The Positive forms of
Segregation are, I judge, assumed to be present, though their
co-operation is not distinctly recognized as a necessary condition
for the breeding of forms that are mutually sterile.

I must, however, confess that I do not see how to reconcile his
statement that “ Physiological selection can never have had any-
thing at all to do with the formation of families, orders, or
classes” with what I believe to be the facts concerning Segre-
gate Fecundity ; and if physiological selection is to be understood
as including Seasonal and perhaps other forms of Segregation,
this passage seems to be still more opposed to the principles of
divergent evolution as I understand them. He certainly could
not have intended to say that mutual fertility between allied

* Linn, Soc. Journ., Zoology, vol. xix. p. 396.

re alone ye ge

THROUGH CUMULATIVE SEGREGATION. 199

genera not otherwise segregated would not have stood in the
way of their becoming different families, and that therefore
mutual sterility has had nothing to do with their continued
divergence ; still he seems to have failed to perceive the im-
portant influence this principle must have had on the divergent
evolution of the higher groups of organisms.

The correspondences in the two papers are notwithstanding
more remarkable than the differences. Of these, the most conspi-
cuous is the use of the word Segregation to express the principle
under consideration.* As I have already pointed out, I used this
word for the same purpose in an article in the ‘ Chrysanthemum,’
published in January 1883 ; and again in the ‘ Chinese Recorder’
for July 1885, where I spoke of the “‘ Law of Segregation rising
out of the very nature of organic activities, bringing together those
similarly endowed,” and causing ‘the division of the survivors
of one stock, occupying one country, into forms differing more
and more widely from each other.”

I trust that my discussion of the various forms of Segregation,
both Negative and Positive, though presented in so condensed a
form, will throw light on the subject of the mutual sterility of
species ; and that in other ways my presentation of the subject
will contribute something, not only to the theory of Physio-
logical Segregation, but to other branches of the general theory
of evolution.

I should here acknowledge (what will, I think, be manifest on
every page of my paper) that my obligations to Darwin and
Wallace are far greater than are indicated by quotations and
references.

I very much regret that I have failed of obtaining a copy of
‘Evolution without Natural Selection, by Charles Dixon ; but,
from his letter in ‘ Nature,’ vol. xxxiii. p. 100, I see that he main-
tains “That isolation can preserve a non-beneficial variation as
effectually as natural selection can preserve a beneficial variation.”
He does not there refer to the fact, which I emphasize, that all
divergence of a permanent character, whether beneficial or non-
beneficial, is dependent on Segeneration either Separative or
Segregative.

* See paper on “Physiological Selection,” Linn. Soc. Journ., Zoology,
vol, xix. pp, 354, 356, 891, 395,

200 REV. J. T. GULICK ON DIVERGENT EVOLUTION

PRELIMINARY DEFINITIONS.

Believing that great obscurity has often been introduced into
the discussion of biological subjects by the use of terms of un-
certain import, I have endeavoured to obtain greater precision by
giving definitions of the terms I have introduced; and for the
sake of indicating what words are thus used with special and
definite meanings, they have been distinguished by capitals. A
few of these definitions are here given, and others will be given in
the body of the paper.

An Intergenerant, or Intergenerating Group, is a group of
individuals so situated and so endowed that they freely cross
with each other.

Segeneration, or Independent Generation. In harmony with the
fundamental doctrines of evolution, I assume that each species
was at one time a single intergenerant ; but we find that many
species are now divided into two or more intergenerants, between
which there is little or no intercrossing. This state of freedom
from crossing I call Segeneration. Segeneration is of two kinds,
Separate Generation and Segregate Generation.

Separate Generation, or Separation, is the indiscriminate
division of a species into groups that are prevented from freely
crossing with each other.

Segregate Generation, or Segregation, is the intergeneration
of similar forms and the prevention of intergeneration between
dissimilar forms.

Select Generation, or Selection, is the partial or complete
exclusion of certain forms from the opportunity to propagate,
while others succeed in propagating. The generation of any
form is select with reference to the non-generation of forms that
fail of propagating, and segregate with reference to the genera-
tion of forms that propagate successfully, but separately.

Adaptational Selection is exclusive generation that depends
upon superior adaptation either to the environment or to other
members of the same species.

Natural Selection is the exclusive generation of those better
fitted to the natural environment, resulting from the failure to
generate of those less fitted.

Artificial Selection is the exclusive generation of those. better
fitted to the rational environment.

Reflexive Selection is the exclusive generation of those better

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THROUGH CUMULATIVE SEGREGATION. 201

fitted to the relations in which the members of the same species
stand to each other. Sexual, Social, and Institutional Selection
are forms of Reflexive Selection.

The Environment is nature lying outside of the Intergenerant.
The influence of the Environment is the sum of the influences
that fall upon the members of an Intergenerant, exclusive of
their influence upon each other. The environment of an inter-
generant includes members of the same species, only when these
members are so near that they exert an influence through com-
petition or otherwise, while at the same time they are so far dif-
ferentiated that they do not intercross; in other words, the
members of the same species can mutually belong to the environ-
ment, only when they have acquired some of the characteristics
of independent species. The same environment extends as far as
the activities that affect or may affect the species extend without
undergoing change.

Change in the Environment is change in the external activities
affecting the species.

Entering a new Environment is change in the territorial dis-
tribution of the species, bringing either all or a portion of its
members within the reach of new influences. This may also be
called Change of Environment.

Change in the Organism, whether producing new adaptations to
the environment or not, should be carefully distinguished from
both of the above described changes.

Change of Relations to the Environment may be produced by
Change in the Environment, or by Entering a new Environment,
or by Change in the Organism.

As great confusion has been occasioned by the terms ‘“ Con-
ditions of Life” and “ External conditions” being used, some-
times for activities outside of the species under consideration,
and sometimes for those within the species (as for example the
influence upon the seed produced by its position in the capsule),
I have tried to avoid their use.

Monotypie Hvolution is any transformation of a species that
does not destroy its unity of type.

Polytypic Evolution or Divergent Evolution is any transforma-
tion of a species in which different types appear in different
sections.

202 REV. J. T. GULICK ON DIVERGENT EVOLUTION
CHAPTER I.
Tur Errects OF SELECTION AND INDEPENDENT GENERATION
CoNTRASTED.

In as far as any theory of evolution fails of giving an explanation
of divergence of character, in so far it fails of explaining the origin
of species. This is the crucial test which must decide the strength
or weakness of every theory that is brought forward to account
for the derivation of many species from one original species, <A
satisfactory theory will not only point out the conditions on
which divergence depends, but will show that these conditions
are the natural result of causes that are already recognized by
science as having influence in the organic world, or that are now
shown to have such influence.

In the present chapter I shall present some reasons for believing
that neither “ Natural Selection,” nor “ Sexual Selection,” nor
“the Advantage of Divergence of Character,” nor ‘“ Difference
of External Conditions,” nor all these taken together, nor any
form of Selection that may be hereafter discovered, is sufficient
to account for Divergence of Character, but that another factor
of equal if not superior importance must be recognized. In sub-
sequent chapters I shall try to trace the causes on which this
additional factor depends, and to indicate as far as possible the
laws and relations under which they appear.

DIVERGENT EVOLUTION NOT EXPLAINED BY NATURAL
SELECTION.

Natural Selection is the exclusive generation of certain forms
through the failure to live and propagate of other kinds that are
less adapted to the environment.

In the case of the breeder, no selection avails anything that
does not result in some degree of exclusion. In the case of
natural selection, where we are not considering ineffectual inten-
tions, the selection is measured by the exclusion. Where there
is no exclusion, there is no selection, and where the exclusion is
great the selection is severe. Moreover, it is self-evident that there
can be no crossing between the best fitted that survive and propa-
gate, and the least fitted that perish without propagating. To this
extent, therefore, the prevention of crossing is complete. And
further, it is evident that those whose meagre fitness gives them
but little opportunity for propagating will have a correspondingly

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THROUGH CUMULATIVE SEGREGATION. 2038

diminished opportunity for crossing with the best fitted ; and so
on through the different grades of fitness, the power to affect the
next generation through having a share in propagating will
measure the power to affect the progeny of the best fitted by
crossing with them. It therefore follows that the freest cross-
ing of the fittest is with the fittest.

Natural Selection therefore proves to be a process in which
the fittest are prevented from crossing with the less fitted through
the exclusion of the less fitted, in proportion to their lack of fitness.
Through the premature death of the least fitted, and the inferior
propagation of the less fitted, there arises a continual prevention
of crossing between the less fitted and the better fitted; and
without this separation the transforming influence of the laws
of organic life would have no power to operate. As Darwin
has pointed out, the results produced by this removal of the
less fitted and separate propagation of the better fitted, closely
correspond with those produced by the breeder, who kills off the
less desirable individuals of his stock before they have opportunity
to breed. The selection of the breeder avails nothing unless it
leads to the determining of the kind that shall breed; and this
he cannot accomplish without preventing free crossing with those
that he does not desire. He must use some method to secure
the separate breeding of the form that he desires to propagate.
We therefore find in both Natural and Artificial Selection the
same fundamental method. In either case, the kind that is to
propagate is determined by the selection, and those that are
not to propagate are in some way excluded. The process may
therefore be called the exclusive breeding of certain kinds; and
Natural Selection may be defined as the exclusive breeding of
those better adapted to the environment.

But if from one stock of horses we wish to develop two distinct
breeds, one of which shall excel in fleetness, and the other in
strength for carrying or drawing burdens, the result will not be
gained by simply preventing all that are inferior in strength or
fleetness from breeding. By this process, which is the Exclusive
Breeding of the desired kinds, we should obtain one breed with
fair powers of strength and fleetness; but the highest results
in either respect would not be gained. Suweh experiments show
that the Exclusive Breeding of other than average forms causes
Monotypic Evolution, and that to secure Divergent or Polytypic
Evolution some other principle must be introduced.

204 REV. J. T. GULICK ON DIVERGENT EVOLUTION

In the case of Natural Selection, the separation it introduces
is between the living and the dead, between the successful and the
unsuccessful. In other words, Natural Selection is the exclusion
of all the forms that through lack of adaptation to the environ-
ment fail of leaving progeny, and therefore in the exclusive
generation of the forms that through better adaptation to the
environment are better able to propagate. Variation with the
Natural Selection of other than average forms may therefore ac-
count for the transformation of an ancient species into a series of
successive species the last of which may now exist in full force ;
but without the aid of Segeneration it will by no means account
for the divergent evolution of any one of these species into a family
of coexisting species.

As I have just shown, Natural Selection is the exclusive genera-
tion of those better fitted to the environment ; and it tends to
the modification of species simply through the generation of the
better fitted forms, while they are prevented from crossing with
the less fitted, which fail of propagating through their lack of
fitness. Now from the very nature of this process, which results
from the success and failure of individuals in appropriating the
resources of the environment, it follows that it cannot be the
cause of separation between the successful competitors, and there-
fore any divergence of character that arises between the different
groups of the successful cannot be attributed to Natural Selec-
tion. Natural Selection explains the prevention of crossing
between the fitted and the unfitted, and shows how the succes-
sive generations of a species may gradually depart from the
original type, becoming in time a different species; but 7% can-
not explain the divergences that arise between those that have,
by the fact of successful propagation, proved their fitness. It
depends on superiority of adaptation to the environment, and
tends to produce increasing adaptation; but divergent kinds of
adaptation are not necessary conditions for it, and it cannot be
the cause of increasing divergence between the incipient kinds that
otherwise arise.

DIVERGENT EVOLUTION NOT EXPLAINED BY “ THE ADVANTAGE
OF DIVERGENCE OF CHARACTER.”

Two sections of the 4th chapter of the ‘Origin of Species ’
are given to the discussion of the ‘ principle of benefit being
derived from divergence of character,” which it is maintained

ee ee ee, >»

THROUGH CUMULATIVE SEGREGATION. 205

“will generally lead to the most different or divergent varia-
tions being preserved and accumulated by natural selection.”
Now it cannot be doubted that ability to appropriate unused
resources would be an advantage to any members of a com-
munity pressed for food ; but I do not see how the divergence
that would enable them to appropriate, for example, a new kind
of food can be accumulated while free crossing continues; and
Natural Selection cannot prevent the free crossing of competitors
who leave progeny.

Having found that the evolution of the fitted is secured
through the prevention of crossing between the better fitted
and the less fitted, can we believe that the evolution of a
special race, regularly transmitting a special kind of fitness,
can be realized without any prevention of crossing with other
races that have no power to transmit that special kind of
fitness? Can we suppose that any advantage, derived from
new powers that prevent severe competition with kindred, can
be permanently transmitted through succeeding generations to
one small section of the species while there is free crossing
equally distributed between all the families of the species? Is
it not apparent that the terms of this supposition are inconsis-
tent with the fundamental laws of heredity ? Does not inheri-
tance follow the lines of consanguinity ; and when consanguinity
is widely diffused, can inheritance be closely limited? When
there is free crossing between the families of one species will not
any peculiarity that appears in one family either be neutralized
by crosses with families possessing the opposite quality, or being
preserved by natural selection, while the opposite quality is
gradually excluded, will not the new quality gradually extend to
all the branches of the species; so that, in this way or in that,
increasing divergence of form will be prevented ?

If the advantage of freedom from competition in any given
variation depends on the possession, in some degree, of new
adaptations to unappropriated resources, there must be some
cause that favours the breeding together of those thus specially
endowed, and interferes in some degree with their crossing with
other variations, or, failing of this, the special advantage will in
succeeding generations be lost. As some degree of Independent
Generation is necessary for the continuance of the advantage, it
is evident that the same condition is necessary for the accumu-
lation through Natural Selection of the powers on which the

LINN. JOURN.—ZOOLOGY, VOL. XX. Ls

206 REV. J. T. GULICK ON DIVERGENT EVOLUTION

advantage depends. The advantage of divergence of character
cannot be retained by those that fail to retain the divergent charac-
ter ; and divergent character cannot be retained by those that are
constantly crossing with other kinds; and the prevention of free
crossing between those that are equally successful is im no way
secured by Natural Selection.

NATURAL SELECTION witH GREAT DIFFERENCE IN EXTERNAL
CONDITIONS NOT SUFFICIENT TO EXPLAIN DIVERGENT EVOLUTION.

The insufficiency of Natural Selection without Segeneration
to account for divergent evolution in an area where the external
conditions are nearly uniform may be admitted by some who
will claim that the case is quite otherwise when a species ranges
freely over an area in which it is subjected to strongly contrasted
conditions. It may be claimed that Diversity of Natural Selection
resulting from a great difference in external nature is sufficient
to account for divergent evolution without any Segeneration.

In the discussion of this subject, important light can be gained
_ by referring to the experience of the breeder. This experience,
in as far as it relates to the subject of Separation in the
production of divergent breeds, may be arranged under three
heads. 1st. Diversity of Selection without Separation. 2nd.
Separation without Diversity of Selection. 8rd. Separation more
or less complete with Diversity of Selection.

As the full discussion of these points is impossible here, and as
there is probably but little difference of opinion in regard to what
the results would be, I shall content myself with a simple state-
ment of what I believe the experience of breeders shows. Dif-
ference in the standards of Selectiou without Separation can
avail nothing in creating divergence of types; while Separation
without difference in the standards of Selection will avail some-
thing, though food and external conditions are kept the same;
but to secure the greatest divergence, in a given time, there must
be both Diversity of Selection and complete Separation. In the
case of Separation without Diversity of Selection there is room
for difference of opinion ; for the examples that some would claim
as proving that there is often divergence without Diversity of
Selection and without difference in external conditions may be
attributed by others to unconscious Selection. It is granted by
everyone that no skill in selecting the animals that possess the
desired qualities will have any effect in establishing a new breed,

ss,

THROUGH CUMULATIVE SEGREGATION. 207

unless the selected animals are prevented from breeding with
others that are deficient in the desired qualities. We further
find that while Separation is an absolutely essential condition for
this divergence, Diversity of Selection is not so essential. This
is illustrated in the case of the slightly different types that
are presented by the wild cattle found in the different parks of
England,* a phenomenon which can hardly be attributed to any
diversity in the environment.

In artificial breeding universal experience teaches that Varia-
tion and Selection, without Separation, do not produce diver-
gence of races. The separate breeding of different ciasses
of variation is a necessary condition for the accumulation of
divergent variation; and wherever the separate breeding of
different classes of variation is secured, there divergence of
character is the result. In other words, Segregate Breeding is
necessary to divergent evolution in gamogenetic animals.
Moreover, we have every reason to believe that the same law
holds good throughout the whole organic world. The generating
together of similars, with the exclusion or separation of dissimi-
lars, is the central necessity in all evolution by descent, whether
monotypic or polytypic; and whatever causes the separate genera-
tion of different classes of variation will be the cause of divergent
evolution. That is, wherever this condition is added to the per-
manent laws of organic life, there divergence will follow. As
we have already seen, Natural Selection or the Survival of the
Fittest necessarily separates between the survivors and the non-
survivors, between the best fitted and the least fitted, and is,
therefore, the cause of monotypic transformation; but it cannot
be the cause of separation between the different families of those
that survive, and, therefore, cannot be the cause of divergence
of character between these families. But we find that divergence
of character often arises between the branches of one stock,
and in many cases this divergence increases till well-marked
varieties are established. If, therefore, the general principle we
have just stated is true, there must be certain causes producing the

* See Darwin’s ‘ Variation under Domestication,’ chapter xv. 2nd page.

+ In a subsequent paper I shall show how it is that Separate Breeding, long
continued, inevitably ends in Segregate Breeding. In this chapter I confine my
attention more especially to Separate Breeding when combined with Diversity
of Selection in the different sections, for it is evident that this will produce
Segregate Breeding.

Lie

208 REV. J. T. GULICK ON DIVERGENT EVOLUTION

Independent Generation of these forms; and, if we can discover
these causes and trace them to general principles, they will, in
connection with the laws of Variation and Selection, explain
divergent evolution, that is the transformation of one form into
many forms, of one species into many species. As community
of evolution arises where there is community of breeding between
those that through superior fitness have opportunity to propa-
gate, so I believe it will be found that divergent evolution arises
where there is separate breeding of the different classes of the
successful. In other words, Exclusive Breeding of other than
average forms causes Monotypic Evolution, and Segregate
Breeding causes Divergent or Polytypic Evolution.

The facts of geographical distribution seem to me to justify
the following statements :-—

(1) A species exposed to different conditions in the different
parts of the area over which it is distributed is not represented
by divergent forms when free interbreeding exists between the
inhabitants of the different districts. In other words, Diversity
of Natural Selection without Separation does not produce diver-
gent evolution.

(2) We find many cases in which areas, corresponding in the
character of the environment, but separated from each other by
important barriers, are the homes of divergent forms of the same
or allied species.

(3) In cases where the separation has been long continued,
and the external conditions are the most diverse in points that
involve diversity of adaptation, there we find the most decided
divergences in the organic forms. That is, where Separation
and Divergent Selection have long acted, the results are found
to be the greatest. The lst and 8rd of these propositions will
probably be disputed by few if by any. The proof of the 2nd
is found wherever a set of closely allied organisms is so distri-
buted over a territory that each species and variety occupies its
own narrow district, within which it_is shut by barriers that
restrain its distribution, while each species of the environing
types is distributed over the whole territory. The distribution
of terrestrial mollusks on the Sandwich Islands presents a great
body of facts of this kind.

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THROUGH CUMULATIVE SEGREGATION. 209

SELECTION OF EVERY KIND INSUFFICIENT TO ACCOUNT FOR
DIVERGENT EVOLUTION.

Though I have no reason to doubt the importance of Sexual
Selection in promoting the transformation of many species, I
think I can show that unless combined with some separative or
segregative influence, that prevents free intercrossing, it can
avail nothing in producing a diversity of races from one stock.
In the nature of its action Sexual Selection is simply exclusive.
It is the exclusive breeding of those better fitted to the sexual
instincts of the species, resulting from the failure to breed of the
less fitted. It therefore indicates a method of separation between
the better fitted and the less fitted; but it gives no explanation
of separation between those that are equally successful in pro-
pagating.

I maintain that in a great number of animal species there are
sexual and social instincts that prevent the free crossing of
clearly marked races; but as these segregative instincts are
rarely the cause of failure to propagate, and since when they are
the cause of failure the failure is as likely to fall on one kind as
on another, I conclude that the Segregate Breeding resulting
from these instincts cannot be classed as either Sexual or Social
Selection. Reflexive Selection in all its forms is, ike Natural
Selection, the result of success and failure in vital processes
through which the successful propagate without crossing with
the unsuccessful; but it in no way secures the breeding in
separate groups of those that are successful in propagating. The
exclusion of certain competitors from breeding is a very different
process from the separation of the successful competitors into
different groups that are prevented from intercrossing, and
whose competition even is often limited to the members of the
same group. Sexual Selection, like other forms of Reflexive
Selection, can extend only as far as members of the same species
act on each other. If the individuals of two groups have
through difference in their tastes ceased to compete with each
other in seeking mates, they are already subject to different and
divergent forms of Sexual Selection ; and is there any reason to
attribute this difference in their tastes to the fact that, when
there was but one group and the tastes of all were conformed to
u single standard, some of the competitors failed of propagating,
through being crowded aside by those more successful? Jf the

210 REV. J. T. GULICK ON DIVERGENT EVOLUTION

failure of the unsuccessful cannot be the cause of separation be-
tween the different kinds of the successful, then Selection, whether
Natural, or Reflexive, or of any other kind, cannot be the cause of
Divergent Evolution, except as co-operating with some cause of
Independent Generation.

The failure of Sexual Selection, without Separation or Segrega-
tion, to account for divergent evolution, will perhaps be made
clearer to some minds by considering some of the particular
conditions under which it occurs. Suppose, for instance, that in
some species of humming-bird there occurs a slight variation in
the form or colour of the tail-feathers of the male that adds to
the beauty of the individuals possessing the new character and
rendering them more attractive to the females. We can see
that they might have an advantage over their rivals in leaving
progeny, and that the variety might in that way gradually gain
the ascendency, and the beauty of the markings become more
and more completely defined; but under such conditions what
could prevent the whole species from being gradually transformed ?
Unless there was some separative or segregative principle that
prevented the new variety from crossing with the others, the
species would remain but one, though changed in some of its
characters. We should have transformation without divergence.

The same must be true of Institutional Selection. It may be
the cause of transformation ; but it cannot be the cause of diver-
gent evolution, unless there are added to it other causes that pro-
duce divergence in the character of the forms selected, and the
separate breeding of the different groups of forms thus selected.
A single illustration will set in a clear light the limitation in
the influence of Institutional as well as all other Selection. In
primitive communities the deaf are but little cared for, and owing
to the great disadvantages of their position their opportunities
for gaining subsistence, and therefore for rearing families, are
ereatly diminished: this is Natural Selection. Again, those who
are at so great a disadvantage in communicating with their com-
panions will be also at a disadvantage in finding consorts: this
we may call Social Selection. Again, a community might either
by law or by strict custom prevent the marriage of the deaf:
this would be Institutional Selection. Any one of these forms of
selection might be pressed so far as to be the meaus of increasing
the average power of hearing in the community in succeeding
generations; but it could never be the cause of two divergent

THROUGH CUMULATIVE SEGREGATION. 211

races, one with good powers of hearing and the other with an
increasing liability to deafness. To secure such divergence it is
necessary that segregative influences should be introduced, such
as have been most amply furnished by the modern system of
education for the deaf. Under these influences those endowed
with hearing and those without hearing have been separated into
two communities, the members of each having but little oppor-
tunity for acquaintance beyond the limits of that community ;
each community having separate schools, separate newspapers,
and to some extent a separate language. As the result of this
segregation marriages between the two classes have been greatly
diminished ; and little by little two races are arising, the hearing
race and the deaf race.*

REASONS oF A GENERAL CHARACTER FOR CONSIDERING SELECTION
witHouT INDEPENDENT GENERATION AN UNSATISFACTORY
Expnanation oF Divercent EvoLurion.

1. The divergence is often confined to characters which seem
to have no possible relations of adaptation either to the environ-
ment or to other members of the species, and, therefore, to be
independent of both Natural and Reflexive Selection.

2. Divergence relating to adaptive characters successfully
propagated involves different kinds rather than different degrees
of adaptation and advantage; and, as Adaptational Selection de-
pends on the difference of degrees of advantage, it cannot account
for the divergence of forms possessing equal degrees of ad-
vantage.

3. In the very nature of its action, we see that Adaptational
Selection unaccompanied by Independent Generation must pro-
duce essentially monotypic transformation.

4. In artificial breeding, Independent Generation is found to be
an essential condition for the production of divergent races ; and
there is no reason to doubt that the same law holds good in the
divergence of natural forms.

5. The general fact that species possessing high powers and
large opportunities for migration occupy large areas, while those

* See paper by Alexander Graham Bell, read before the National Academy
of Sciences, November 13, 18838, upon the “ Formation of a Deaf Variety of the
Human Race;” also a review of the same in ‘ The Popular Science Monthly,’
vol. xxvii. p. 15, entitled “Can Man be Modified by Selection ?”

214 REV. J. T. GULICK ON DIVERGENT EVOLUTION

the progenitors of man were deprived of their hairy coat by
Sexual Selection that was, in its earlier stages, disadvantageous.

It is therefore evident that the simple fact of divergence in
any case is not a sufficient ground for assuming that the divergent
form has an advantage over the type from which it diverges. We
may, however, be sure that there is some cause or combination of
causes that facilitates the intergenerating of those similarly
endowed, and hinders their crossing with other kinds; and if we
can discover the cause of this Segregate Generation, we shall
have an explanation of one part of the process by which the
forms thus endowed are becoming a distinct race.

SEPARATION AND SEGREGATION WITH THE PRINCIPLE
oF INTENSION.

It will contribute to clearness in our discussion if we can gain
definite conceptions of the conditions that are necessarily in-
volved in Separate and Segregate Breeding.

Separate Generation, which for convenience I call Separation,
implies :—

1st. The indiscriminate separation of the members of a species
into different sections that are prevented from freely crossing
with each other.

2nd. The aggregation of the members of one section ; that is,
their being brought into conditions of time and place that allow
of their freely crossing.

3rd. The integration of the members of each section into one
intergenerating group, through the operation of functional adap-
tations by which the members of each section freely cross with
each other. This analysis of the process shows that it may
depend upon a great variety of causes, working together in a
very complex way. We shall hereafter find that the causes of
separation may operate in such a way that no aggregation or
propagation takes place amongst the members that are separated
from the old stock; but in such cases there is no Separate
Generation, and therefore no Separation in the sense in which I
use the word.

Segregate Generation also consists of separation, aggregation,
and integration; but it differs from Separate Generation in that
in the latter the separation is indiscriminate, while in the former
there is a more or less pronounced bringing together of those
that are similarly endowed, with separation of those that are

= THROUGH CUMULATIVE SEGREGATION. 215

dissimilar. Segregate Generation is therefore the separation of
dissimilars, with the aggregation and integration of similars. As
we have already seen, Segregate Breeding may be produced by
Separate Breeding accompanied by Diversity of Natural Selection
in the different sections. It is also evident that any other cause
that develops in one or more of the separate sections of the
species characters that are not found in the other sections will
produce Segregate Breeding. Such cases are Diversity of Selec-
tion of other forms than Natural Selection, Diversity in the
inherited effects of Use and Disuse (unless physiologists have
been mistaken in supposing that there are any such effects), and
Diversity in the inherited characters derived from the Direct
Effects of the Environment (unless, again, Weismann is right and
the general belief wrong). Segregate Breeding may, moreover,
be produced directly by the very way in which the separation of
the different sections is secured. One of the best examples of
this kind of Segregation is seen in what I call Industrial Segre-
gation, where the members of a species are distributed according
to their endowments, those of similar endowments being brought
together. In such cases, Segregation is introduced as soon as
the Separation, without depending on the subsequent action of
the environment, or on diverse forms of Use, or of Selection ;
though there can be no doubt that,in the great majority of
cases, Diversity of Use and Diversity of Selection of some kind
will in time come in to intensify the result.

There is another invariable sequence which it is necessary we
should keep in mind, if we would understand the relation in
which these two principles stand to each other. I refer to the
certainty that all prolonged Separate Breeding will be trans-
formed into Segregate Breeding. In other words, indiscriminate
separation, in which there is no apparent difference in the dif-
ferent groups, 1s in time found to be a separation in which there
is a decided difference in the different groups. Whenever a
sufficient number of the same species to ensure propagation are
brought together in an isolated position, Separate Generation is
the result ; and, if this Separate Generation is long-continued,
we have reason to believe, it always passes into Segregate Genera-
tion with divergent evolution. The fundamental cause for this
seems to lie in the fact that no two portions of a species possess
exactly the same average character, and that the initial differences
are for ever reacting on the environment and on each other in

214 REV. J. T. GULICK ON DIVERGENT EVOLUTION

the progenitors of man were deprived of their hairy coat by
Sexual Selection that was, in its earlier stages, disadvantageous.

SEPARATION AND SEGREGATION WITH THE PRINCIPLE
oF INTENSION.

It will contribute to clearness in our discussion if we can gain
definite conceptions of the conditions that are necessarily in-
volved in Separate and Segregate Breeding.

Separate Generation, which for convenience I call Separation,
implies :—

1st. The indiscriminate separation of the members of a species
into different sections that are prevented from freely crossing
with each other.

2nd. The aggregation of the members of one section ; that is,
their being brought into conditions of time and place that allow
of their freely crossing.

~ THROUGH CUMULATIVE SEGREGATION. 215

dissimilar. Segregate Generation is therefore the separation of
dissimilars, with the aggregation and integration of similars. As
we have already seen, Segregate Breeding may be produced by
Separate Breeding accompanied by Diversity of Natural Selection
in the different sections. It is also evident that any other cause
that develops in one or more of the separate sections of the
species characters that are not found in the other sections will
produce Segregate Breeding. Such cases are Diversity of Selec-
tion of other forms than Natural Selection, Diversity in the
inherited effects of Use and Disuse (unless physiologists have
been mistaken in supposing that there are any such effects), and
Diversity in the inherited characters derived from the Direct
Effects of the Environment (unless, again, Weismann is right and
the general belief wrong). Segregate Breeding may, moreover,
be produced directly by the very way in which the separation of
the different sections is secured. One of the best examples of
this kind of Segregation is seen in what I call Industrial Segre-
gation, where the members of a species are distributed according
to their endowments, those of similar endowments being brought
together. In such cases, Segregation is introduced as soon as
the Separation, without depending on the subsequent action of
the environment, or on diverse forms of Use, or of Selection ;
though there can be no doubt that, in the great majority of
cases, Diversity of Use and Diversity of Selection of some kind
will in time come in to intensify the result.

There is another invariable sequence which it is necessary we
should keep in mind, if we would understand the relation in
which these two principles stand to each other. I refer to the
certainty that all prolonged Separate Breeding will be trans-
formed into Segregate Breeding. In other words, indiscriminate
separation, in which there is no apparent difference in the dif-
ferent groups, is in time found to be a separation in which there
is a decided difference in the different groups. Whenever a
sufficient number of the same species to ensure propagation are
brought together in an isolated position, Separate Generation is
the result ; and, if this Separate Generation is long-continued,
we have reason to believe, it always passes into Segregate Genera-
tion with divergent evolution. The fundamental cause for this
seems to lie in the fact that no two portions of a species possess
exactly the same average character, and that the initial differences
are for ever reacting on the environment and on each other in

216 REV. J. T. GULICK ON DIVERGENT EVOLUTION

such a way as to ensure increasing divergence in each successive
generation, as long as the individuals of the two groups are kept
from intergenerating. In my paper on Diversity of Evolution
under one Set of External Conditions, I spoke of this principle of
divergence as “ Separation with Variation ;” but in order to dis-
tinguish the antecedent condition, which is Separation, from the
result, which is something more than Variation, I now call the
certainty that some form of divergent transformation will arise
when intergeneration is prevented the principle of Intension ;
and Segregation produced by independent transformation I call
Intensive Segregation.

As Separate and Segregate Generation are so closely related,
I have, in order to avoid a multiplication of terms, classified the
two principles together under the general term Segregation. In
my discussion of the causes of Segregation I shall, however,
endeavour to determine concerning each class of causes whether
they are primarily Separative or Segregative.

A full discussion of tie causes of Segregation would require
that under each combination of causes to which we give a dis-
tinctive name we should show :—

1. How the Independent Generation is produced.

2. How the difference of character in the different sections is
produced.

3. How the aggregation in place bringing together the members
of each section is produced.

4. How the correspondence in times and seasons necessary for
intergeneration is secured within each section.

5. How the correspondence of community and of Sexual and
Social instincts necessary for intergeneration is secured within
each section.

6. How the correspondence in structure, in dimensions, and
in the mutual potentiality of the sexual elements necessary for
intergeneration is secured within each section.

It will, however, be observed that, with the exception of the
two first, these questions relate to the necessary conditions that
must always exist in the case of every intergenerating group;
and as it is evident that intergeneration in some degree must be
the normal condition in every sexual, that is in every gamo-
genetic, species, we may here assume that all the conditions
necessary to intergeneration exist, except so far as they have
been disturbed by causes producing Segeneration. In tracing

a ee ee ae

RR elgie tans esate > ooa) aera dese fo)

THROUGH CUMULATIVE SEGREGATION. 217

the causes of Segregation it will therefore be sufficient if in
each class of cases we give the cause of Segeneration, showing
why the same cause does not prevent all intergeneration, and
explain the difference of character in the different sections pro-
duced by the Segeneration. In full accord with the implications
of the theory of evolution, we proceed on the assumption that
intergeneration was the original condition of every species, and
that the intergeneration of those that are brought together under
favourable circumstances may be taken for granted, unless there
is some special cause that prevents. All that is necessary to
produce Separation is the failure of any one of the many con-
ditions on which free-crossing depends, in such a way, and to
such a degree, that the species falls into two or more sections
between which crossing is interrupted, without its being inter-
rupted within the bounds of each section. And all that is
necessary to produce Segregation is that to Separation should be
added some cause that secures difference of character in the
different sections. And as Separation long continued inevitably
ends in Segregation through the development of difference of
character in the different sections, we need not in our classifi-
cation set them wholly apart, though for the sake of clearly
recognizing the difference it will be well to note in each class of
causes whether the primary effect is Separation or Segregation.

CUMULATIVE SEGREGATION AND THE CLASSIFICATION OF
Ivs DIFFERENT Forms.

The fundamental law to which I would call attention may be
expressed in the following formula :—Cumulative Segregation
produces accumulated divergence; and accumulated divergence
produces permanent Segregation; and the Segregate subdivision
of those permanently Segregated produces the divisions and sub-
divisions of organic phyla. If, then, we can discover the causes
of Segregation, we shall understand the causes of a wide range
of phenomena; for this is the fundamental principle in the for-
mation of varieties, species, genera, families, orders, and all greater
divergences that have been produced in the descendants of
common ancestry.

In treating of the causes of Segregation, I have found it con-
venient to make two distinct classifications. In the one the

218 REV. J. T. GULICK ON DIVERGENT EVOLUTION

fundamental distinction is between segregation produced by the
purpose of man, which I call

RaTIONAL SEGREGATION, in its two forms:

ARTIFICIAL SEGREGATION,
INSTITUTIONAL SEGREGATION,

and that produced by nature outside of man, which I call

RESPONSIVE SEGREGATION ;

while any of these forms of Segregation may be intensified by
Independent transformation through the principles of Diversity
of Selection, Diversity of Use, or Diversity of Direct Effects of
the Environment; and the combined action of Segregation witl
these and other principles of transformation I call

INTENSIVE SEGREGATION.

In the other classification, the fundamental distinction is be-
tween segregation arising from the relations in which the organism
stands to the environment, which I call

Environal Segregation,

and segregation arising from the relations in which the members
of the same species stand to each other, which I call

Reflexive Segregation ;

while any form of segregation belonging to either of these classes
may be enhanced by one or more of the forms of Intension, and
thus present what I call

Intensive Segregation.

THe EFFECTS OF SEGREGATION.

The effects of Segregation can be studied to advantage in the
vast experience that has been accumulated in the domestication
of plants and animals. |

Artificial Segregation is caused by the relations in which the
organism stands to the rational environment, that is to the
purposes of man. In other words, Artificial Segregation is the
rational form of Environal Segregation. Though the bearing of
Segregation on the evolution of species in a state of nature has
been for the most part overlooked, its effects have been quite
familiar to the breeder of domestic races.

THROUGH CUMULATIVE SEGREGATION. 219

As a convenient method of illustration, let us consider the
different results that will be gained according as we subject the
same ten pairs of wild rock-pigeons to one or the other of the
following methods of treatment.

In the first experiment let the treatment be as follows :—Let
ten aviaries be prepared ; and in each aviary put one male with
the female that most nearly resembles it. When the young of
each aviary arrive at maturity, let them be inspected, and if any
individual resembles the inmates of one of the other aviaries
more than the inmates of the aviary in which it was produced,
let it be placed with those it most closely resembles. If any
unusual variation arises, let it be placed in a new aviary, and let
the one of the other sex that most closely resembles it in that
respect be placed with it. When the crowding in any aviary
becomes injurious to the health of the birds let the numbers be
indiscriminately reduced. Let this process be continued many
generations, the inmates in all the aviaries being fed on the same
food, and in every respect treated alike, and what will be the
result P

No experienced breeder will hesitate in assuring us that under
such treatment a multitude of varieties will be formed, many of
which will be very widely divergent from the original wild stock.
In other words, Cumulative Segregation will produce accumulated
divergence, though there is no Selection in the sense in which
Natural Selection is Selection.

Again, let us take the same ten pairs, and putting them into
one large aviary, let them breed freely together without any

' Segregative influence coming in to affect the result ; and who

does not know that the type would remain essentially one
though a considerable range of individual variation might arise.
That is, without Segregation no divergence of type will arise.

THE Natrurat Law or CUMULATIVE SEGREGATION.

I shall now show that there is in nature a law of Cumulative
Segregation. There are large classes of activities in the organism
and in the environment that conspire to produce Segregate
Breeding ; and to produce it in such a way that, in a vast mul-
titude of cases, it becomes a permanent fact, which no cause
that we are acquainted with can ever obliterate. Moreover,
when one form of Segregation has become fully established, we

220 REV. J. T. GULICK ON DIVERGENT EVOLUTION

find that the different branches are liable to be again subjected
to segregative influences, by which each branch is subdivided,
and in time differentiated into divergent forms that are not liable
to intercross in a state of nature.

Now, as we have just pointed out, we know, from the funda-
mental laws of the organic world, that Cumulative Segregation
of this kind must produce Cumulative Divergence of Types.

The Segregation that results from the natural causes enume-
rated in this paper is cumulative in two respects. In the first
place, every new form of segregation that now appears depends
on, and is superimposed upon, forms of Segregation that have
been previously induced ; for when Negative Segregation arises,
and the varieties of a species become less and less fertile with
each other, the complete infertility that has existed between them
and some other species does not disappear, nor does the Positive
Segregation (that is, the prevention of the consorting of the
species characterized by this mutual incapacity) cease. The
means by which the males and females of one species find each
other are not abrogated when the species falls into segregated
varieties. In the second place, whenever Segregation is directly
produced by some quality of the organism, variations that possess
the endowment in a superior degree will have a larger share in
producing the segregated forms of the next generation, and
accordingly the Segregative endowment of the next generation
will be greater than that of the present generation ; and so with
each successive generation the segregation will become in-
creasingly complete.

The principle of Cumulative Segregation, first in its inde-
pendent action, and still further when combined with the differ-
ent principles by which the divergence of the segregated branches
is intensified, gives a formal explanation of the ever-expanding
diversities of the organic world. It shows how varieties arise
and pass into species, how species pass into genera, genera into
families, families into orders, and orders into classes and the
higher divisions, as far as evolution by descent extends. It
brings to light the dependence of this whole process on the
influences that produce segregation; and shows how these
influences, added to Variation, Heredity, and the other acknow-
ledged powers residing in organisms, must produce the phe-
nemena of divergent evolution.

Pe ee ee Ee ee ee

THROUGH CUMULATIVE SEGREGATION. 221

CoMPETITIVE DiIsRUPTION.

Before entering upon the discussion of the direct causes of
Cumulative Segregation, let us briefly consider a law resulting
from the competition of kindred with each other, which brings to
light the fact that such competition is one of the most important
factors in preparing the way for, and in giving intensity to, the
activities that lead to Segregation and divergent evolution. It
is manifest that competition for identical resources and Geogra-
phical Segregation are conditions which cannot exist at the same
time between the same members of any species; but it is also
manifest that, when there are no natural barriers separating the
different districts of an area part of which is occupied by a
species, pressure for food through a great increase in the popu-
lation will tend to distribute the species over the whole area;
and, if the available resources in the different districts are consi-
derably diverse, the overflow of population from the crowded
district will be subjected to a necessary change of habits; and
thus, through competition, there will be the disruption of old
relations to the environment, and the bringing in of conditions
that give the highest efficiency and the fullest opportunity to all
the activities that produce Segregation. In the case of animals,
no condition can tend more strongly to produce migration than
scarcity of food in the old habitat ; and in the case of both plants
aud animals, a great increase in the numbers that are exposed to
the winds, currents, and other transporting influences of the
environment increases the probability that individuals will be
carried to new districts where circumstances will allow of their
multiplying, and where they will, at the same time, be prevented
from crossing with the original stock. In many cases the segre-
gation thus brought about will be in districts where the environ-
ment is the same, and in other cases the pressure for food or
other resources will lead portions of the species to take up new
habits in the effort to appropriate resources not previously used ;
and through these new habits they will often be segregated from
those maintaining the original habits. I shall hereafter show
that in both these cases there is a tendency to divergent
evolution.

Tat one time thought of describing this principle as a form
of Segregation, calling it dominational segregation ; but fuller
reflection convinces me that the domination of the strong over

LINN. JOURN.— ZOOLOGY, VoL, Xx. 18

222 REV. J. T. GULICK ON DIVERGENT EVOLUTION

the weak is not a form of Segregation, but rather a cause that
prepares the way for Segregation, by forcing portions of the
community out of their inherited relations to the environment.

CHAPTER III.

DESCRIPTION AND CLASSIFICATION OF THE CAUSES OF
CUMULATIVE SEGREGATION.*

A. ENVIRONAL SEGREGATION.

Environal Segregation is Segregation arising from the relations
in which the organism stands to the environment.

It includes four classes, which I call Industrial, Chronal,
Spatial, and Artificial Segregation.}

(a) INDUSTRIAL SEGREGATION

is Segregation arising from the activities by which the organism
protects itself against adverse influences in the environment, or
by which it finds and appropriates special resources in the
environment. .

The different forms of Industrial Segregation are Sustentational,
Protectional, and Niditicational Segregation.

For the production of Industrial Segregation it is necessary
that there should be, in the same environment, a diversity of
fully and of approximately available resources more or less
separated from each other, and in the organism some diversity
of adaptation to these resources, accompanied by powers of
search and of discrimination, by which it is able to find the
resources for which it is best fitted and to adhere to the same
when found.

The relation in which these causes stand to each other and
through which they produce segregation may be described as
separation according to endowment produced by endeavour
according to endowment.

It is evident that if initial variation presents in any case a
diversity of adaptations to surrounding resources that cannot be

* In the following chapters numerals are attached to what I consider sepa-
rate causes of segregation independent of human purpose.

t Francis Galton has suggested another class, which might appropriately be
called Fertilizational Segregation.

THROUGH CUMULATIVE SEGREGATION. 923

followed without separating those differently endowed, we shall
have, in the very nature of such variation, a cause of segregation
and of divergent evolution. Some slight variation in the di-
gestive powers of a few individuals makes it possible for them to
live exclusively on some abundant form of food, which the species
has heretofore only occasionally tasted. In the pressure for
food that arises in a crowded community these take up their
permanent abode where the new form of food is most accessible,
and thus separate themselves from the original form of the
species. These similarly endowed forms will therefore breed
together, and the offspring will, according to the law of Diver-
gence through Segregation, be still better adapted to the new
form of food. And this increasing adaptation, with increasing
divergence, might continue for many generations, though every
individual should come to maturity and propagate; that is, though
there were no enhancing of the effect through Diversity of
Selection, or indeed through any other cause producing Intensive
Segregation. And when different forms of Intension do arise,
they may be entirely independent of change in the environment,
the only change being in the forms or functions of the organism.

In choosing a name for this form of Segregation I first thought
of calling it Physiological or Functional Segregation. But such
a name is, on closer examination, found to imply both too much
and too little; for on the one hand there is probably no form of
segregation that is not in some way or in some degree due to
physiological or functional causes, and on the other hand this
special form of segregation is as dependent on psychological
causes which guide the organism in finding and in adhering to
the situation for which it is best fitted, as it is on the initial
divergence of the more strictly physiological adaptations by
which it is able to appropriate and assimilate the peculiar form
of resource. In the case of freely moving animals, the psycho-
logical guidance is an essential factor in the success of the in-
dividual ; while in the case of plants and low types of animal life,
the suitable situation is reached by a wide distribution of a vast
number of seeds, spores, or germs, and the same situation is
maintained by a loss of migrational power as soon as the germs
begin to develop. In these lower organisms it is evident that
the success of the individual must depend on its physiological
rather than on its psychological adaptations; and if an initial
divergence of adaptations results in a slight difference in the kinds

18*

224, REY. J. T. GULICK ON DIVERGENT EVOLUTION

that succeed in germinating in contrasted situations, the difference
is directly due to a Diversity in the forms of Natural Selection
affecting the seed, and the Separation is what I hereafter describe
as Local Separation passing into Local Segregation. We there-
fore see that what I here call Industrial Segregation depends on
psychological powers acting in aid of divergent physiological
adaptations to the environment, or in aid of adaptations that are
put to different uses.

Observation shows that there is a multitude of cases in which
Endeavour according to Endowment brings together those simi-
larly endowed and causes them to breed together; and when the
species is thus divided into two or more groups somewhat differ-
ently endowed, there will certainly be an increased divergence in
the offspring of the parents thus Segregated ; and so on in each
successive generation, as long as the individuals find their places
according to their endowments, and thus propagate with those
similarly endowed, there will be accumulated divergence in the
next generation. Indeed it is evident that Endeavour according
to Endowment may produce under one environment what Natural
Selection produces when aided by local separation in different
environments. As it produces the separate breeding of a diver-
gent form without involving the destruction of contrasted forms,
it is often the direct cause of divergent transformations; while
Natural Selection, which results in the separate breeding of the
fitted through the failure of the unfitted, can never be the cause
of divergence, unless there are concurrent causes that produce
both divergent forms of Natural Selection, and the separate
breeding of the different kinds of variations thus selected.

Suetudinal Intension——Another law is usually believed to be
connected with Endeavour which, if it exists, must conspire to
enhance its tendency to produce divergent evolution. I refer
to the influence which the habitual endeavour of the parents has
on the inherited powers of the offspring. We may call it the
law of Endowment of Offspring according to the Exercise or
Endeavour of Parents, or more briefly Suetudinal Intension.
The inherited effects of use and disuse have been fully recognized
by Darwin, Spencer, Cope, Murphy, and others, and need not
here be discussed. The one point to which I wish to call atten-
tion is, that in order that diversity of use should produce diver-
gent evolution, it is necessary that free crossing should be pre-
vented between the different sections of the species in which the

Se a a

THROUGH CUMULATIVE SEGREGATION. pi)

diversity of use is found. Now this condition of separate breed-
ing is often secured by Industrial Segregation. In other words,
the law of Endeavour according to Endowment, often secures
Separation according to endowment; and this gives an oppor-
tunity for the inheritable effects of diversity of endeavour to be
accumulated in successive generations; and in this way both
laws conspire to produce divergent evolution.

In the relation of these two factors we have a striking example
of the peculiar interdependence of vital phenomena. Diversity
of endowment is the cause of diversity of endeavour and of
Segregate Breeding, and diversity of endeavour with Segregate
Breeding is the cause of increased diversity of endowment. It
is very similar to the relation between power and exercise in the
individual. Without power there can be no exercise, and with-
out exercise there can be no continuance or growth of power.

We, therefore, see that the effects of Industrial Segregation
are specially liable to be enhanced by that form of Intensive
Segregation which I have suggested should be called Suetudinal
Intension.

Simple and familiar as the principles of Industrial Segregation
and Suetudinal Intension may seem, their consistent application
to the theory of evolution will throw new light on a wide range
of problems. This law of divergent evolution through Industrial
Segregation rests on facts that are so fully acknowledged by all
parties, that it seems to be a superfluous work to gather evidence
on the subject. It may, however, be profitable to consider briefly
whether the cases are frequent in which different habits of feed-
ing, of defence, or of nest-building become the cause of separate
breeding by which the same habits are maintained in one line of
descent without serious interruption for many generations. It
is important to remember, (1) that the separate breeding will
arise with equal certainty whether the diversity in the habits
has been initiated by original diversity in the instincts and
adaptations of the different variations, or by the crowding of
population inducing special efforts to find new resources, and
leading to diversity of endeavour ; and (2) that in either case the
result is what is here called Industrial Segregation. In the first
case the process is directly Segregative, while in the second case
it is primarily Separative, but (according to the principle dis-
cussed in the second section of last chapter) inevitably passes
into Segregate Breeding. Suetudinal Intension, or Divergent

226 REV. J. T. GULICK ON DIVERGENT EVOLUTION

Evolution through Diversity of Use, will operate as surely in the
one case as in the other.

1. Sustentational Segregation arises from the use of different
methods of obtaining sustentation by members of the same
species.

There can be no doubt that of the innumerable cases where
phytophagic varieties (as they are sometimes called) of insects
exist, a considerable proportion would be found on investigation
to be permanent varieties producing offspring that are better
adapted to the use of the special form of food consumed by the
parents than are the offspring of other varieties ; and it is evi-
dent that if the peculiar habits of each variety had no tendency
to produce segregative breeding this result would not be reached ;
for each variety would be promiscuously mingled with every
other, and, though the tendency to variation might be greatly
increased, the regular production of any one variety of young
would be prevented.

A large mass of facts could be easily gathered illustrative of
Sustentational Segregation ; but as the principle will probably
be denied by no one, we shall pass on without further expansion
of this part of the subject.

2. Protectional Segregation is Segregation from the use of
different methods of protection against adverse influences in the
environment.

When a new enemy enters the field occupied by any species
different methods of escape or defence are often open to the
members of the one species; and the use of these different
methods must sometimes result in the segregation of the mem-
bers according to the methods adopted. Some may hide in
thickets or holes, while others preserve themselves by flight.
Supposing the species to be an edible butterfly occupying the
open fields, and the new enemy to be an insectivorous bird also
keeping to the open country, certain members might escape by
taking to the wood-lands, while others might remain in their old
haunt, gaining through Protectional Selection more and more
likeness to some inedible species.

3. Nidificational Segregation.—Let us now consider the effects
of divergent habits in regard to nest-building. It is well known
to American ornithologists that the Cliff Swallow of the eastern
portions of the United States has for the most part ceased to
build nests in the cliffs that were the original haunts of the

THROUGH CUMULATIVE SEGREGATION. 997

species, and has availed itself of the protection from the weather
offered by the eaves of civilized houses; and that with this
change in nest-building has come a change in some of its other
habits. Now there is reason to believe that if the number of
houses had been limited to a hundredth part of those now exist-
ing, and if that limited number had been very slowly supplied,
this gradual change in some of the elements of the environment
would have resulted in divergent forms of adaptation to the en-
vironment in two sections of the same species. One section
would have retained the old habit of building in the cliffs, with
all the old adaptations to the circumstances that depend on that
habit ; while another section of the species would have availed
itself of the new opportunities for shelter under the eaves of
houses, and would have changed their inherited adaptations to
meet the new habits of nest-building and of feeding. It is also
evident that the prevention of free interbreeding between the
different sections caused by the diversity of habits would have been
an essential factor in the divergence of character in the sections.

Jt simply remains to consider whether the industrial habit
that separates an individual from the mass of the species will
necessarily leave it alone, without any chance of finding a consort
that may join in producing a new intergenerant. The answer is
that there is no such necessity. Though it may sometimes
happen that an individual may be separated from all companions
by its industrial habit, it is usually found that those that at one
time and one place adopt the habit are usually sufficient to keep
up the new strain, if they succeed in securing the needed sus-
tenance.

(6) CHronat SEGREGATION

is Segregation arising from the relations in which the organism
stands to times and seasons.

I distinguish two forms—Cyclical and Seasonal Segregation.

4. Cyclical Segregation is Segregation arising from the fact
that the life cycles of the different sections of the species do not
mature in the same years.

A fine illustration of this form of Segregation is found in the
case of Cicada septemdecim, whose metropolis is in Virginia,
Maryland, and Delaware, though many out-lying broods are
found in other regions east of the Mississippi River. The typical
form has a life-cycle of seventeen years, but there is a special

228 REV. J. T. GULICK ON DIVERGENT EVOLUTION

race (Cicada tredecim, Riley) that is separated from the typical
form, both locally and chronally. As the life-cycle of this race
is thirteen instead of seventeen years, even if occupying the same
districts and breeding at exactly the same season, interbreeding
could occur between the two forms only once in 221 years, or
once in 13 generations of the longer lived race, and once in 17
generations of the shorter lived race. During the year 1885 the
two races appeared simultaneously. The opportunity for testing
whether they would freely interbreed if brought together has,
therefore, passed not to return till the year 2106; but the dis-
tribution of the two races in different districts seems to indicate
that Local Segregation has had an important influence in the
development of the race. It is manifest, however, that if during
a period of local separation, or if during the period of 221 years
of Cyclical separation after the thirteen-year race was first
formed, this race should become modified in the season of its
appearing, there would after that be no mingling of race, though
brought together in the same districts. This would be Seasonal
Segregation, which we shall consider in the next section; but
what is of special interest here, as an example of complete
Cyclical Segregation, is the fact that at Fall River, Massachu-

setts, there is a brood of the septemdecim form, due a year later.

than the universal time of appearing.*

In any species where the breeding of each successive genera-
tion is separated by an exact measure of time which is very
rigidly regulated by the constitution of the species, Cyclical
Segregation will follow, if, through some extraordinary combina-
tion of circumstances, members sufficient to propagate the species
are either hastened or delayed in their development, and thus
thrown out of synchronal compatibility with the rest of the
species. If, after being retarded or hastened in development so
that part of a cycle is lost or gained, the old constitutional time
measure reasserts itself, the Segregation is complete.

So far as this one point relating to the time of maturing is
concerned the constitutional difference is segregative, while in

every other respect it will be simply separative, except as sepa- |

ration passes into Segregation. The Fall-River brood of Cicada

* See statement by Prof. C. V. Riley, in ‘ Science,’ vol. vi. p. 4. For par-
ticulars concerning the distribution and habits of this species see a paper by
Prof, Riley, read before the Biological Society of Washington, May 380, 1885,
extracts from which are given in ‘ Science,’ vol. v. p. 518.

‘
a, - = = m
a es el i ii

THROUGH CUMULATIVE SEGREGATION. 229

septemdecim being entirely separated from all other broods of the
same race by being belated a year, may be modified by forms of
Natural Selection that never arise in these other broods. And
this may be the case even if a brood observing the ordinary time
is always associated with it in locality.

5. Seasonal Segregation is produced whenever the season for
reproduction in any section of the species is such that it cannot
interbreed with other sections of the species. It needs no argu-
ment to show that if, in a species of plant that regularly flowers
in the Spring, there arises a variety that regularly flowers in the
autumn, it will be prevented from interbreeding with the typical
form. The question of chief interest is, under what circum-
stances are varieties of this kind likely to arise? Is a casual
sport of this kind likely to transmit to subsequent generations a
permanently changed constitution? If not, how is the new con-
stitution acquired? One obvious answer is that it may arise
under some special influence of the environment upon members
of the species that are geographically or locally segregated from
the rest of the species.

But may not the variation in the season of flowering be the
cause of segregation that will directly tend to produce greater
variation in that respect in the next generation, and so on till
the divergence in the constitutional adaptation to season is
carried to the greatest extreme that is compatible with the en-
vironment? I believe that it not only may, but must have that
effect; but we should remember that the average form which
flowers at the height of the season will so vastly predominate
over the extreme forms that the latter will be but stragglers in
comparison.

In regard to the one point of the season of readiness for pro-
pagation, this principle is segregative ; but in other respects it is
simply separative, unless through the principle of correlated
variation other characters are directly connected with the con-
stitution that determines the season. It will be observed that
Seasonal Segregation is produced by a parallel and simultaneous
change in the constitution of members in one place sufiicient to
propagate the species; while Cyclical Segregation is produced
by a simultaneous acceleration or retardation in the development
of members in one place sufficient to propagate the species with-
out disturbing the regular action of the constitution under
ordinary circumstances.

230 REV. J. T. GULICK ON DIVERGENT EVOLUTION

I distinguish two forms, viz. Geographical and Local Segre-
gation.

Geographical Segregation is Segregation that arises from the
distribution of the species in districts separated by geographical
barriers that prevent free interbreeding. Decided differences of
climate in neighbouring districts and regions may be classed as
geographical barriers.

Local Segregation is Segregation that arises when a species
with small powers of migration and small opportunities for trans-
portation has been, in time, very widely distributed over an area
that is not subdivided by geographical barriers. The Segrega-
tion in this case is due to the disproportion between the size of
the area occupied and the powers of communication existing
between the members of the species occupying the different parts
of the area. Though it is often difficult to say whether a given
case of Segregation should be classed as Geographical or Local,
still the distinction will be found useful; for the results will
differ according as the Segregation is chiefly due to barriers or
to wide diffusion of the species. In Geographical Segregation
the result is usually the development of well-defined varieties or
species on opposite sides of the barriers; but in Local Segrega-
tion it often happens that the forms found in any given locality
are connected with those in surrounding localities by individuals
presenting every shade of intermediate character ; and in general
terms it may be said that the forms most widely separated in
space are most widely divergent in character. It is of course
apparent that when the divergence has reached a certain point
the differentiated forms may occupy the same districts without
interbreeding, for they will be kept apart by some, if not all, of
the different forms of Industrial, Chronal, Conjunctional, and
Impregnational Segregation.

Three different forms of Spatial Segregation may be distin-
guished according to the causes by which they are produced, viz.:—

6. Migrational Segregation, caused by powers of locomotion
in the organism.

7. Transportational Segregation, caused by activities in the
environment that distribute the organism in different districts,
(prominent among these are currents of atmosphere and of

THROUGH CUMULATIVE SEGREGATION. 231

water, and the action of migratory species upon those that can
simply cling).

8. Geological Segregation, caused by geological changes di-
viding the territory occupied by a species into two or more
sections. For example, geological subsidence may divide the
continuous area occupied by a species into several islands,
separated by channels which the creatures in question cannot
pass.

Migration differs from transportation simply in that the former
is the direct result of activities in the organism, and the latter
of activities in the environment; and though the distribution of
every species depends on the combined action of both classes of
activities, it is usually easy to determine to which class the
carrying power belongs. The qualities of the thistle-down
enable it to float in the air, but it is the wind that carries it afar.

Some degree of Local Segregation exists whenever the members
of a species produced in a given area are more likely to interbreed
with each other than with those produced in surrounding areas,
or whenever extraordinary dispersal plants a colony beyond the
range of ordinary dispersal. In other words, when those pro-
duced in a given district are more nearly related with each other
than with those produced in surrounding districts, there local
segregation has existed.

There is one important respect in which Spatial Segregation
differs from all other forms of Environal Segregation, namely, in
its ordinary operation it does not depend directly upon diversity
in the qualities and powers of the organism. ‘The dispersion of
the members of a species would not be prevented if each was
exactly like every other ; though, of course, if there were no
power of variation, separate breeding would have no influence in
producing divergence of character. It follows that every species
is, or is more or less liable to be, affected by Spatial Segregation ;
and it often happens that other forms of Segregation arise
through the previous operation of this form; but as Spatial
Segregation prevents organisms from crossing only when sepa-
rated in space, it must always be reinforced by other forms
of segregation before well-defined species are produced that
are capable of occupying the same district without interbreeding.
The vast majority of the divergent forms arising through Local
Segregation are reintegrated with the surrounding forms, new
divergences constantly coming in to take the place of the old ;

232 REV. J. T. GULICK ON DIVERGENT EVOLUTION

but if, during its brief period of local divergence, Industrial
or Chronal Segregation is introduced, the variety becomes
more and more differentiated, and, as one after another the
different forms of Reflexive Segregation arise, it passes into a
well-defined species. There is, however, reason to believe that
the order of events is often the reverse, Reflexive forms of
Segregation being the cause of the first divergences.

As Spatial Segregation does not depend upon diversity in the
qualities and powers of the organism, so also it does not usually
result in distributing the organism in different localities according
to their differences of endowment. The causes that produce it
are primarily separative, not segregative.

Migration is produced by the natural powers of the organism,
acting under the guidance of instincts that usually lead a group
of individuals, capable of propagating the species, to migrate
together; while the organisms that are most dependent on
activities in the environment for their distribution, are usually
distributed in the form of seeds or germs, any one of which is
capable of developing into a complete community.

The causes of Separation between the different sections, and
of Integration between the members of one section, are therefore
sufficiently clear ; but what are the causes of difference of cha-
racter in the different sections, especially when they are exposed
to the same environment? These causes all come under what I
call Intensive Segregation, which, for the sake of saving repetition ,
will be fully discussed in a separate paper.

(2d) 9. FerrrnizaTIONaL SEGREGATION.

Since writing this chapter on Environal Segregation, I have
seen Francis Galton’s short article on “ The Origin of Varieties ”
published in ‘ Nature,’ vol. xxxiv. p. 395, in which he refers to a
cause of segregation that had not occurred tome. He says :—“ If
insects visited promiscuously the flowers of a variety and those
of the parent stock, then—supposing the organs of repro-
duction and the period of flowering to be alike in both, and
that hybrids between them could be produced by artificial cross-
fertilization—we should expect to find hybrids in abundance
whenever members of the variety and those of the original stock
occupied the same or closely contiguous districts. It is hard to
account for our not doing so, except on the supposition that insects
feel repugnance to visiting the plants interchangeably.”

Se ee

THROUGH CUMULATIVE SEGREGATION. 233

Following the form of nomenclature adopted in this paper, I
venture to call this principle Fertilizational Segregation.

It is evident that Segregation of this form depends on diver-
gence of character already clearly established, and therefore on
some other form of Segregation that has preceded. It is also
segregative rather than separative, in that it perpetuates a segre-
gation previously produced, which might otherwise be obliterated
by the distribution of the different forms in the same district.
The form of Segregation that precedes Fertilizational Segregation,
producing the conditions on which it depends, must, from the
nature of the case, be Local Segregation. Chronal and Impreg-
national Segregation, when imperfectly established, might be
fortified by Fertilizational Segregation ; but, in the case of plants,
these are all dependent on previous Local Segregation.

(e) ARTIFICIAL SEGREGATION.

Artificial Segregation is Segregation arising from the relations
in which the organism stands to the rational environment. As
the operation of this cause is familiar, and as it was considered
in the last chapter when discussing the effects of segregation, we
pass on, simply calling attention to the fact that it is a form of
Environal Segregation.

Tue ImwporTANCE OF ENVIRONAL SEGREGATION.

‘We must not assume that the various forms of Environal
Segregation are of small influence in the formation of species
because Sexual or Impregnational Incompatibility is a more
essential feature, without which all other distinctions are liable
to be swept away. The importance of the forms of segregation
discussed in this chapter lies in the fact that they often open
the way for the entrance of the more fundamental forms of
segregation, even if they are not essential conditions for the
development of the same. Though myriads of divergent forms
produced by Local and Industrial Segregations are swept away
in the struggle for existence, and myriads are absorbed in the vast
tides of crossing and intercrossing currents of life, the power of
any species to produce more and more highly adapted variations,
and to segregate them in groups that become specially adapted to
special ends, or that grow into specific forms of beauty and
internal harmony, is largely dependent on these factors.

234 REV. J. T. GULICK ON DIVERGENT EVOLUTION

CHAPTER IV.

DESCRIPTION AND CLASSIFICATION OF THE CAUSES OF
CUMULATIVE SEGREGATION (continued).

B. REFLEXIVE SEGREGATION.

Reflexive Segregation is Segregation arising from the relations
in which the members of one species stand to each other.

It includes three classes, which I call Conjunctional, Impreg-
national, and Institutional Segregation.

It is important to observe that Intergeneration requires com-
patibility in all the circle of relations in which the organism
stands ; but, in order to ensure Segeneration between any two
or more sections of a species, it is sufficient that incompati-
bility should exist at but one point. If either sexual or social
instincts do not accord, if structural or dimensional characters
are not correlated, if the sexual elements are not mutually
potential, or if fixed institutions hold groups apart, Intergen-
eration is prevented, and Segeneration is the result, either as
Segregation, or as Separation that is gradually transformed
into Segregation.

(a) CoNJUNCTIONAL SEGREGATION.

Conjunctional Segregation is Segregation arising from the
instincts by which organisms seek each other and hold together
in more or less compact communities, or from the powers of
growth and segmentation in connection with self-fertilization,
through which similar results are gained.

I distinguish four forms—Social, Sexual, Germinal, and Floral
Segregation.

10. Social Segregation is produced by the discriminative action
of social instincts.

The law of social instinct is preference for that which is
familiar in one’s companions ; and, as in most cases the greatest
familiarity is gained with those that are near of kin, it tends to
produce breeding within the clan, which is a form of Segregate
Breeding. If the clan never grows beyond the powers of
individual recognition, or if the numbers never become so great
as to impede each other in gaining sustenance, there will be but:
little occasion for segregation ; but multiplication will lead to
segmentation. Wherever the members of a species, ranging freely

THROUGH CUMULATIVE SEGREGATION: 235

over a given area, divide up into separate herds, flocks, or swarms,
of which the members produced in any one clan breed with each
other more than with others, there we have Social Segregation.

It should always be kept in mind that Social Segregation
arises at avery early stage, holding apart groups not at all or
but very slightly differentiated; while in the case of many
animals, the eager sexual instincts of the males constantly tend
to break up these minor groups. Though the barriers raised by
social insincts are often broken over, their influence is not
wholly overcome; and in many intstances the Social Segregation
becomes more and more pronounced, till in time decided Sexual
Segregation comes in to secure and strengthen the divergence.

11. Sexual Segregation is produced by the discriminative action
of sexual instincts.

There can be no doubt that sexual instincts often differ in such
a way as to produce segregation. But how shall we account for
these differences? In the case of Social Segregation there is no
difficulty, for it seems to be, like migration, due to a constant
instinct, always tending to segregation. We also see that an
endowment which prevents the destruction of the species through
the complete isolation of individuals, and which co-operates with
migrational instincts in securing dispersal without extinction,
may be perfected by the accumulating effects of its own action.
And is there any greater difficulty in accounting for the law that
regulates sexual instincts ? Ifit can be shown that Vigour and
Variation, the conditions on which adaptation depends, are in
their turn dependent on some degree of crossing, there will be no
difficulty in attributing the development of an instinct that
secures the crossing to the superior success of the individuals
that possess it in even a small degree. On the other hand,
whenever there arises a variety that can maintain itself by crosses
within the same variety, any variation of instinct that tends to
segregation will be preserved by the segregation. It needs no
experiments to prove that, if the members of a species are im-
pelled to consort only with the members of other species, they
will either fail to leave offspring, or their offspring will fail to
inherit the characteristics of the species. The same is true con-
cerning the continuance of a variety that is not otherwise segre-
gated. The power of variation on the one hand, and the power
of divergent accumulation of variations on the other hand, are
prime necessities for creatures that are wresting a living from a

236 REV. J. T. GULICK ON DIVERGENT EVOLUTION

vast and complex environment ; and the former is secured by the
advantage over rivals possessed by the variations that favour
crossing, and the latter by the better escape from the swamping
effect, and sometimes from the competition of certain rivals,
secured by the more segregative variations. We must there-
fore believe that, whenever in the history of an organism there
arise segregative variations which are able to secure sufficient
sustentation and propagation to continue the species, the segre-
gative quality of the forms thus endowed will be preserved and
accumulated through the self-accumulating effect of the segre-
gative endowments.

It is probable that in many of the higher vertebrates sexual
instincts tend to bring together those of somewhat divergent
character, but the difference preferred is within very narrow
limits ; and beyond those limits, it may be said that the general
law for sexual attraction is, that it varies inversely as the dif-
ference in the characters of the races represented, if not inversely
as some power of such difference. The action of such a law
is necessarily segregative, whenever the divergence has, through
other causes, passed beyond the limit of higher attraction. Before
Sexual Segregation can arise, there must arise distinctive charac-
teristics by means of which the members of any section may
discriminate between those of their own and other sections. If
there are no constant characteristics, there can be no. constant
aversion between members of different groups, no constant pre-
ference of those of one’s own group. From this it follows, that
before Sexual Segregation can arise, some form of Segregation
that is not dependent on accumulated divergence of character
must have produced the divergence on which the Sexual Segre-
gation depends. Such forms are Local, Social, and some kinds of
Industrial Segregation. When varieties have arisen through
these causes, it often happens that Sexual Segregation comes in
and perpetuates the Segregation which the initial causes can no
longer sustain. As long as the groups are held apart by diver-
gent sexual instincts, it is evident that divergent forms of Sexual
Selection are almost sure to arise, leading to a further accumu-
lation of the divergence initiated by the previous causes.

If there is any persistent cause by which local and social
groups are broken up and promiscuously intermingled before
recognizable characters are gained, the entrance of Sexual Segre-
gation will be prevented. I therefore conclude that the chief

EE SEeEeEE—————_—

THROUGH CUMULATIVE SEGREGATION. 237

influence of this latter factor is found in its prolonging and
fortifying the separate breeding of varieties that have arisen
under Local, Social, or Industrial Segregation, and in thus con-
tinuing the necessary condition for the development of increas-
ingly divergent forms of Intensive Segregation, under which the
organism passes by the laws of its own vital activity when dealing
with a complex environment in groups that never cross.

12. Germinal Segregation is caused by the propagation of the
species by means of seeds or germs any one of which, when
developed, forms a community so related that the members breed
with each other more frequently than with the members of other
communities. If the constitution of any species is such that the
- ovules produced from one seed are mbre likely to be reached and
fertilized by pollen produced from the same seed than by pollen
produced from any other one seed, then Germinal Segregation is
the result.

In order to secure this kind of Segregation it is not necessary
that the flowers fertilized by pollen from the same plant should
be more fertile, or the seeds capable of producing more vigorous’
plants than the flowers fertilized by pollen from another plant.
All that is required is that of the seeds produced a larger number
shall be fertilized by the pollen of the same plant than by the
pollen of any other one plant.

This form of Segregation is closely related to Local Segre-
gation on one side, and to Social Segregation on the other. It,
however, differs from the former in that it does not depend on
Migration or Transportation, and from the latter in that it does
not depend on social instincts.

13. Floral Segregation is Segregation arising from the closest
form of self-fertilization, namely the fertilization of the ovules of
a flower by pollen from the same flower.

Many plants that in their native haunts are frequently crossed
by the visits of insects depend entirely on self-fertilization when
transported to other countries where no insect is found to per-
form the same service for them. The common pea (Pisum
sativum) is an example of a species that habitually fertilizes
itself in England, though Darwin found that it was very rarely
visited by insects that were capable of carrying the pollen.*
Darwin also mentions Ophrys apifera as an orchid which “has

* See ‘Cross- and Self-Fertilization in the Vegetable Kingdom,’ p. 161.
LINN. JOURN.—ZOOLOGY, VOL. Xx. 19

238 REV. J. fT. GULTCK ON DIVERGENT EVOLUTION

almost certainly been propagated in a state of nature for thousands
of generations without having been once intercrossed.”’ *

General Observations on Germinal and Floral Segregation.

A fact of great importance in its bearing on the origin of
varieties should be here noted. Any variation, arising as a so-
called sport, in any group of plants where either of these prin-
caples is acting strongly will be restrained from crossing, and will
be preserved except in so far as reversion takes place. Now
there is always a possibility that some of the segregating branches
of descent will not revert, and that, through the special character
which they possess in common, they will some time secure the
services of some insect that will give them the benefit of cross-
fertilization with each other without crossing with other varieties.
The power of attaining new adaptations may be favoured by self-
fertilization occasionally interrupted by interbreeding with indi-
viduals of another stock ; for the latter is favourable as intro-
ducing vigour and variation, and the former as giving opportunity
for the accumulation of variations.

(6) IMPREGNATIONAL SEGREGATION.

Impregnational Segregation is due to the different relations in
which the members of a species stand to each other in regard to
the possibility of their producing fertile offsprmg when they
consort together.

In order that Impregnational Segregation should be established
and perpetuated it is necessary, 1st, that variation should arise
from which it results that those of one kind are capable of pro-
ducing vigorous and fertile offspring in greater numbers when
breeding with each other than when breeding with other kinds ;
2nd, that mutually, compatible forms should be so brought to-
gether as to ensurg propagation through a series of generations.
In order to secu:/e this second condition, it is necessary that, in
the case of piants, there should be some degree of Local, Germinal,
or Floral Segregation, and, in the case of animals that pair,
either pronounced, Local Segregation, or partial Local Segre-
gation supplemented by Social or Sexual Segregation. The first
of these factors I call Negative Segregation, as contrasted with all

* See ‘Cross- and Self- Fertilization in the Vegetable Kingdom,’ p. 489.

THROUGH CUMULATIVE SEGREGATION. 239

other forms of Segregation, which I group together as Positive
Segregation.

Of each form of Segregation which we have up to this point
considered, the segregating cause has been one that distributes
individuals of the same species in groups between which free
intergeneration is checked ; while the propagation of the different
groups depends simply on the original capacity for intergenera-
ting common to all the members of the species. The intercrossing
has been limited not by the capacity, but by the opportunity and
inclination of the members. Coming now to cases in which the
lack of capacity is the cause that checks the production of
mongrels, we find a dependence of a very different kind; for to
ensure the propagation of the different groups it is not enough
that the general opportunity for the members to meet and con-
sort remains unimpaired. There must be some additional segre-
gating influence bringing the members together in groups corre-
sponding to their segregate capacity, or they will fail of being
propagated.

A partial exception must be made in the case of Potential
and Prepotential Segregation, the latter being due to the pre-
potency of the pollen of a species or variety on the stigma of the
same species or variety, and the former to the complete impo-
tence of the foreign pollen. When allied species of plants are
promiscuously distributed over the same districts, and flowering
at the same time, prepotency of this kind is one of the most
direct and efficient causes of Segregate Breeding. The same
must be true of varieties similarly distributed whenever this
character begins to affect them. In the case, however, of
dicecious plants and of plants whose ovules are incapable of being
impregnated by pollen from the same plant, no single plant can
propagate the species. If, therefore, the individuals so varying
as to be prepotent with each other are very few and are evenly
distributed amongst a vast number of the original form, they
will fail of being segregated through failing to receive any of the
prepotent pollen. It is thus apparent that when the mutually
prepotent form is represented by comparatively few individuals,
their propagation without crossing will depend on their being
self-fertile and subject to Germinal or Floral Segregation, or on
their being brought together by some other form of Positive
Segregation.

When a considerable number of species of plants are commingled

19*

240 REV. J. T. GULICK ON DIVERGENT EVOLUTION

and are flowering at the same time, their separate propagation is
preserved, in no small degree, by the Prepotential Segregation of
those that are most nearly allied, and by the complete Potential
Segregation of those that belong to different families, orders, and
classes. The same principle must come in to prevent the crossing
of different species, genera, families, and orders of animals whose
fertilizing elements are distributed in the water. We must,
therefore, consider it a form of Positive as well as Negative
Segregation ; for the free distribution of the fertilizing element,
with the superior affinity of the two sexual elements when pro-
duced by those that are mutually prepotential, secures the inter-
breeding of those that are mutually prepotential.

Impregnational Segregation generally exists between the dif-
ferent species of the same genus, almost always between species
of different genera, and always between species of different
families, orders, classes, and all groups of higher grade. And in
all these cases it is associated with other forms of segregation,
and whenever it has once become complete, it has never been
known to give way. Though complete mutual sterility never
gives place to complete mutual fertility, in every case where the
descendants of the same stock have developed into different
classes or orders, and in most cases where they have developed
into different families or genera, the reverse process has taken
place, and complete mutual fertility has given place to complete
mutual sterility.

Under Impregnational Segregation I distinguish five principles:
namely, Segregate Size, Segregate Structure, Potential and Pre-
potential Segregation, Segregate Fecundity, and Segregate Vigour.

14. Segregate Size is caused by incompatibility in size or
dimensions.

As familiar illustrations of this form of Segregation, I may
mention the following :—The largest and smallest varieties of the
ass may run in the same pasture without any chance of crossing.
I have also kept Japanese bantam fowls in the same yard with
other breeds without any crossing. In many other species indi-
viduals of extreme divergence in size are incapable of inter-
breeding. .

15. Segregate Structure is caused by the lack of correlation in
the proportionate size of different organs and by other incom-
patibilities of structure.

Darwin suggests that the impossibility of a cross between

THROUGH CUMULATIVE SEGREGATION. 241

certain species may be due to a lack of correspondence in
length of the pollen-tubes and pistils. Such a lack of harmony
would perhaps account for difference of fertility in reciprocal
crosses.

Segregate Structure does not usually arise till other forms of
Segregation have become so well established that difference of
structure does not make any essential difference in the amount of
intergeneration. It is not, however, impossible that species that,
would otherwise be fertile inter se are thus held apart. In
Broca’s work on ‘Human Hybridity’* there is a passage
quoted from Prof. Serres, showing that it is very possible that
this form of incompatibility may exist between certain races
of man.

16. Potential Segregation and Prepotential Segregation.—These
are caused by more or less free distribution of the fertilizing
element together with the greater rapidity and power with which
the sexual elements of the same species, race, or individual com-
bine, as contrasted with the rapidity and power with which the
elements of different species, races, or individuals combine.
Potential Segregation is caused by the mutual impotence of the
contrasted forms, as is always the case between different orders
and classes; and Prepotential Segregation is caused by the
superior influence of the fertilizing element from the same
species, race, or individual, as contrasted with that from any
other species, race, or individual, when both are applied to the
same female at the same time, or sometimes when the prepotent
element is applied many hours after the other.

For the operation of this principle the fertilizing element from
different males must be brought to the same female.

When pollen from a contrasted genus, order, or class has no
more effect than inorganic dust, it seems appropriate that we
should call the result Potential Segregation rather than Prepo-
tential Segregation, which implies that the foreign as well as the
home pollen is capable of producing impregnation. Prepotential
Segregation may be considered the initial form of Potentrl
Segregation, the former passing through innumerable grades of
intensity into the latter. We may, therefore, consider the
principles as fundamentally one, though it will be convenient to
retain both names.

* English translation published by the Anthropological Society of London,
p. 28.

242 REV. J. T. GULICK ON DIVERGENT EVOLUTION

The importance of this principle in producing and preserving
the diversities of the vegetable kingdom can hardly be over-
stated. If pollen of every kind were equally potent on every
stigma, what would the result be? What distinctions would
remain? And if Potential Segregation is necessary for the
preservation of distinctions,.is it not equally necessary for their
production? Amongst water-animals that do not pair, the
same principle of Segregation is probably of equal importance.
Concerning this form of Segregation many questions of great
interest suggest themselves, answers to which are not found in
any investigations with which I am acquainted. Some of these
questions are as follows :—

(1) Arethere many cases of Prepotential as wellas of Potential
Segregation between different forms of water-animals ?

(2) Is Prepotential Segregation always accompanied by Segre-
gate Fecundity and Segregate Vigour ?

(3) If not always associated, which of the three principles
first appears? And what are their relations to each other?

(4) When allied organisms are separated by complete Environal
Segregation, are they less liable to be separated by these three
principles ?

Darwin has in several places referred to the influence of pre-
potency in pollen, and in two places I have found reference to
the form of prepotency that produces segregation ; but I find no
intimation that he regarded this or any other form of segregation
as a cause of divergent evolution, or as a necessary condition for
the operation of causes producing divergent evolution. The
effect of prepotency in pollen from another plant in preventing
self-fertilization is considered in the tenth chapter of his work
on ‘Cross- and Self-Fertilization in the Vegetable Kingdom,’
pp. 891-400. Some very remarkable observations concerning
the prepotency of pollen from another variety from that in which
the stigma grows are recorded in the same chapter; but no
reference is there made to the effect that must be produced when
the pollen of each variety is prepotent on the stigma of the
game variety. In the sixteenth chapter of ‘ Variation under
Domestication,’ it is suggested that prepotency of this kind
might be a cause of different varieties of double hollyhock repro-
ducing themselves truly when growing in one bed; though
there was another cause to which the freedom from crossing in
this case had been attributed. Again, in chapter viil. of the

THROUGH CUMULATIVE SEGREGATION. 243

fifth edition * of ‘The Origin of Species,’ in the section on “ The
Origin and Causes of Sterility,’ Darwin, while maintaining that.
the mutual sterility of species is not due to Natural Selection,
refers to prepotency of the kind we are now considering as a
quality which, occurring in ever so slight adegree, would prevent
deterioration of character, and which would therefore be an ad-
vantage to a species in the process of formation, and accordingly
subject to accumulation through Natural Selection. In order to
construct a possible theory for the introduction of sterility
between allied species by means of Natural Selection, he finds it
necessary simply to add the supposition that sterility is directly
caused by this prepotency. He, however, for several reasons
concludes that there is no such dependence of mutual sterility on
the process of Natural Selection. Concerning the prepotency
he makes no reservation, and I accordingly judge that he con-
tinued to regard it as strengthened and developed through the
action of Natural Selection.

It is concerning this last point that I wish to give reasons for
a different opinion. I believe that qualities simply producing
Segregation can never be accumulated by Natural Selection ;
for :—

(1) When separate generation comes in between two sections
of a species they cease to be one aggregate, subject to modifi-
cation through the elimination of certain parts. Both will be
subject to similar forms of natural selection only so long as the
circumstances of both and the variations of both are nearly the
same, but they will no longer be the members of one body
between which the selecting process is carried out. On the con-
trary, if they occupy the same district each group will stand in
the relation of environment to the other, modifying it, and being
modified by it, without mutually sharing in the same modifi-
cation.

(2) Though one may exterminate the other, the change that
comes to the successful group through the contest is not due to
its superiority over the other, but to the superiority cf some of
its own members over others.

(8) When any Segregate form begins to arise we cannot attri-
bute its success to the advantage of segeneration, tor the inter-

* Since my comments on this passage were written, I have discovered that
Darwin has omitted it from the sixth edition.

244 REV. J. T. GULICK ON DIVERGENT EVOLUTION

generating forms are at the same time equally successful; where-
fore it is not the success, but the separateness of the success,
that is due to the segeneration.

(4) The continuance of the descendants of a group in a special
form will depend on its Segregation ; but this is a very different
thing from the special success of its descendants. The preser-
vation of a special kind of adaptation is never due to natural
selection, which is the superior suecess of the higher degrees of
adaptation of every kind.

(5) The power of migration, or any other power directly related
to the environment, may be accumulated by natural selection,
and afterward lead to Segregation ; but, according to my method
of judging, the continuous advantage of Segregation over Inte-
gration can never be shown, for both are equally essential in the
economy of nature; and though one process may at one time
predominate over the other, the comparative advantage of Segre-
gation, if there be such advantage, cannot be the cause of the
preservation of forms endowed with segregative qualities, for
they will certainly be preserved as long as they are able to win &
bare existence, which is often a lower grade of success than the
one from which they are passing.

(6) According to my view, instead of the accumulation of the
Segregative prepotency depending on natural selection, the accu-
mulation of divergent forms of natural selection depends on some
form of Segregation.

But if the accumulation of Prepotential Segregation is not due
to Natural Selection, how shall we explain it? It is, I think,
due to the fact that those forms that have the most of this
character are, through its action, caused to breed together. We
have already seen, when considering Seasonal and Sexual Segre-
gation, that, if Segregation is directly produced by the instincts
or physiological constitution of the organism, there is a tendency
toward an increasing manifestation of the character in successive
generations. Those that have but a slight degree of Segregate
prepotency eventually coalesce, forming one race, while those
possessing the same character in a higher degree remain more
distinct, and their descendants become still more segregate and
still more permanently divergent. As long as the segregate
forms are able to maintain vigour and secure fair sustentation,
the process continues and the separation becomes more pro-
nounced. Of this form of the Law of Cumulative Segregation

THROUGH CUMULATIVE SEGREGATION. 24.5

we may say that, as the descendants of the best fitted necessarily
generate with each other and produce those still better fitted, so
the descendants of those possessing the most segregative endow-
ments necessarily generate with each other and produce those
that are still more segregate.

It may at first appear that a slight degree of prepotence will
prevent crossing as effectually as a higher degree; but further
reflection will show that the efficiency of the prevention will vary
in direct proportion with the length of time over which the pre-
potent pollen is able to show its prepotence, and this will allow
of innumerable grades. If, in the case of certain individuals,
the prepotency is measured by about twenty minutes, while with
other individuals it enables the pollen of the same variety to
prevail, though reaching the stigma an hour after the pollen of
another variety has been applied, the difference in the degree of
Segregation will be sufficient to make the persistence of the
Jatter much more probable than that of the former. This form
of Segregation is evidently one of the important causes prevent-
ing the free crossing of different species of plants. It probably
has but little influence on terrestrial animals; but how far it is
the cause of Segregation among aquatic animals is a question of
no small interest, concerning which I have but small means for
judging. I have, however, no hesitation in predicting that, unless
we make the presence of this Segregative quality the occasion for
insisting that the forms so affected belong to different species,
we shall find that amongst plants the varieties of the same species
are often more or less separated from each other in this way. I
do not know of any experiments that have been directed toward
the determining of this point ; but on the general principle that
physiological evolution is not usually abrupt, and that race
distinctions are the initial forms under which specific differences
present themselves, I can have no doubt that feeble prepotence
precedes that which is more pronounced, and that part of this
divergence in many cases takes place, while the divergent branches
may be properly classed as varieties. Another reason for believ-
ing that Prepotential Segregation will be found on further inves-
tigation to exist in some cases between varieties, is the constancy
with Which, in the case of species, this character is associated
with Segregate Fecundity and Segregate Vigour, which we know
are sometimes characteristics of varieties in their relation to each
other. The importance of these latter principles when occurring

246 REV. J. T. GULICK ON DIVERGENT EVOLUTION

in connection with different forms of partial Segregation will
now be considered.

17,18. Segregate Fecundity and Segregate Vigour.—By Segre-
gate Fecundity I mean neither Segregation produced by Fecun-
dity nor Fecundity produced by Segregation, but the relation in
which species or varieties stand to each other when the intergen-
eration of members of the same species or variety results in
higher fertility than the crossing of different species or varieties.
In like manner Segregate Vigour is the relation in which species
or varieties stand to each other when the intergeneration of
members of the same species or variety produces offspring more
vigorous than those produced by crossing with other species or
varieties. Integrate Fecundity and Integrate Vigour are the
terms by which I indicate the relation to each other of forms in
which the highest fertility and vigour are produced by crossing,
and not by independent generation.

Before discussing these principles through which the influence
of Segregation is greatly increased, it will be an advantage if we
can gain some idea of the nature of Cumulative Fertility in its
relations to a law of still wider import. I refer to the fourfold law
of antagonistic increase and mutual limitation between (1) In-
tegration, (2) Segregation, (3) Adaptation, (4) Maultiplication—
in other words between (1) General invigoration and power of
variation through crossing, (2) The opening of new opportunities
and independent possibilities, (3) Special adaptation to present
circumstances, (4) Powers of multiplied individualization. Darwin
has considered at length the 1st and the 8rd, though I do not
remember that he has anywhere pointed out that their develop-
ment is due to a kind of self-augmentation. I believe this is so
emphatically the case that the former might well be called the
law of Self-Cumulative Vigour, and the latter the law of Self-
Cumulative Adaptation. Corresponding to these two laws, I
find the additional laws of Self-Cumulative Segregation and Self-
Cumulative Fertility. Darwin’s theory, that Diversity of Natural
Selection is directly and necessarily dependent on exposure to
different external conditions, tends to obscure, though not to
deny, the fact that the breeding together of the better adapted,
which causes the increase of adaptation, is due to the different
degrees of endowment in the organism, rather than to diversity
in the environment. It is also true of segregative endowment
and of fertility that they are necessarily cumulative whenever

THROUGH CUMULATIVE SEGREGATION. 247

they belong in different degrees to members of the same Inter-
generant that are equally fitted. The cumulation of vigour, as
that of adaptation, is, I think, rightly classed as a form of Selec-
tion; for in both cases it depends on the power of the more
highly endowed to supplant the less endowed without allowing
them full opportunity to propagate; but the increase of segre-
gative endowments and of fertility is due to principles quite
different from this, and differing from each other. The segre-
gative endowments augment through the inherent tendency of
the more highly endowed to breed more exclusively with those of
the same form, and therefore in the long run to breed more
exclusively with each other ; while the fertility of the more fer-
tile neither drives out the less fertile nor holds the two classes
apart, but simply multiplies the offspring of the more fertile,
making it sure that in each generation they will predominate.
But all these forms of augmentation correspond in that they
secure the breeding together of those possessing higher degrees
of the special endowment, and so increase the average endow-
ment, either of the whole number of the offspring, or of the
segregated portion. Vigour increases through the breeding to-
gether of the more vigorous, resulting from their overcoming and
crowding out the less vigorous without allowing them full oppor-
tunity to propagate. Adaptation increases through the breeding
together of the better adapted, resulting from their supplanting
their rivals without allowing them full opportunity to propagate.
Segregative endowments increase through the breeding together
of the more highly endowed, resulting from the fact that as long
as Segregation is incomplete more than half of each generation of
pure descent are necessarily the offspring of parents whose segre-
gative endowments were above the average. Fertility increases
through the breeding together of the more fertile, resulting from
the fact that more than half of each generation are the offspring
of parents of more than average fertility. As the breeding to-
gether of the more vigorous and the better adapted, caused by their
superior success, tends to increase and intensify the vigour and
adaptation of successive generations, so the breeding together of
those more highly endowed with Segregative powers, caused by
the Segregation, tends to strengthen and intensify the Segregative
powers in successive generations ; and so the breeding together of
the more fertile, caused by the larger proportion of offspring
produced by the more fertile, tends to increase the fertility of

248 REV. J. T. GULICK ON DIVERGENT EVOLUTION

successive generations. Among those that would be equally pro-
ductive if equally nourished, the ratio of propagation varies
directly as the degree of sustentation above a certain minimum
(and perhaps below a certain maximum), and therefore directly
as the degree of adaptation that secures this sustentation. This
propagation according to degrees of adaptation to the environment
is what I understand by natural selection. But among those
that are equally adapted to the environment the ratio of propa-
gation varies directly as the ratio of fertility. This propagation
according to degrees of fertility 1s what I call the Law of Cumu-
lative Fertility. It is not due to different degrees of success, or
to any advantage which the individuals of one form have over
those of other forms; but simply to the higher ratio of multipli-
cation in the more fertile forms securing the intergeneration of
the more fertile. Jn connection with natural selection it ensures,
in the descendants, the predominance of the better adapted of the
more fertile, and the more fertile of the better adapted.

At the close of the previous chapter I called attention to the
fact that innumerable Local Segregations and other imperfect
forms of Segeneration are being constantly broken down, partly
by the increase of numbers and partly by the superior fertility
and vigour of offspring produced by crossing. It seems to bea
fundamental law that vigour and variation in the offspring depend
on some degree of diversity of constitution in the parents, and
diversity of constitution that is not entirely fluctuating depends
on some degree of Positive Segregation; therefore vigour and
variation depend on the breaking-down of incipient Segrega-
tions, and on the interfusion of the slightly divergent forms
that had been partially segregated. But in the history of
every race that is winning success by its vigour and varia-
tion there is liable to come a time when some variety, inher-
iting sufficient vigour to sustain itself, even if limited to the
benefits of crossing with the individuals of the same variety,
becomes partially Segregated. As we have already seen, Segre-
gation, in so far as it depends on the qualities of the organism,
tends ever to become more and more intense; but, in the very
nature of things, not only will the Segregation be for many
generations only partial, but partial Segregation, though it may
greatly delay the submerging of different groups in one common
group, will never prevent that result being finally reached.
Though the siphon that connects two tanks of water be ever so

OE a -

THROUGH CUMULATIVE SEGREGATION. 249

small, the water will in time find a common level in both tanks,
unless there are additions or subtractions of water that prevent
such a result. So, in the case under consideration, final fusion
will take place, unless differentiation progresses more rapidly than
the fusion, or some other influence comes in to counteract the
levelling influence of occasional crosses. If, under such condi-
tions, some branch of the partially Segregated variety becomes
more fertile when generating with members of the same variety,
and Jess fertile when generating with other varieties, a principle
will be introduced tending to strengthen any form of partial
Segregation that already exists between the varieties. This
principle when co-operating with partial Segregation will produce
pure masses of each variety, when, without the action of this
principle, all distinctions would be absorbed by the crossing.
We know that a transition from Integrate Fecundity to Segre-
gate Fecundity usually takes place at a point in the history of
evolution intermediate between the formation of an incipient
variety and a strongly-marked species; and though the causes
that produce this transition may be very difficult to trace, I
believe the results that must follow can be pointed out with
considerable clearness and certainty.

Darwin’s investigations have shown that in many cases, if not
in the majority, the relation of varieties to each other is that
which I have called Integrate Fecundity and Integrate Vigour;
that is, the highest fertility is attained when varieties are crossed,
and the vigour of offspring thus produced is greater than when
the intergeneration is within the limits of one variety. He,
however, gives in ‘ Variation under Domestication,’ chapter xvi.,
some special cases, in which “ varieties of the same species behave,
when crossed, like closely allied but distinct species”; and re-
marks that similar cases “may not be of very rare occurrence ;
for the subject has not been attended to.” The same cases are
also mentioned in all the editions of the ‘ Origin of Species.’ *

The problems that arise in considering the different results
produced by different degrees of Positive Segregation and Segre-
gate Fecundity are of a nature.suitable for mathematical treat-
ment. Before, however, computing the effects of Segregate
Fecundity when co-operating with Positive Segregation, it will be
in place to show that it is of itself only a negative form of

* See Ist edition, p. 288; 5th edition, p. 259; 6th edition, p. 258.

250 REV. J. T. GULICK ON DIVERGENT EVOLUTION

Segregation, having no power to ensure the propagation of the
varieties thus characterized, though they are fully adapted to
the environment. This is most easily brought to light by con-
sidering the effect of a high degree of this quality when Posi-
tive Segregation is entirely wanting, or when it is sufficient to
give simply a chance of Segregate Breeding by bringing each
individual near to its natural mate. For example, let us suppose,
1st, that a male and a female each of several allied but mutually
sterile species are brought together on one small island, all other
tendences to Positive Segregation being removed, while mutual
sterility still remains; 2nd, that a male and female when
once mated remain together for the breeding-season ; and 3rd,
that all find mates. Now, if we have 7 species, each represented
by one individual of each sex, what is the probability that all the
species will be propagated ? And what the probability for the
propagation of none, or of but one, or of but two, or of but three
of the species ? The answers, as I have computed them, Bre es
follows :—The probability that none will be propagated is £834 ;
that 1 species will be is Boo ; that 2 species B04} ; that 3 species
3040 ; that 4 species =/2,; that 5 species -24,; that 7 species
sory: These numerators are found in the 7th line of a table of
figures which I call the Permutational Triangle. If we have 10
species, the probability that in any one trial no species will
match truly and be propagated is $234261; that 1 species will
match truly and propagate is $334269; that 10 will is sgghyaz-
This means that if 3,628,800 trials are made, one of them will
probably be a case in which each male pairs with the female of
the same species, while 1,834,961 will be cases in which none are
so matched, and 1,334,960 will be cases in which one pair is so
matched. It therefore appears that more than 8 of the proba-
bilities are against the continuance of more than one of the ten
species.

There will perhaps be some hesitation in receiving these
figures before I have given the method by which the results have
been reached ; but the necessary length of this paper, even when
restricted to the briefest discussion of general principles, induces
me to reserve my computations for another occasion. It is
not, however, necessary to have a complete solution of this
problem, in order to reach the conclusion that the origin of
separate races and species depends not only upon their adap-
tation to the environment and their mutual sterility when

THROUGH CUMULATIVE SEGREGATION. O51

crossing with each other, but also upon their Positive Segrega-
tion. We can further see (when considering an extreme case, like
either of the above-supposed cases) that Segregate Fecundity,
without the aid of Positive Segregation, must lead to extinction.
We have already seen that partial Segregation cannot by itself
prevent the fusion of species. It therefore follows that in order
to account for the continuance of divergent races we must sup-
pose either that the Segregation is complete, or that the divergent
evolution is strong enough to more than counterbalance the
influence of the occasional crossing, or that the partial Segre-
gation is aided by Segregate Fecundity or Segregate Vigour.
Between the members of species belonging to different orders
we find not only complete Segregation, but complete sterility
when attempts at crossing are made; but hope of gaining an
explanation of how these characteristics have arisen is found,
not in the study of those cases in which the process has been
completed, but in the study of the relations to each other of
species and varieties that are characterized by partial Segrega-
tion and mutual sterility, that is not complete. Here, again,
mathematical analysis will help us in understanding the subject.
Though I have not succeeded in constructing a complete mathe-
matical representation of all the grades of intermingling that
will take place, I have found a general formula that gives a close
approximation to the proportion in which two species will breed
pure as contrasted with the proportion of first crosses and their
descendants that will be produced, in any case in which the
degree of Segregation and the ratios of fertility for the pure and
crossed breeds are known. As my object is simply to show
under what conditions the pure races will continue without
being swamped by crossing, it is not necessary that I should
follow the action and reaction between the three-quarter-breeds.
I wish, however, to call attention to the fact that when the
number of the pure forms and of the half-breeds is constantly
decreasing, without a general decrease in the sum of the de-
scendants, it is evident that the three-quarter-breeds and their
descendants are increasing; and when a three-quarter-breed on
one side crosses with a three-quarter-breed on the other side, the
offspring will usually be about intermediate between the two
species ; therefore, where the two species are equally numerous,
if we find that the pure forms will disappear through fusion, we

252 REV. J. T. GULICK ON DIVERGENT EVOLUTION

may expect that the three-quarter-breeds will also disappear
through fusion.

In constructing my formula, it was found necessary to com-
mence by placing in the 1st generation of the half-breeds a more
or less arbitrary symbol; for the true symbol in each case is the
final one reached in the nth generation when 7 is a very high
number. The chief interest therefore centres in what can be
accomplished through the use of this formula for the nth gene-
ration. It seems to me to furnish a method of reaching the
final proportion of pure breeding that will be produced by any
form of combination between Positive Segregation and Segregate
Fecundity, and to give results that would require thousands of
years of continuous experimenting to reach in any other way.

Method of using Table III. (see p. 255).

By supposing 2 to be an indefinitely high number, and by
giving different values to M, m, and c, we shall have the the means
of contrasting the number of the pure-breeds with that of the
half-breeds, when the process has been long continued under
different degrees of Positive Segregation and Segregate Fecundity.

In the first place let us take a case in which there is no Segre-
gate Fecundity, that is M=m; and for convenience in computa-
tion let us make M=1, m=1. In every case where m is not
larger than M the fraction hat is less than unity, and the
sum of the geometrical progression of our formula will fall within
the limits of a number that can be easily computed by the well-

known formula S=;—_, in which a is the first number of the

progression, which in this case is 1, and qg is the fraction we

are now considering. Supposing c= 74, the fraction will be

(<7 dt Boe ee Oe 1) eee i
paper ery gt dem becomes ‘= 75 eee This

number 9 is therefore equal to the sum of this progression and
can therefore be used as the value of the infinite progression in
the formula for the th generation when x is a very high number.
Substituting these values we find that the zth generation of the
half-breeds equals the xth generation of the pure forms, each
being equal to 3% of A (M-—Mc)"-. A(M—Mcoe)"—! is a
vanishing quantity, for M—Mc is less than1. Every form is
therefore in time fused with other forms. But let us try higher

253

THROUGH CUMULATIVE SEGREGATION.

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JaquINt TeIyUT
"uorn.iauab

snyn JO

er(9-T)
o1(8-T)

¥,
¥,

ad os puv ‘spaaaq-oind jo

[

“¢ Surye} Lq punog st spooaq-jeq 9q} JO UoleioUEs pug ou,

= 212 1Le'6e
o1(8-1) = GLG-LEE6E
- OS0'LES °°

‘80 Aq payndurod o7(g.1) =¢0-168

e(-T)

2(8-T)

(8-T)

as ZES‘G

g-T
= OFS

8-1
= 008‘T

8-1
= 000‘T
"SPI9LQ-IANT
‘T “ON Apa

‘CIT oTqey, 998) T=w ‘g=py OL=

oyy [enbe ATdurts suoran poxtm wr0F
ATESwM9y} Suoure suoran wus0f AjouvA

“ON OF YIM pojsetzloo sv spoorq-jrey Jo soquinu oyy SULMOYS ‘worngndwop 1091j9UWY IIL

‘[ Wavy,

LINN. JOURN.—ZOOLOGY, VOL. Xx.

REV. J. T. GULICK ON DIVERGENT EVOLUTION

254

“MOTyeLoUEd pug IY} JO Sspaotq-F[eY ore Suridsyo tey} pur ‘ova LOT }O

ayy Uji SuUOIUN UILOJ ‘quay —AWIDV Afoueu ‘1epureutor OY} ‘ poqoola.t oq Jsnul §
zeqtinu penba ue YL suOTUN WULOF aWIV Eset} FO gUOUN PeXIUI ULLOF JVY} UOTE

poeaq-1oy4enb-sery} SuIEq Suridsyo toy} pus ‘speeaq-F[Vy_ JO
{gua JST oy} JO spoetq-oand jo roquunu [e}0} Ot} sjussoidet

AWU)V {Speetq-jTey Sureq Surrdsyo oy} ‘soapestuoy} SuoMME suOTUN WAOF JVI} spooiq-jTey JO toquinu otf} sjuesordor ywuoVy We} OUT, x

‘ON —IDUay +R — [)woy =Uorjze.1oued pug

eAvy OM ‘AT 10} (9— T) siq} Ur sulynyzTYsqng
CIN MOY + U(9— W)Moy =
(ypwoy twowy—awy)= “ — PUG
eux (Moy—-ANDV+HoVvy= “ PVG
"MOY =UOlye1oues 4ST

-spoadg-fyoq 9yp burjuasardan
sppnprmarpur fO saquUeanny

-sqnMoULUMOD UOT}eINduI09 oT} UoT{M soloods aand oy} 3
-royjo#0} Surpooiq vay spriqdy ot} 1oF puB sMOTUN-SsO19 otf
“pULY UAO ATOM YILM pootg yey} sot}

“uoryetoues pug=,(9IN—W)V
aavy om “ued pug oy} UL YZ LOF (9— J) Surynzrysqng

“ wp = sONWV
«“ pag = .CNWV
“pug = ANU)V

‘uoretoued JST = CNU)V
‘roquinu [VIuy = N:

‘mol aing ayn burguasacdas
syunprarpur fo LaquUenny

utjuosaader sTeNPTATpUT JO Toquinu [eT}UE oy = V 4'T
| LOF UOTVaoues Yous ul AjITIAoF JO OLPVA OHM POT
Loy UOTyeLoUIEs Yowe ur ApTILIF JO O1et OUI=W eT

‘9—T Tenbo sivas

Tea t= “=a saloods party UB YITM a18 suOTUN 9t[} JO OL puv soroods of} JO SHUT] Ot} UTGFTA Orv suolun oy} Jo “i Ua A “A

‘Surpooiq-ssoro JO oTVl OY} =I 9'T

“uoIyeSaideg ou} ‘a “2 ‘SuTpee.tq oand jo o1jet oy} =U PT

‘spaaig-aing 0} spaelq-fVH $0 uorysodoig ay, buinoys of vynusog havuruyard

‘TL TIaV I],

255

THROUGH CUMULATIVE SEGREGATION.

‘oures OF} OP VG} Spaorq-s[ey Jo uorrodo.ad oy} Sutoq @ pure ‘oyeFudoad pur Aqranyeur
04 MOIS 4eq} spaoiq-ernd jo uoyetoues yore Jo uorsodoard oy} Sutoq 4 {we TOF Mw pue "TT IOf ATT oyNyHsqns ysnut om ‘pasepisuoo st
Speedq sso10 puv aand jo anoSia ovyeyzer 04} ToyAd q[nsot [La 4VI} Suortodoad oyy oyeorpur qeqs 41 WU} O8 “B[NUIIOF SI} 4yoeII00 OF —aynay pue

*spo0iq-a.ng JO Jequinu oq} Sutaq d pur ‘spacrq-J,ey Jo xoquint ayy
om19q H ‘( eset! a 2N —W +1 )< Mom _ 4 a

ea = LT 8! 4say oy} YOIyM Jo ‘surzay w a ‘2 "2 “UOT}VIOUES OY} Jo JoquNuU £,
t : oW—W =
ey} Aq possordxo yey} sv suze} Luvm sv Suraieyuoo 6 ++: ET Ct +T soltes 04} Jo X ums oxy Aq peydyjnu jo —Wowy=
z— yar

Speciq-jTeq JO Uorvsoues yyw oy} pue ‘ (OW—W) X 1-«(21—W) ¥ = u9N—W) V= speeaq-eand JO wuoHnvioues YW eyT,—‘anar pus

‘wa £q porpdiyjnur speeaq-aand
JO ToHRIaUEs snotastd oy Surppe pur w(9Z—T) Aq spooaq-jpey jo uoNR«ouad snoraord oy} Surktdryqnur fq punoj orev woryetoues Aue jo
Spserq-jrey eyy pure ‘op — pl Lq spooaq-oand JO uoTet9u09 snotacad oy} Sutéydryynur Aq punos o18 UoTyetoued AUB JO spooaq-aing OU L— yng IST

(1+ (25*) +4 ) +f ) + ( ) al )) Xiu W)owy= “maw « y= —-W) V
ON-W), OW OW — oW-- ‘
‘(Gas ¥ Be = ‘. roe = =i (ee =) RON ODE ss. eae
“eC — IW) Moy + w(0Z— T)e(ON — Woy +-,(wloz—1))( PM — WMV + g(w(og—T))owy= “ aap va Wr= ON —W V
CIN W) Moy +H(9B— 1 (OW —W) wYt,(w(og—P))owy= pag . Pg=-(°N—W) V
‘PIN—W) woy+w(og—T)owy= “ pug “ pug==,(0—W) ¥
‘OMY =UOl}VIVUIT 4ST “aoljyeieues ysT= (OW —W)V
*Leq TInU [eryUay = V
“spoaug-f/OHT "SP90LQ-OLNT

‘Spaadq-ain_ 07 spaaig-fyory fo uowsodoud yg burarb ‘Aypunsagy ajnbauhay pun uornbashay sof vjnumsog padojaaag
‘TIL @1avy,

256 REV. J. T. GULICK ON DIVERGENT EVOLUTION

degrees of Segregation. If we make c= 1, or yp, we still
find that Half-breeds= Pure-breeds, while the latter are con-
stantly decreasing, which shows that imperfect Pcsitive Segre-
gation, without the aid of some quality like Segregate Fecundity,
cannot prevent a species being finally fused with other species, as
long as the whole number of each successive generation does not
increase.

Let us now consider cases in which the Segregation is incom-
plete but Segregate Fecundity comes in to modify the result.
Let M=2, m=1, c=). Substituting these values in our
formula, we shall tind that the sum of the infinite progression is
9$=18, And M—Mc=1%, which makes the half-breeds = the
pure formsxXem; and cem=7,. Let M=2, m=1, c=7h;
then Half-breeds=Pure formsx 75. Let M=2, m=1, c=};
then the infinite progression=1, M—Mc=1, and the pure forms
in each generation will equal A, and the half-breeds A x4,
Therefore Half-breeds= Pure-breeds x 4.

Let M=3, m=2, c=4; then the sum of the infinite pro-
gression=1, and the Half-breeds=4}x2xA(M—Me)»-1, and
the Pure-breeds=14 x A(M—Mc)"-!; therefore Half-breeds=
Pure-breeds x 2.

Let M=3, m=2, c=4; then Half-breeds= Pure-breeds x Z.

Let M=3, m=2, c=}; then Half-breeds=Pure-breeds x 2.

Let M=3, m=2, c=+; then Half-breeds= Pure-breeds x 2.

Let M=3, m=2, c=; then Half-breeds= Pure-breeds x 74.

Let M=3, m=2, c=7},; then Half-breeds = Pure-breeds
x ToT:

Tase IV.
Simplified Formulas for the Proportions in which Half-breeds

and Three-quarter-breeds stand to Pure-breeds when all are
equally vigorous.
From Table ITI. we learn that

EP me (1—2c)m )
Poem (1+ MoMe * 7 au
Whem (1—2c)m is less than M—Me¢, the series within the brackets is a de-
creasing geometrical progression, and we may obtain the value of the whole

series by the formula S=;—> Applying this formula we have
He Some 1

P M—Mc 1 _G—2e)m
M—Me

THROUGH CUMULATIVE SEGREGATION.

me * M—Mce
M—Me*M—Mo—mp ome

~ M—m-+(2m—Mye’

N0€

=?
*M—m+(Qm—Mye

257

(Formula 1)

(2)

If m'= the ratio of fertility for the Three-quarter-breeds, then according to

the reasoning given in Tables VII. and VIIL.,

a Ime
Ho M—m!'+(2m'—M)c :
and Le =
let ease i

(3)

(4)

The following solutions, as well as those given in Table V., are
obtained by substituting values for M, m, and c in formula (2) :—

When M=4, m=8, thenif
c=4, Half-breeds= Pure-breeds x #,

c=3, ” San ” x 3,
c=4, ” = ” x 3,
c=t, ” oa 9 x 3,
c=4, ” — 9 Xx 3,
c=7, ” = ” x 3,
C=%; ” = 9 x >
c=%, ” = ” x 3D
c= 'p 9 et 9 xX er
c=10) ” = ” x Tis
When M=5, m=4, then if
e= 3, Half-breeds= Pure-breeds x 4,
c=3, ” — ” x 4,
c=4, ” = ” x 4,
c=, ” _ ” x 4,
cC=% ” — 9 x 4,
c=7, ” iis ” x o>
c=%, ” — "99 x th
c=, ” — ” x i
c= ” = ” x ED
c=Thn ” == 9 Xx tos
= 100 ” = 9 x TOF

258

TABLE V.

REV. J. T. GULICK ON DIVERGENT EVOLUTION

TeRRtREE RS
le ER ee ee E
fe heat ee ee
Teeneee ee $
1D

Hi. Se ofS Ba ee “eo “Ses Mae

nN ko ia |S
(ies eae a a ae
= “ 23 4 is
He Sim. ale i a “leu a mh ots “b
S
ss ko
Go fit oe eR ok oe
S
> — ls) bry | of of a kao
a A a of’ of? a i — fs re
§
S a = SSS SS SO OS ae
e
I! | x
m2 Xx x x x x X 2X &xX
a 2 . 2 J 4 7 e
= 5 Q ripe Aaj rR AY nisAy AicAy Rie Ay na Ay “Pay Ay
o S68 Itt Ul 4k UU US be
SF owe SoA OR OR OR oe oe om oe
= Ts = ros ele Ae == aia == eines lies = [ie
© FY Ay
ey
me

8
2006

—— SS.

THROUGH CUMULATIVE SEGREGATION, 259

Observations on Table V.

This mathematical analysis of the effects of Positive Segregation
and Segregate Fecundity when co-operating brings distinctly into
view several important relations.

Ist. Incomplete forms of Segregation, that avail little or
nothing in preventing a form from being absorbed in the course
of time, become very efficient when strengthened by moderate
degrees of mutual sterility. Take, for instance, the line of the
table in which c=;4,. If 1 in every 100 unions is a cross with
some other form, the form will in time be overwhelmed, unless
other causes come in to counteract; but here we see that, if
Segregate Fecundity occurs in the ratio of 10 to 9, the pure
form becomes 12 times as numerousas the half-breeds; and if in
the ratio of 10 to 5, it becomes 100 times as numerous.

2nd. Again, if we take the proportional differences between
the different terms of the top line opposite c=, we shall find
them very unlike the differences that appear in the bottom line
opposite c= zo'5y- In the former the first term is 9 times as
large as the last; while in the latter the first term is more than
80 times as large as the last. This shows that when Segregation
is intense, differences in the degree of Segregate Fecundity pro-
duce greater contrasts than the same differences do when the
Segregation is slight.

3rd. A similar distinction is found when we compare the right-
hand column with the left-hand column. The smallest term in
the former is to the largest term in the same column as 1 to 899,
while in the left-hand column the greatest difference is as 1 to
100. This shows that when Segregate Fecundity is strongly
developed, differences in the degrees of Segregation produce
ereater contrasts than the same differences produce when the
Segregate Fecundity is but slightly developed.

4th. Once more let us consider the relations to each other of
the four terms that stand in the upper left-hand corner of the
table. Suppose that of some one variety of a plant species, cha-
racterized by Prepotential Segregation and Segregate Fecundity,
we have occurring in equal numbers four variations whose rela-
tions to other varieties are indicated by the figures given in these
four terms, while in their relations to each other they are com-
pletely fertile and not Segregated. Which variation will leave
the greatest number of pure offspring, that is the greatest number

260 REV. J. T. GULICK ON DIVERGENT EVOLUTION

of offspring belonging to the one variety to which the four varia-
tions alike belong? Evidently the variation represented by the
fraction 58 will have the greatest influence on the following
generation. But as the supposed conditions allow of exact com-
putation, let us look at the problem a little closer. If each varia-
tion numbers say a thousand individuals, then the number of
each that will breed true will be as follows:—Of the one repre-
sented by 59,, 526 will breed true and 474 will cross, —

i, 550 H 450°
ay, BSS'S ‘i 4445 ,,
8, 600 : 400" 9

And the next generation of each kind will be as follows: multi-
plying the pure parents by 10, and the hybrid parents by 8 or 9,
according to the value of m, we have of those represented by

+o, pure offspring 5260, hybrids 4266,

Tp ; 5500, ,, | Gams
To : 5555, ,, 8556,
Tp . 6000, .,, 8200.

There can, therefore, be no doubt that under such conditions the
average Prepotential Segregation and Segregate Fecundity of
the next generation will be considerably advanced, and so with
each successive generation till the average of the Pure forms is
represented by the fraction 58, and is surrounded by a circle of
variations, of which one will be represented by the fraction 75.
And from this new point continuous advance will be made toward
ever higher and higher grades of Segregation and Segregate
Fecundity ; though of course the process will be subject to
antagonisms and limitations arising from the principles of Self-
accumulating Vigour and Self-accumulating Adaptation. Let it,
however, be carefully noted that we have in this process the
manifestation of a new principle, for it rests not only on Self-
accumulating Positive Segregation but on Self-accumulating
Segregate Fecundity.

261 .

THROUGH CUMULATIVE SEGREGATION,

‘AvM owes oy} Ur posserdxa spoaaq-F[e]T oT} Jo INOSIA oy} =a ‘oyeSedord puv Azranyeur
0} Moad Jey} UOTyeAoUES Yows jo uoysodosd oy} sasi8 yvuy uoyovry v Xq possoidxe speorq-oing JO anost, = A ‘eTNuTAoy oAoge ou} UT

(--- _2AW—AW , 7) ?AW—AW _ Specaq-oing y pur { ~ 4s =speony-
( + au0e— 1) +1) oa = spacaq-ypeyy nO PUP * CAN — AW) X 1-«(AW— AN) V=sp20qq-erng Jo uoyvaeued yu

(., °AW-AW , (AW-AW i AW AW 4 + aioe 4
Gare oe eed a )+ + ow Gmegey) TH CAH ATONE, og OE. pe

‘MOljeIeUes YI — wu 1—-“9AW—-AWV
) FOANW — AW )20uy =

PAN — AW) (2m(0Z—T)) * -(aw(og—T)) * (aw(og—T))

(CANAD 4 CAW—AW) , PAW—AW) , AW- AW)

"(2A — AW)2AWY +am(0Z— TA — AW)20wWy + ,(amu(og — DICAW —AW)20wy + ,(aw(9Z—]))oamy — ‘uoryeroed UF *CANM— AW)V

PAN — AWW +am0F— EAN AW)wy + (0m(og—])oawy — -wonereweS pag «(0A W— ATR)
‘PAN — AN)202wy +aw(og—T)joawy ‘uornvseued pug (°AW—ATDV
“QU ‘HOVIIUET 4ST ‘OANW—-AN)V

sequina [eruy y
"spaatg-{) OFT *SP9dlQ-ILNT

('g o[NI 04 SUIPLOD0V “ITT qe, Mosz poyonazysu09) "Spaa4q-aINg
07 spaaiq-fvH Jo uornsodorg ay; buzarh ‘unobr, agnbaubay pun ‘hppunoay apnbashay ‘uornbashay sof njynusog

‘TA aTavy,

REV. J. T. GULICK ON DIVERGENT EVOLUTION

262

29AW— AW
f °AW— AWN eae ees ‘ ‘

Fede i
( PAW AW (CAN AN OAW—AW)\,...4 (2AN—AW d
linea) T Nau Ge m) flee (26— 0) oor or (o3- ) (AW) alee 5 AS ee core
Pork a] otal d
ee. \OAW AW + (24 (23—1 2S CAW—AW)V + (.2:0 (6-1) Ja ee AN Any = uoryeroues YI(E+w)
awe © (AI AWW + 0.0(9B—1)% 210 7 a Can Aiy— connie ira

* Qub OG (OAN—AIDV= toyetouss yw
[-u
‘spaaig-lagdonb-aaLyT,

—; sd1108 SULMOTIOF OY} UIEIGO OM “TTT PGT, SuTJON.YstIOD UI PESN SvAL
sv poyjom owes oy} Summsand pue ‘YOTYVAOMOS YW O[} UT ToqUINU STYy YA SuryAVyg ‘AqLIngvur oF MOIS [ILM Jeq} uOoTyeAoUed YWw OT{} JO eee

posrq-tojaenb-oeiyy, JO Toqumu 4vy SOUT 0% eonpord [IM yey} ‘Moroes TIT—w¢ eyy UT syuoand 9% dy (OAW—AIDY sureu

‘speorq-edng Jo toquinu [enbo uv YFTA yAOSMOO T[Las ToqtANU sTY} JO 9 pure {TA pue “A “AT SoTquy, UL UMOYS SB Pe Yor ‘speoiq-aing 04
Lid 2 : ss fe
pureys sposdeiq-j[eH] Gory Ul OLVeY oy Ssutaq an Xx A? AIW— AI) V = Uorqetoues oumes 011] Jo spoorqg-j[e Fy OY} pue : on AIN— ATID V¥= spe7tq

-oINT Jo UOIyVsoues snorsoad or} 4VYY “TA STARE Wor MOU OAL WOTPLAOUOS YYW OY} YILA WOTPeIndtMo9 ANO SULUSTAMMOA) *JOT[}O TOV YALA Pootq foyy
sv avy st UT spoorq-toqtenb-ooly.Z, JO suoljereues snotaoad Jo sjuvpusdsop oy} suid ‘spoorq-etng Jo tequinu [enbo we TIA 4.L0su09 ous spoo.iq-}[@H
jo uoryetgues snoracad oY} Jo (9 a0) 2f jo Suradsyo oy} oie UOTJVAOUES Youre Jo spooig-royavnb-soly,y, ey} Jey} puy om “T o[qvy, 0} Suruany,
‘spooaq-aing JO Joquinu oyj=q ‘“spoodq-JTeET Jo oquinu oyj= FF *ApANgVUt 07 UIod yey} Spoe.tq
-sojavnb-901G J, 94} Jo uoTjAodoad oy} Sulats uororsy v ‘a soulvs OY} AF AIPAC JO oVy= wu ‘ spacaq-toqtenb-seayy, JO taquunu eyi= J,
“spaalg-aiNd ay} 02 Spaatq-a,lvnb-aaLyT,
ayy fo uonwodorg ay7 burarh ‘enobry ayvbashay pun ‘kppunsag ayobauhagy ‘uornbashag sof opnu.coy

“ITA FTE4V TG

THROUGH CUMULATIVE SEGREGATION. 2638

TasreE VIII.
Simplified Formulas, giving the Proportions in which Half-breeds
and Three-quarter-breeds stand to Pure-breeds when we have
_ both Segregate Fecundity and Segregate Vigour.
From Table VI. we learn that
H muc (1—2c)mv
Bov—mvex (+itv—uvet *-- )
When the numerator, (1—2c)mv, is less than the denominator, MV—MVe,
the sum of the whole series within the brackets may be obtained in accordance
with the formula s=,—, in which § = the sum of the series, a= the first

term, and g = the constant multiplier.

| ah me « 1
*" P-MV—MVe j= 2e)me
MV—MVe
MvUE MV—MVe

MV —MVe* MV—MVe—mv42mve

MvC

MV —mv+(Qmv—MV)c erecta Reh Boh irs) - 46) its (Formula 1)
Applying the same method to the formula in Table VII., we find that
Ly EL ae m'v'e
i: ae MV —m'v' + (2in'v' —MYV)c’
| 9
j P—P * MV—m'v'+ (2m'v'—MV)ec dS a
and
Aes 2m'v'c (3)
MM ii@ay—Mys °°

1 M=10, m=5, m'=5, V=}, v= 75, ' =, C= To;

thoes 1 tbo +" tbo tet tag
soe Tie Tea Ta A To _ — — ?
Pte —ot(is— 42) tsi — ws | 88
and (as m=m', and v=v')
£ H 1 ris 7” H 1 ; ee |
Ho pte ; and pp XA sXe oae eae
if M=10, m=10, ta 30); V=i, v= 359 y= 3/55 c=;
then a 310 90 a ae
ae ee — sere eee = Tae ACE Pa a: mak. | >
Pet tro tt sete — 38

a EL,
and Fa ee—7rs and p> p* AT Bee

In this latter case, where the Vigour of Hybrids is ,'5 that a Pure-breeds,
while their Fecundity is equal to that of Pure-breeds, we find P = gz, Which is
the same result as that given in the 8th line of the last column of Table V.,
where the Fecundity of cross unions and of Hybrids is ,'5 that of Pure-breeds,
while their Vigour is equal.

264 REV. J. T. GULICK ON DIVERGENT EVOLUTION

The Influence of Segregate Vigour.

I think we may say we have here come in sight of one form of
the still wider fourfold law already mentioned; for on the same
principle that Segregate Fecundity increases when once allied
with partial Segregation in vigorous forms, Segregate Vigour
must also tend to increase when brought into the same alliance ;
and I believe it will be found that there is a similar principle
tending to the self-accumulation of Segregate Adaptation.

At the point where they both arise, that is during the
period that immediately follows the act of impregnation, it is
difficult to distinguish between the two principles, and the mor-
tality of the hybrid embryo before birth, or before it leaves the
egg, may be conveniently classed as Segregate Fecundity. *

Though the two principles are so closely related, it would be a
great mistake not to distinguish them; for there is no close
correspondence between the degrees in which the two qualities
occur in the relations of individuals or varieties ; and in some cases
we find Segregate Fecundity associated with Integrate Vigour.
The mule, though absolutely sterile, possesses vigour equal, if
not superior, to that of either parent. In the record of experi-
ments given by Darwin in ‘ Cross- and Self-Fertilization in the
Vegetable Kingdom’ mention is made of certain species in which
self-fertilized flowers are more fertile than the cross-fertilized,
while the plants produced from the crossed seed are the more
vigorous ; and of other species in which cross-fertilized flowers are
by far the most productive, while the plants produced from the
crossed seed are neither taller nor heavier than the self-fertilized.t
In the same work the common pea (Pisum sativum), the common
tobacco (Nicotiana tabacum), and Canna Warscewiczi are shown
to be more vigorous when raised from self-fertilized seed than
when raised from seed crossed with other individuals of the same
strain ; but in the case of the tobacco and the pea, great increase of
vigour is produced by a cross with a slightly different variety
while the fertility is increased but little if any.

But the most interesting of all his experiments as bearing on
the subject of Segregate Vigour, is given in the history of “ The
Descendants of the self-fertilized Plant, named Hero, which ap-
pearedin theSixth Self-fertilized Generation of Ipomea purpurea.’
“A cross between the children of Hero did not give to the

* See ‘ Origin of Species,’ 6th edition, p. 249, t See pages 322-329.

THROUGH CUMULATIVE SEGREGATION. 265

grandchildren any advantage over the self-fertilized grandchildren
raised from the self-fertilized children.” “And, what is far
more remarkable, the great-grandchildren, raised by crossing the
grandchildren with a fresh stock, had no advantage over either
the intercrossed or the self-fertilized great-grandchildren. It
thus appears that Hero and its descendants differed in consti-
tution in an extraordinary manner from ordinary plants of the
same species.” ‘If we look to the [ordinary] plants of the ninth
generation in table x., we find that the intercrossed plants [of
the same stock] were in height to the self-fertilized as 100 to 79,
and in fertility as 100 to 26; whilst the Colchester-crossed
plants [raised by crossing with a fresh stock| were in height to
the intercrossed as 100 to 78, and in fertility as 100 to 51.”*
The Colchester-crossed plants were therefore in height to the
self-fertilized as 1 to °78 x ‘79, or as 1000 to 616, and in fertility
as 1 to *51x°‘26, or as 1000 to 183; while the self-fertilized
descendants of Hero when crossed with the same fresh stock
not only had no advantage over those that had been continuously
self-fertilized for nine generations, but, as the details of the
experiment show, the advantage was on the side of the plants
raised from the self-fertilized seed. The experiment was con-
ducted under conditions decidedly unfavourable for the production
of healthy plants ; but, as it is usually found that the superiority
of crosses between varieties is most clearly brought to light when
the competitors are subjected to unfavourable circumstances, it
seems to furnish even stronger evidence of Segregate Vigour
being occasionally produced in the earliest stages of divergent
evolution, than would have been furnished if the same degree of
superiority in the self-fertilized plants had been obtained under
a less severe test. As the case is of unusual interest, I give the
details as recorded by Darwin :—

“Several flowers on the self-fertilized grandchildren of Hero
in table xvi. were fertilized with pollen from the same flower ;
and the seedlings raised from them (great-grandchildren of
Hero) formed the ninth self-fertilized generation. Several other
flowers were crossed with pollen from another grandchild, so
that they may be considered as the offspring of brothers and
sisters, and the seedlings thus raised may be called the inter-
crossed great-grandchildren. And, lastly, other flowers were
fertilized with pollen from a distinct stock, and the seedlings

* *Cross- and Self-Fertilization,’ pp. 47, 60, 61.

266 REV. J. 1. GULICK ON DIVERGENT EVOLUTION

thus raised may be called the Colchester-crossed great-grand-
children. In my anxiety to see what the result would be, I
unfortunately planted the three lots of seeds (after they had
germinated on sand) in the hothouse in the middle of winter,
and in consequence of this the seedlings (twenty in number of
each kind) became very unhealthy, some growing only a few
inches in height, and very few to their proper height. The
result, therefore, cannot be fully trusted; and it would be
useless to give the measurements in detail. In order to strike
as fair an average as possible, I first excluded all the plants
under 50 inches in height, thus rejecting all the most unhealthy
plants. The six self-fertilized thus left were on an average 66°86
inches high, the eight intercrossed plants 63°2 high, and the
seven Colchester-crossed 65°37 high; so that there was not
much difference between the three sets, the self-fertilized plants
having a slight advantage. Nor was there any great difference
when only the plants under 36 inches in height were excluded.
Nor, again, when all the plants, however much dwarfed and
unhealthy, were included.

“In this latter case the Colchester-crossed gave the lowest
average of all; and if these plants had been in any marked
manner superior to the other two lots, as from my former
experience I fully expected they would have been, I cannot but
think that some vestige of such superiority would have been
evident, notwithstanding the very unhealthy condition of most
of the plants. No advantage, as far as we can judge, was
derived from intercrossing two of the grandchildren of Hero,
any more than when two of the children were crossed. It
appears therefore that Hero and its descendants have varied
from the common type, not only in acquiring great power of
growth and increased fertility when subjected to self-fertilization,
but in not profiting from a cross with a distinct stock ; and this
latter fact, if trustworthy, is a unique case, as far as I have
observed in all my experiments.” *

Let us now consider for a moment what must be the result
when such a variation occurs in a wild species subject to the
ordinary conditions of competition. In the first place, it would
gradually prevail over other representatives of the same local
stock, both by its more vigorous growth and by its greater

* «Oross- and Self-Fertilization in the Vegetable Kingdom,’ pp. 50, 61.

THROUGH CUMULATIVE SEGREGATION. 267

fertility, especially in the case of flowers that failed of securing
across. And afterwards, when it came into competition with the
equally adapted variety from which it was partially protected by
Segregate Vigour, it would neither be driven out nor lose its
separate existence in a commingled race. It will be observed
that we have in such a case Local, Germinal, and Floral Segre-
gation, each producing partial effects which are enhanced by the
Segregate Vigour. In order to bring out the relation of these
factors to each other, let us assume definite values for each.

Let us suppose that 54, of the flowers are self-fertilized, 7% are
fertilized with pollen from another flower of the same plant, ;%5
are fertilized with pollen from other plants of the same new
variety, and =), are fertilized with pollen from the older variety
occupying contiguous areas. Therefore the sum of the segre-
gating influences, which is called the “ Ratio of pure breeding,”
and is represented by R in Table II. » equals 7 ;’;; and the “ Ratio
of cross-breeding,” represented by c in all the tables, equals +5.
Again, let us suppose that the fertility of the pure breeds is the
same as that of the half-breeds, but that the superior vigour of
the former is such that any one of the pure seeds has twice as
gooda chance of germinating, growing to maturity, and producing
seed as any one of the crossed seeds. The general effect on the
final result will in that case be the same as if the “ Ratio of
increase for the pure unions” (which I call M) equalled 10,
while the “ Ratio of increase for the cross unions” (which I call
m) equalled 5. Turning now to Table V., we can easily find the
ratio in which the number of pure-breeds will stand to the half-
breeds, if the conditions continue Jong; for in the column in
which m equals 5 and in the line marked c=;5 we find »,
which means that the half-breeds will equal the pure-breeds
multiplied by 7, or by 7b.

Segregate Vigour and Segregate Fecundity between Human Races.

My attention has recently been called to the following facts
relating to the Japanese and Aino races, who have for many
centuries met under circumstances favourable for interfusion
without any apparent effect of this kind. I quote from ‘ Me-
moirs of the Literature College, Imperial University of Japan,’
No. 1: “ The Language, Mythology, and Geographical Nomen-
clature of Japan viewed in the Light of Aino Studies,” by
Basil Hall Chamberlain, p. 43 :—

268 REY. J. T. GULICK ON DIVERGENT EVOLUTION

“With what logic, it may be urged, do you invite us to accept
a great extension of the Aino race in early Japan, when it is
a physiological fact, vouched for by so high an authority as
Dr. Baelz, that there is little or no trace of Aino blood in
the Japanese people? In reply to this some would perhaps
quote such examples as New England, whence the Indians have
vanished, leaving nought behind them but their place-names. In
Japan, however, the circumstances are different from those of
New England. There has undoubtedly been constant inter-
marriage between the conquerors and the native race upon the
Aino border. We can infer this from history. Those who have
travelledin Yezo know it by personal experience to-day. Never-
theless, these intermarriages may well consist with the absence
of any trace of Aino blood in the population. As a matter of
fact, the Northern Japanese, in whose veins there should be most
Aino blood, are no whit hairier than their compatriots in Central
and Southern Japan. Anyone may convince himself of this by
looking at the coolies (almost all Nambu or Tsugaru men)
working in the Hakodate streets during the summer months,
when little clothing is worn. But the paradox is only on the
surface. The fact is that the half-castes die out—a fate which
seems, in many quarters of the world, to follow the miscegenation
of races of widely divergent physique. That this is the true
explanation of the phenomenon was suggested to the present
writer’s mind by a consideration of the general absence of
children in the half-breed Aino families of his acquaintance.
Thus, of four brothers in a certain village where he staid, three
have died leaving widows without male children, and with only
one or two little girls between the three. The fourth has
children of both sexes; but they suffer from affections of the
chest and from rheumatism. Mr. Batchelor, whose opportunities
for observation have been unsually great, concurs in considering
this explanation as sufficient as it is simple. There are scores of
mixed marriages every year. There are numerous half-breeds
born of these marriages. But the second generation is almost
barren; and such children as are born—whether it be from two
half-breed parents, or from one half-breed parent and a member
of either pure race, are generally weakly. In the third or
fourth generation the family dies out. It may be added that
the half-breeds have a marked tendency to baldness, and that
their bodies are much less hairy than those of the genuine

THROUGH CUMULATIVE SEGREGATION. 269

Ainos. This fact has doubtless helped to cause the divergence
of opinion with regard to Aino hairiness. For the com-
paratively smooth half-breeds usually speak Aino, dress Aino-
fashion, and are accounted to be Ainos, so that travellers are
likely to be misled, unless constantly on their guard. There
seem to be half-breeds in all the villages whither Japanese
pedlars and fishermen have penetrated. There have therefore
probably, at some time or other, been half-breeds in every
portion of Japan where the two races have come in contact.”

If these two races were equal in civilization and in natural
adaptation to the environment, or if one race was specially
adapted to mountain life and the other to life by the sea-shore,
it seems probable that they might permanently occupy adjoining
countries without losing any of their distinctive characteristics.
Broca, after careful collation of all the information that could be
gathered from the publications of travellers and historians, reaches
the conclusion “ that alliances between the Anglo-Saxon race and
the Australians and Tasmanians are but little prolific; and that
the mulattoes sprung from such intercourse are too rare to have
enabled us to obtain exact particulars as to their viability and
fecundity.”* I have no means of knowing whether later investi-
gations in Australia and other parts of the world have thrown
fuller light on the mutual fertility or sterility of the more diver-
gent human races, but I am inclined to think that the interest in
the subject has declined since Darwin has shown that such data
ean never afford proof that the different races of man are not
descended from commou ancestry. There are, however, signs
that a renewed interest in the subject is being awakened through
the realization that it has a direct bearing on the theory of the
origin of species.

Impregnational Segregation a Cause of Divergence in both its
Earlier and Later Stages.

As we have already seen, the negative factors+ Segregate
Vigour and Segregate Fecundity would tend to produce extinc-
tion if not associated with positive forms of Segregation. But

* See ‘Phenomena of Hybridity in the Genus Homo.’ By Paul Broca.
English translation, published for the Anthropological Society of London by
Longman, Green, Longman, and Roberts (1864), pp. 45-60.

+ For a definition of Negative Segregation see page 238 of this paper.

LINN. JOURN.—ZOOLOGY, VOL. Xx. 21

270 REV. J. T. GULICK ON DIVERGENT EVOLUTION

in the case of organisms whose fertilizing elements are distributed
by wind and water, the qualities that produce these negative
forms of Segregation are usually accompanied by those that pro-
duce Prepotential Segregation, which is in an important degree
positive. But even Prepotential Segregation, when produced by
mutual incompatibility between a few individuals and a numerous
parent stock, depends for its continuance and development on
Local, Germinal, or Floral Segregation, partially securing the
intergeneration of the few that are mutually compatible. On
the one hand, Impregnational Segregation depends on some
degree of Local, Germinal, or Floral Segregation which is a con-
stant feature in most species; but, on the other hand, not only
do these initial forms of Positive Segregation fail of producing
any permanent divergence till associated with Impregnational
Segregation, but the more effective forms of Positive Segregation,
such as Industrial, Chronal, Fertilizational, Sexual, and Social
Segregation, often depend on Impregnational Segregation, inas-
much as the divergence of endowments which produces these
depends on Impregnational Segregation. Moreover, in all such
cases, increasing degrees of diversity in the forms of adaptation,
and consequently of diversity in the forms of natural selection,
must also depend upon these negative factors, which in their
turn depend on the weak, initial forms of Positive Segregation .
Divergent evolution always depends on some degree of Posi-
tive Segregation, but not always on Negative Segregation.
Under Positive Segregation of a rigorous form (as, for example,
complete Geographical Segregation), considerable divergence may
result without any sexual incompatibility. Darwin has shown,
by careful experiments, that Integrate Vigour and Fecundity is
the relation in which the varieties of one species usually stand to
each other. This fact does not, however, prove that the more
strongly divergent forms, called species, which are prevented from
coalescing by Segregate Vigour and Fecundity, did not acquire
some degree of this latter character before any permanent diver-
gence of form was acquired. Their having acquired this segre-
gating characteristic may be the very reason why their forms are
now so decidedly different, for without it they would have been
swallowed up by the incoming waves of intergeneration. Again,
we must remember that forms only moderately divergent are
habitually classed as different species if they are separated by
Segregate Vigour and Fecundity (that is by some degree of

THROUGH CUMULATIVE SEGREGATION. Tit

mutual sterility), unless observation shows that they are of
common descent. These two considerations sufficiently explain
why the varieties of one species are so seldom reported as
mutually infertile. Notwithstanding this, the experiments of
Gartner and of Darwin, already referred to at length, seem to
show that Segregate Fecundity and Vigour may arise between
varieties that spring from one stock. In view of these cases, we
must believe that in the formation of some, if not many, species,
the decisive event with which permanent divergence of allied
forms commences is the intervention of Segregate Fecundity or
Vigour between these forms. Positive Segregation, in the form
of Local, Germinal, or Floral Segregation producing only tran-
sitory divergences, always exists between the portions of a species
that has many members, but as it does not directly produce the
Negative Segregation which is, in such cases, the necessary ante-
cedent of permanent divergence, we cannot, in accordance with
the usage of language, call it the cause of the permanent diver-
gence. Moreover, though it may be in accordance with ordinary
language to call the Negative Segregation, which is the immediate
antecedent of the permanent divergence, the cause of the same, it
will be more correct to call the coincidence of the Negative and
Positive Segregations the cause, and still more accurate to say that
the whole range of vital activities (when subjected to the limita-
tions of any sexual incompatibility that corresponds in the groups
it separates to some previous but ineffectual Local, Germinal,
or Floral Segregation), will produce permanent divergence.

In many cases not only is the entrance of Impregnational
Segregation the cause of the commencement of permanent diver-
gence, but its continuance is the cause of the continuance of the
divergence. The clearest illustration of this is found in the case
of plants that are fertilized by pollen that is distributed by the
wind. All the higher, as well as the lower, groups of such plants
would rapidly coalesce if each grain of pollen was capable of
producing fertilization, with equal certainty, promptness, and
efficiency, on whatever stigma it might fall. We may also be
sure that, with organisms that depend upon water for the dis-
tribution of their fertilizing elements, Impregnational Segrega-
tion is an essential factor in the development of higher as well
as of lower taxonomic groups.

It is important to observe that, in the cases under considera-
tion, the inferior fertility or vigour resulting from the crossing of

272 REV. J. 1. GULICK ON DIVERGENT EVOLUTION

the incompatible forms is as truly a cause of divergence as the
inferior opportunity for crossing which from the first existed
between the members occupying different localities or between
the flowers growing on different trees of the same species. The
former has been called Negative, and the latter Positive, Segre-
gation, not for the sake of distinguishing different grades of
efficiency, but for the sake of indicating the different methods of
operation in the two classes of Segregation.

Institutional Segregation is the Reflexive form of Rational
Segregation. It is produced by the rational purposes of man
embodied in institutions that prevent free intergeneration be-
tween the different parts of the same race.

As the principal object of the present paper is to call attention
to the causes of Segregation acting independently of effort and
contrivance directed by man to that end, it will be sufficient
to enumerate some of the more prominent forms under which
Institutional Segregation presents itself, noting that some of
these influences come in as supplemental to the laws of segrega-
tion already discussed, simply reinforcing by artificial barriers
the segregations that have their original basis in nature. The
chief forms that should be enumerated are National, Linguistic,
Caste, Penal, Sanitary, and Educational Segregation ; and if we
had not already considered Industrial Segregation in the previous
chapter, that might be added.

ConcLUDING REMARKS.

Besides Artificial and Institutional Segregation, which depend
on the rational purpose of man, we have now considered numerous
forms of Segregation, resting on no less than 18 groups of purely
natural causes. Owing to the length of this paper I deem it
wise to bring it to a close without discussing the laws that co-
operate in intensifying the effects directly produced by the
segregative causes already considered. As I have shown in
Chapter II., Segregation is not simply the Independent Genera-
tion of the different sections of a species, but the Independent
Generation of sections that differ; and though no one will
believe that any two sections of a species are ever exactly equi-
valent, it is evident that the degrees of difference may be greater

THROUGH CUMULATIVE SEGREGATION. 273

or less, and that whatever causes a greater difference in two
sections that are prevented from intergenerating will also be a
cause of increased Segregation.

It has been observed that some of the causes enumerated in
this and the previous chapter are primarily separative, and that
no one of those that are primarily segregative is at any one time
segregative in regard to many classes of characters. As several
forms of Segregation may co-operate in securing a given division
of a species, and one form is superimposed upon another, the
aggregate effect must be incalculably great; but we easily per-
ceive that it may be indefinitely enhanced by causes producing
increased divergence in the segregated branches. The causes
which produce monotypic evolution when associated with Inter-
generation must be equally effective in producing polytypic
evolution when associated with Segeneration, whether in its
separative or segregative forms. But the discussion of Intensive
Segregation must be reserved for another occasion.

Believing that the study of Cumulative Segregation in its re-
lations to the other factors of evolution will throw light on the
origin of species far beyond what I have been able to elicit, I trust
the subject will secure the attention of those who enjoy better
opportunities than I do for carrying forward such investigations.

26 Concession, Osaka, Japan,
May 12, 1887.

APPENDIX.
Classified Table of Forms of Segregation.

A.
Environal Segregation.

(a) Industrial Segregation.
Sustentational.
Defensive.
Nidificational.

(b) Chronal Segregation.
Cyclical.

Seasonal.

274 EVOLUTION THROUGH CUMULATIVE SEGREGATION.

: Migrational.
eieuza } Transportational.
heh Geological.

(d) Fertilizational Segregation.
(e) Artificial Segregation.

B.
Reflexive Segregation.

(a) Conjunctional Segregation.

Social.
Sexual.
Germinal.

Floral.

(6) Impregnational Segregation.
Segregate Size.
Segregate Structure.
Prepotential Segregation.
Segregate Fecundity.
Segregate Vigour.

C.
Intensive Segregation.

(a) Assimilational Intension.
(6) Stimulational Intension.
(c) Suetudinal Intension.
(d) Correlated Intension.
(e) Integrational Intension.
(f) Selectional Intension.
(g) Fecundal Intension.
(h) Eliminational Intension.

OVICELLS OF CYCLOSTOMATOUS BRYOZOA. 275

On some Ovicells of Cyclostomatous Bryozoa.
By Artaur Wm. Waters, F.LS., F.GS.

[Read 5th April, 1888.]

(Puate XIV.)

For the determination of the Cyclostomatous Bryozoa the ovi-
cells are certainly most important characters, and in many cases
are undoubtedly of generic value; but how far this is universally
the case we are not yet in a position to decide. It may, how-
ever, safely be said that the Cyclostomata will never be rescued
from their present confusion until we are more fully acquainted
with these receptacles. Though of such paramount importance,
they are not known in a large majority of species. This is partly
on account of insufficient search, but more from the fact that
large numbers of specimens are often found without any ovi-
cells ; so that in some common species they are as yet unknown.
For instance, Hornera concatennata, Reuss, a fossil found abun-
dantly in the Miocene of Italy, Germany, Austria, and Hungary,
has, so far as we know, never furnished an ovieell.

It will thus be seen that with the Cyclostomata it is often the
case that abundant material is necessary for true appreciation of
the characters, and therefore it was a considerable disappoint-
ment that the results of the ‘ Challenger’ Expedition were not
richer in this respect. Jam, however, now able to add figures of
the ovicells of three species found by the ‘ Challenger,’ one from
‘Challenger’ material, and two from specimens in my own collec-
tion from other localities.

Hornera Fissunara, Busk. (Pl. XIV. figs. 1, 3, 4, 7.)
Taking them in order: when, through the kindness of Mr. John
Murray, | received two colonies of “Zdmonea fissurata,” Busk, from
‘Challenger’ “Station 320,” near Monte Video, I was surprised
to find a dorsal ovicell upon one, giving sufficient ground for
removing it to Hornera; but the structure of both the front and
back would have led me to do this independently of the ovicell.
As Miss Busk had kindly sorted and forwarded the specimens,
I informed her what I had found ; and she replied that she had
also noticed them, and mounted some for the British Museum,
and added that it was to be regretted that “ there did not happen
to be any among the rather numerous specimens selected for
LINN. JOURN.—ZOOLOGY, VOL, XX. 22

-—se
276 MR. A. W. WATERS ON SOME

examination by Mr. Busk”; who would have placed it, she
considered, under Hornera if he had noticed the ovicell.

In Idmonea I believe the ovicell is only known in TI. radians,
Lam., I. atlantica, Forbes, I. gracillima, Busk, I. concava*, Reuss,
I. serpens, L., I. eboracensis, Busk, I. radicata, Kirkpatrick ; and
in all is on the anterior surface ; but both in Zdmonea and Hor-
nera the position may be axillary, or far away from the bifurca-
tion ; and the position in relation to the axils can seldom or never
be looked upon as characteristic. I have before me a MS. list of
130 Idmonee, some of them synonyms, though all have been con-
sidered as separate species ; but only in the few cases mentioned
is the ovicell described. An ovicell has been figured by Hagenow
as Celophyma granulata on “ Idmonea”’ lichenoides, Goldf.; but
T am not sure that this is a true Jdmonea.

From a list of 74 species of Hornera, it seems that the ovicell
is only known in H. frondiculata, Lam., H. foliacea, MacG..,
fH, violacea, Sars (practically dorsal), H. galeata, Sm., H. robusta,
MacG., H. ramosa, MacG., H. lichenoides, L.t Goldstein (Trans.
Roy. Soc. Vict. 1881, pl. i. figs. 6, 7, 8) has named a ‘ Chal-
lenger’ specimen from Marion Island AH. subdubia, with “ ovi-
cells’; but from the figure it might be Hydrocorallina. Reuss
- has described as Ceelophyma striata an ovicell which, as pointed
out by Manzoni, probably belongs to H. hippolyta, Defr. In
all these cases the ovicell is dorsal, forming a distinct chamber ;
and this is also the case in Hornera fissurata. In this last it
is longitudinally ridged, and in the furrows are pits, giving,
when not calcined, an areolated appearance.

The position and nature of the opening of the ovicell is most
‘important, often, in fact, more so than the shape of the ovi-
cell. The opening is, however, formed after the ovicell is com-

* Not described ; but a recent specimen from Naples has the ovicell as a
raised chamber at the bifurcation. It nearly encloses one of the series, and
the ovicellular opening is by the side of the series. In another specimen
there is a slight central inflation between the series; but this does not seem to
be mature.

t I find that H. lichenoides, L., occurs in Naples, together with H. frondi-
culata, and has an ovicell with a rib down the middle, as figured by Smitt
and Alder, whereas Hincks figures a transverse rib. It has been looked upon
as an exclusively northern form; but the ‘Challenger’ found it off Monte
Video; and, as mentioned, it lives in the Mediterranean. Among the ‘ Chal-
lenger’ material in Edinburgh I have since seen the ovicells on specimens from
Simon’s Bay. These have a central rib, and also a transverse one from the
ovicelluiar opening to the median rib.

OVICELLS OF CYCLOSTOMATOUS BRYOZOA. 277

plete, so that frequently this important character cannot be made
out ; and inmy specimen I could not find any opening, but one
of the lateral zocecial tubes is much larger than usual, and looked
as though this change was connected with the functions of the
ovicell. I therefore wrote to Mr. Kirkpatrick, of the British
Museum, explaining what I had seen in my specimens, and asked
him if he would look at those in the Museum. He kindly did so
at once, and wrote that he examined ten ovicells, “and found
in each instance that the last one or two zoewcia of a series in
proximity to an ovicell are not only enlarged, but considerably
altered in direction. In eight instances the tube curved for-
wards and upwards so as to open on the anterior surface.” In
two the tubes opened laterally. “In every case the wide tube
is to some extent connate to a zoecial series ; but in some cases
the former curves away from the latter in such a way as to make
the wide tube appear to be part of the ovicell, and not a meta-
morphosed zocecium ”’ *.

I have already referred (Ann. Mag. Nat. Hist. ser. 5, vol. xx.
p- 255) to the fact that the ovicells of the Cyclostomata have
the surface nearly always perforated with much more numerous
pores than the rest of the zoarium. In the present case the
ridges are narrower and the pits somewhat finer than on the rest of
the dorsal surface ; and at the base of these pits, which are some-
times spoken of as pores, there are several fine pores, whereas in
the pits on the dorsal surface, except when elongated, there is
but one pore at thebase. It will thus be seen that in this case
the pores on the ovicell are much more numerous than on the
rest of the zoarium.

Figures (1 and 3) are given from calcined specimens of the
front aud back of the zoarium, and I consider that preparations
should be thus made wherever there is available material; for
figuring with all the organic integument is like taking a photo-
graph of a lady with a thick veil down. Calcined specimens can
be directly compared with fossils.

The anterior surface has large pores, or rather pits, following the
lines of the zoccia, with one or two pores at the base of the pit.
The way in which the ornamented appearance is formed is shown
more clearly in this species than in any other I have seen. At

* In the ‘Challenger’ specimens in Edinburgh ovicells occur in a large
number of cases, with a large lateral tube curving forwards,

278 MR. A. W. WATERS ON SOME

the growing end the zocecial tubes are more or less angular, and
down each side there is a row of rather small pores (fig. 3);
as growth proceeds, calcareous matter is added in ridges both on
the front and back surfaces, and then these small pores are left
at the bottom of the pits. I figured and referred to these pits
in Hornera frondiculata in the Quart. Journ. Geol. Soc. vol. xl.
p. 677, pl. xxx. fig. 8.

The external structure of the growing ends of the outside is
the same as seen in the inside in other parts when broken down, or
when sections are made. The position of the interzoccial pores
is often very characteristic, and can also sometimes be used in
fossil species. These internal pores may be in parallel rows
close together or far apart, and may be approximately regular
or irregular; and the distance apart is a point of great import-
ance; but as yet little attention has been given to these pores
in published papers. An absolute regularity does not seem ever
to occur; but the plan can very frequently be seen.

There is one structure which lam not able to explain. In the
interior of the zocecial tubes there are several protuberances
projecting forwards (fig. 7), either short or half as long as the
width of the zoccial tubes, and much curved forwards. They
do not seem to be the commencements of “closures,” and
whether they are used for attachments cannot be decided from
my dried specimens. Although I have sections of a very large
number of Cyclostomata, I have not come across anything of the
kind before. It does not seem likely that it should be compared
with the rays of Entalophora intricaria, Lichenopora, and Hetero-
pora (see Quart. Journ. Geol. Soe, vol. xlii. p. 340, &c.); but
both are as yet unexplained. In the coarsely pitted structure
this species resembles Idmonea radians, Lam.; but the genus
Idmonea usually has a finely punctured surface.

The zocecia are arranged in very distinct series, suggesting at
first Jdmonea; but the serial arrangement is not uncommon in
Hornera; for in H. lichenoides it occurs decidedly in some spe-
cimens, and in the fossil H. concatenata, Rss., and H. serrata,
Rss. (non d’Orb., non Menegh.), it is very marked ; again in
H. lichenoides the amount to which the zocecial tubes are exserted
is variable, and no doubt this and H. fissurata are closely allied.

Ipmonrea Meneouint, Heller. (Pl. XIV. fig. 2.)
The ovicell taking the place of one of the lateral series is a

OVICELLS OF CYCLOSTOMATOUS BRYOZOA. 279

unique structure ; and, so far as I am aware, nothing of the kind
has before been found. The base is narrow, but the sac widens
out, and then again becomes narrower; the surface has a few
large oval pores, and the ovicellular opening is a raised tube at
one upper corner. The zocecial tubes spread out at the ends.

It is placed, with hesitation, under I. Meneghini, as there are
only four zocecia to a series; but Heller only figures four, and
the number is not always constant in a colony. In a specimen
from Naples, without an ovicell, of what I considered J. Mene-
ghint, the zocecial tubes do not spread out at the end in the same
way ; but sometimes a trace of this structure can be seen. The
dorsal surface near tue ends of the branches is finely punc-
tured, but near the base there are larger openings. I have
only the one specimen with ovicells.

IDMONEA IRREGULARIS, Meneghini. (Pl. XIV. figs. 5 and 6.)

This was found by the ‘ Challenger’ off the Azores; and Mr.
Busk (p. 14) says, “ ocecial chamber ? ”

I have, however, previously (see Ann. Mag. Nat. Hist. ser. 5,
vol. xx. p. 257, and Quart. Journ. Geol. Soe. vol. xl. p. 687)
referred to the dorsal ovicell of JZ. irregularis; but as it has
never been figured, a normal ovicell is given (fig. 5) and a short
abnormal one (fig. 6).

The ovicellular aperture is wide, with a raised irregular funnel-
shaped openivg ; but in the young ovicells the upper part is flat,
sloping inwards. One colony has eight ovicells, and five of
these are intact, without showing any opening; and from this
we see how misleading this specimen alone would have been,
as the young ovicells are without external orifices.

A somewhat similar ovicell occurs on Filisparsa orakeiensis,

Stol. (doc. ett. p. 687).

Ipmonea Mitneana, d’Orb. (Pl. XIV. fig. 8.)

I have already referred to the ovicell of Z. Milneana (Ann.
Mag. Nat. Hist. ser. 5, vol. xx. p. 256), and give a figure from a
Capri specimen, and may repeat that the British Museum speci-
mens of J. notomala, B., and J. Milneana, @Orb., are so similar
that they might be fragments of the same colony. The ovicell
is figured from a rather broken-down specimen, and therefore
there is some restoration of the zocecial tubes.

280 MR. A. W. WATERS ON

EXPLANATION OF PLATE XIV.

Fig. 1. Anterior surface of Hornera fissurata, Busk, drawn from calcined

specimen.

2. Idmonea Meneghini, Heller. Naples.

3. Dorsal surface (growing end) of Hornera fissurata, B.

4. Dorsal surface of Hornera fissurata, B., showing ovicell.

5. Normal ovicell of Filisparsa irregularis, Meneghini. Dorsal surface,
Naples.

6. Short ovicell of Filésparsa irregularis, Menegh.

7. Section of zoccial tubes of Hornera fissurata, B., showing interior
projections.

8. Idmonea Milneana, d’Orb., from Capri.

Note.—I also found that in Gephyrophora polymorpha, B.,
dredged by the ‘ Challenger’ from Simons Bay, there is an entirely
concealed ovicell. Itis around sac quite free except at the border
of the opening; whereas in other Chilostomata, so far as my
experience goes, the wall of the ovicell is for a considerable
part attached to the zocecial walls.

On the Ovicells of some Lichenopore.
By ArtHur Wm. Waters, F.L.S., F.G.S.

[Read 3rd May, 1888. ]
(Prats XV.)

By Lichenopora I understand a genus with eancelli between
the rays; but there are other discoid forms with the zoeecia
arranged in radial series, so that the mode of growth is common
to several genera. For instance, the fossil Actinopora regularis,
d’Orb., does not seem to have any cancelhi, and I cannot agree with
my friend Mr. Hincks in placing it under Lichenopora, but should
consider it ALultitubigera.

Dr. Jullien * would take us back to the name Disporella of
Gray, and points out the curious mistake that was made for so
many years in calling this genus Discoporella, Gray ; whereas
Discoporella was a name given by d’Orbigny to some Chilostomata
which would now be Cupularia or allied genera. The genus
Lichenopora is, however, older than Gray’s, and is now well
established, so that I am unable to understand why Dr. Jullien
wishes us to return to Disporella. On the other hand, another

* Mission du Cap Horn: Bryozoaires.

Hollick. lith.

Linn. Soe. Journ. Zoou. Vor.XX. Pl. 14.

Imp. Camb. Sci. Inst Co.

Waters del.

OVICELLS OF BRYOZOA

OVICELLS OF SOME LICHENOPORS. 281

recent writer, Mr. Th. Marsson*, revives Discocavea and Defrancia
tor Lichenopora, though his Defrancia contains species which,
according to what I have said above, cannot be placed with Licheno-
pora, and his Discocavea is entirely based upon the radii being
uniserial ; but this cannot be looked upon as a generic character,
and even in specific determination must not be used with too much
confidence, as there are some species where the rays are in parts
uniserial, in others biserial.

In some species there are spines arising from various parts of
the surface of the zoarium, but these do not appear to furnish
constant characters; and, further, I have on one or two occasions
called attention to the great variation to which the shape of the
peristome is subject in different parts of the same colony; on
the other hand, which side of the peristome is prolonged seems
to be of considerable diagnostic importance.

The internal “rays”’ in the zoccial tubes and cancelli should
be more studied, and I should consider the denticulation of recent
Heteropora cervicornis, @ Orb. (Journ. R. Micr. Soe. vol. 11. p. 392,
pl. xv. figs. 9-11), Lichenopora bullata, MacG., L. echinata,
MacG., and L. pristis, MacG., as the equivalents of the rays with
knobs in Lntalophora intricaria, B., Lichenopora radiata, Aud.,
L, reticulata, MacG., L. Holdsworthii, B., &c.

Further, the position of the connecting-pores in the interior
walls of the zocecia should be examined.

Having seen how many of the characters used for diagnosis
are variable, it is quite clear that numerous species made on ac-
count of a difference in some unimportant character can never be
recognized again, making a better acquaintance with this genus
very desirable ; and it seems that a knowledge of the ovicells may
often assist us in more fully grasping the amount of variation
caused by the conditions under which the colony grew, and may
show the amount of variation in different parts of acolony. We
are as yet unable to say how far the ovicells may be trusted for
specific determination ; but there is no doubt that they are for this
purpose of value. Also as some species have been described with-
out ovicells and some with, in cases where the author has not
appreciated whether they were present or not, exact studies in both
conditions are required. Inso recent a book as Busk’s ‘ Catalogue
of the British Museum Polyzoa,’ pt. iii, where 14 species are
described, the ovicells are never mentioned, nor are they in the

* Bryozoa der weissen Schreibkreide der Insel Riigen, 1887.

282 MR. A. W. WATERS ON

‘Challenger’ Report. A ‘Challenger’ specimen from Tristan da
Cunha was submitted to me named L. fimbriata. This is L. echi-
nata with an ovicell, so that the name fimbriata must be dropped
The ovicells sometimes occur as inflations over the central area,
as in L. grignonensis, B. (see Ann. Mag. Nat. Hist. ser. 5, vol.
xx. pl. vil. fig. 4), LZ. nove-zelandia, B., L. wanganuiensis, Waters
(Quart. Journ. Geol. Soe. vol. xliii. p. 346), L. Holdsworthi, B.,
L. echinata, B.; or it may be nearer to the edge, showing the
ovicellular ducts as figured by Mr. Hincks in Lichenopora radiata.
Although now keenly on the look-out for ovicells, it is a very
difficult study, which can progress but slowly and only be done
satisfactorily with ample material collected from a few localities,
and one cannot help feeling despair when trying to determine
the Lichenopore ; but it is a genus which, when understood, is
perhaps more likely than any other to -throw light upon the
Cyclostomata generally and upon some fossil forms.

LICHENOPORA CALIFORNICA, Busk. (PI. XV. fig. 1.)

Unicavea ealifornica, d’Orb. Pai. Frang. p. 972.

Diseoporella californica, Busk, Cat. Mar. Poly. pt. ui. p. 32, pl. xxx. fig. 5.

Specimens from Port Western (Victoria) given to me by Miss
Jelly seem to be the species described by Mr. Busk; but as
d’Orbigny’s description might apply equally well to three or four
species, it seems preferable to consider Mr. Busk as the author.

The radii are usually bi- or triserial, though in parts only uni-
serial, and in one specimen most of the rays are uniserial and only
locally biserial ; the interserial spaces have round or angular cells,
but when these interserial cancelli are examined at a level below
the surface they are usually seen to be round. The mouths of the
cells are prolonged on the distal margin, namely, the side nearest
to the border of the zoarium. Zocecial tube about 0:06 mm.
wide inside. When the ovicellular cover is removed the central
area is seen to have large reticulated spaces. The upper part of
the ovicell is formed by a finely perforated plate extending between
the rays like the cogs of a wheel. There is a thick calcareous
rim at the border of this area.

The thin calcareous border of the zoarium is not shown, as this
is very variable in size, and probably of no specific value.

Loe. California (Busk); Port Western (Victoria).

Licuenoprora EcHINATA, MacG. (Pl. XV. figs. 2, 3, 6.)

Discoporella echinata, MacG. Trans. Roy. Soc. Vict. vol. xx. p. 127,
fig. 4.

OVICELLS OF SOME LICHENOPORE. 283

Through Miss Jelly’s kindness I have been able to examine a
considerable number of specimens of Lichenopore from Victoria,
and an abundant one is, I think, the LZ. echinata of MacGillivray.
The zocecia are irregularly arranged, the peristome is much pro-
duced on the proximal edge, namely, the margin nearest to the
centre of the zoarium, sometimes with one process, sometimes
divided into several ; numerous fine spines usually grow from the
zocecial tubes as well as from the cancelli and the surface of the
ovicell. The cancelli are angular with rounded corners, though
when the tube is examined at some depth below the surface it is
often seen to be round, and some of the cancelli may be called
round, though that is not the usual character; the entire inner
surface of the cancelli is denticulate. The ovicell is formed by
an inflated crust covering all the central area of the zoarium, and,
as shown in fignre 6, it starts from numerous places, ultimately
meeting iu the centre: at first this crust is very thin, but in the
mature condition is deeply pitted; the surface is then somewhat
ribbed and there are numerous small pores at the base of the pits.
In my previous paper to this Society (p. 277) I alluded to some-
what similar pits in Hornera.

As soon as the growth of the ovicell commences the basal reticu-
lated cells of the central area are covered over by a thin pellicle
with several perforations to each cancellus. Both the upper and
under surface of the ovicell is shown in figure 6. Figure 3 is drawn
from a specimen bent up at the two sides, and it is impossible to
figure it satisfactorily, as it cannot all be seen in one focus.

The opening of the ovicell is a wide tube low down at the
border of the inflation, directed horizontally, much in the same
way as in L. ciliata (Ann. Mag. Nat. Hist. ser. 5, vol. xx. p. 263,
pl. vu. fig. 5); but I have only seen the one ovicell of L. ciliata
spreading among the zocecial tubes, and these two forms seem so
closely allied that I am in doubt as to the basis of their distinction.

In some specimens the zocecia are slightly ridged, as in L. grig-
nonensis. The spines from the zocecial tubes are not constant,
sometimes occurring in great abundance, in others there are very
few or they may be entirely absent. Both in this and L. pristis,
MacG., there is a semitransparent closure with a dull opaque
disk in the centre, perforated in the middle. It seems probable
that L. pristis and L. echinata are only the simple and confluent
colonies of the same thing.

Loe. Victoria; Tristan da Cunha (‘ Challenger’).

284 ON OVICELLS OF SOME LICHENOPORE.

LICHENOPORA VICTORIENSIS, nom. nov. (Pl. XV. fig. 4.)

Discoporella reticulata, MacGillivray, Trans. Roy. Soc. Vict. vol. xx.
p. 126, fig. 1.

There is already Lichenopora reticulata, since Hagenow described
a fossil as Defrancia reticulata (Bry. der Maast. Kreide, p. 43,
pl. iv. fig. 8, non 4). Marsson has already shown that the descrip-
tion refers to fig. 8 and not to 4. It is possible that this may
have been described under another name, but this is not certain ;
and I certainly cannot agree with Dr. Pergens, who would call it
diadema, uniting the Defrancia disticha, Hag., D. diadema, Gold.,
and D. reticulata, Hag.

As Mr. MacGillivray describes it from Victoria, I would suggest
the name L. victoriensis.

This is one of those species in which without the ovicellular
ducts it would be difficult to say whether there was an oyvicell or
not. The central area is covered by the ovicell, the covering of
which is formed by large raised trabecule, within which there is
a smaller reticulation formed by similar trabecule. There are
spines with knobs from the walls of the trabecule, similar to those
I described in Lichenopora grignonensis (Ann. Mag. Nat. Hist.
ser. 5, vol. xx. p. 262, pl. vii. fig. 4). In the present case I have
only figured a few of these spines, not to complicate the figure
too much. The spaces between the trabecule are filled in with
a perforated cover. Aperture of zocecial tubes about 0-06 mm.
wide.

Loc. Victoria (MacG.); Port Stephens, 5-6 fath., New South
Wales.

DIscoTUBIGERA? LINEATA, MacG. (PI. XV. fig. 5.)

Diastopora lineata, MacGillivray, Trans. Roy. Soc. Vict. vol. xxi. p. 96,
pl. iii. fig. 1. :

Liripora lineata, MacG. Cat. Mar. Polyzoa of Vict., Roy. Soc. Vict.
1887, p. 32.

Discotubigera lineata, Waters, Ann. Mag. Nat. Hist. ser. 5, vol. xx.
p- 260, pl. vi. fig. 24.

I have already (loc. cit.) alluded to the ovicell, but it has never
been figured, and therefore I add a figure of an ovicell occurring
near the border of the zoarium. It will be seen that it is of a
type quite different from that of Lichenopora; but as yet we are iu
ignorance concerning the ovicells of Discotubigera and its allies.

Linn. Soc. Journ. Zoou. Vou XX. Pi. 16d.

; A.W. Waters 7

* #

7 Lb Ge

es,

Mintern Bros. ump.

Michael bth.

A.W. Waters del.

LICHENOPORE

OVICE LIS OF

ON GLYCIPHAGUS DOMESTICUS AND G. SPINIPES. 285

Licuenorvora Hoxtpsworrtuil, Busk. (PI. XV. figs. 7 & 8.)

Discoporella Holdsworthii, Busk, Cat. Mar. Polyzoa, pt. ii.p. 33,
pl. xxx. fig. 4.

Lichenopora Holdsworth, Waters, Quart. Journ. Geol. Soc. vol. xlin.
p. 347 ; Ann. Mag. Nat. Hist. ser. 5, vol. xx. p. 261.

I have already (Quart. Journ. Geol. Soc. xliii. p. 347, and else-
where) alluded to the “ rays”’ in the zoecial tubes and cancelli,
and to Mr. Busk perhaps referring to these when he speaks of
stellate pores

The ovicell occurs as a central inflation with large roundish
depressions closed with a perforated pellicle. It spreads in
between the radii in somewhat the same manner as in L. californica
though not so marked on the surface. I have only seen the one
ovicell; and if this was the only specimen of L. Holdsworthir
known, I doubt if it would be possible without breaking up the
specimen to decide whether there was an ovicell.

Loc. Living: Ceylon; Victoria; ‘ Challenger’ station 142 (S.
Africa). Foss: Waipukurau (New Zealand).

EXPLANATION OF PLATE XV.

Fig. 1. Lichenopora californica, Busk. Showing ovicell extending between the
radii. From Port Western (Victoria). x 12.

2. Cancellus of L. echinata showing denticles. x 85.

3, Lichenopora echinata,MacG. Shows to the left the lower surface of the
ovicell formed by reticulated cells covered with a pellicle. The upper
surface is pitted. x 20.

4. Lichenopora victoriensis, Shows ovicell and two ovicellular ducts. From
Port Stephens, 5-6 fath. (New South Wales). x 25.

5, Discotubigera lineata, MacG. Shows the ovicell at the border of the
colony, and also the ovicellular duct. From Port Jackson (New South
Wales). x 25.

6. Lichenopora echinata. Shows the formation of the ovicell which starts
from several points to meet over the centre. From Port Phillip. x 12.

7. Cancellus of L. Holdsworthvi, showing rays. x 85.

8. Lichenopora Holdsworthii, B., with ovicell. From Port Phillip. x 25.

Researches into the Life-histories of Glyciphagus domesticus and
G. spinipes. By A. D. Micnast, F.L.S., F.Z.8., F.R.MLS.
[Read 3rd May, 1888.]
(Pirate XVI.)
For over three years last passed I have been, as opportunity

offered, endeavouring to elucidate certain obscure points which
I had observed in the post-embryonic development of the common

286 MR. A. D. MICHAEL ON THE LIFE-HISTORIES

Glyciphagi, the specific names of which are mentioned in the
title of this paper; it is the progress and results of those
investigations which I now propose to record.

The inquiry has involved a considerable amount of care and
labour, not only from the inherent difficulties of isolating, rearing,
and watching these minute creatures, but also because it has
uecessitated very numerous dissections of soft hyaline Acari,
some of them less than the fifth of a millimetre in total length,
and of the cast skins of these organisms.

Glyciphagus is a genus of atracheate Acari, belonging to the
family ‘I'yroglypbide, but which is distinguished from the genus
Tyrogiyphus, or cheese-mites, by, inter alia, the rough cuticle,
covered with granulations or vermiform markings, the pec-
tinated or plumose hairs and long, slender tarsi of Glyciphagi,
as opposed to the polished cuticle, setiform hairs, and usually
shorter tarsi of Zyroglyphi, and more especially by the females
of Glyciphagus possessing a central tubular projection from the
posterior margin which is absent from those of Tyroglyphus :
some years ago I showed this to be a bursa copulatrix, a fact
which, I believe, is now generally admitted ; the bursa doubtless
exists in Lyroglyphus, as indeed Dr. Nalepa has proved in one
species, but it does not form an exterior projection.

In order that this paper may be understood it is necessary to
state, as shortly as possible, what a Hypopus is; but as, in a
former paper read before this Society*, I entered fully into the
question, I do not propose to detail the various opinions held
by different authors on the subject, nor the investigations which
led me to the results given, except so far as is absolutely requisite
in order that this paper may be intelligible ; I shall simply state
conclusions, referring to that paper for all proofs and other in-
formation on the subject. Hypopi are minute Acari provided
with a smooth, chitinous carapace, which conceals the whole, or
almost the whole, of the creature; they are somewhat arched on
the back, but still considerably compressed dorso-ventrally ; the
mouth-organs are rudimentary and the posterior pair of legs are
terminated by hairs, not claws. /ypopi are most commonly found
adhering to insects, miriapods, &c., and they have been ultimately
shown to be astage in the life-history of some Acari of the genus
Tyroglyphus and oue or two allied genera, although they are

* «The Hypopus question,” Journ. Linn, Soc., Zool. vol. xvii. (1884) pp.
371-394,

OF GLYCIPHAGUS DOMESTICUS AND G. SPINIPES. 287

very unlike all other stages of Tyroglyphi in appearance. The
stage does not occur in the life-history of every individual of a
species, but in a comparatively limited number of instances.
Méenin considered that the stage occurred when, from drought
or unfavourable circumstances, the colony was in danger of perish-
ing, and that the nymphs of Z'yroglyphus had the power of turning
into Hypopi and so remaining until either the circumstances had
again become favourable in the place where they were, or until
they had emigrated into more suitable quarters—the Hypopus
being better able to endure drought &c., and more capable of
adhering to insects and thus escaping than the other stages of
thecreature. Ithink that I proved by the investigations detailed
in the paper above referred to that this was an error, and that
unfavourable circumstances did not affect the question—the Hy-
popial stage being a provision of nature to ensure the distribution
of the species, occurring irrespective of adverse conditions, and
that it occupied the period between two ecdyses in the life-history,
commencing in the species which I was then studying at the
second nymphal ecdysis.

Glyciphagus is not one of the genera in which a Hypopial stage
has ever hitherto been found to exist, except in the case of
G. Crameri described by me in 1886 * ; and this, as pointed out in
the paper in which I described if, is an extreme species of the
genus, which would hardly be included in the genus were it not
for the connecting-links, and would undoubtedly fall within Hal-
ler’s genus “ Dermacarus”’ if that genus were retained. Of the
two species named in the title, one, G. spinipes, was first recorded
by Koch; and, so far as we know at present, there is not any
difficulty in identifying it, and but little synonymy connected
with it. G. domesticus can, however, hardly be said to be in this
happy position: the synonymy of this species would probably be
voluminous; but this is not the place to investigate it, beyond
what is necessary for identifying the species that I have been
experimenting upon. The species was originated by de Geer,
who called it Acarus domesticus, the division of Zyroglyphus from
Glyciphagus not having been effected at that time; one result of
this has been that numerous writers have identified the creature
with the common cheese-mite (Zyroglyphus siro, Linn.). Now,
however, that the distinction between the two genera is understood,
it is abundantly clear that de Geer’s species was a Glyciphagus ;

* “Upon the Life-history of an Acarus, one stage whereof is known as Labi-
dophorus talpe, Kramer, &e.,” Journ. R. Mic. Soc. 1886, pp. 377-390.

288 MR. A. D. MICHAEL ON THE LIFE-HISTORIES

it appeared in 1778. In 1841 Gervais published a very short
description and very imperfect drawing of a species which he
called Glyciphagus cursor. In 1867 Robin and Fumose pub-
lished a very carefully prepared article in Robin’s ‘Journal de
Anatomie et de la Physiologie,’ in which they, probably correctly,
identified de Geer’s and Gervais’s species ; but, for some reason
which is not quite apparent, they retained Gervais’s name, not
de Geer’s. M. Mégnin, in the observations mentioned below as
having been made by him, has followed Robin and Fumose in
calling the species G. cursor: unfortunately neither of these able
authors gives a figure of their G. cursor; had Robin and Fumose
added such a drawing as they furnish of G. spinipes no doubt
could arise as to what their species is. A good figure of what is
apparently the male of G. domesticus is given by Berlese (Acari
Ital. fase. xiv. no. 3); it is accompanied by a short description,
which, however, is supplemented by the notes to the same work
(fase. i. pp. 9,10). This author, however, does not say whether
he considers it to be identical with G. cursor, and the points
which he gives for identifying his species do not admit of com-
parison with those given by Robin and Fumose for G. cursor. I
imagine the two species to be identical. Under these circum-
stances I have thought it best to give a figure of the female of
the species I have been dealing with, to facilitate identification ;
it is, I think, the G. domesticus of de Geer and Berlese, and, so
far as I can judge at present, it is also the G. cursor of the other
authors above named.

It would be out of place here to give any formal description of
such well-known creatures ; but as these two species, G. spinipes
and G. domesticus, are commonly found together, and are not
easily distinguished at first, although a little practice enables the
student to do so with facility and certainty, it may be useful to
state some of the principal characters by which they may be dis-
tinguished ; these are :—Firstly, that the tarsi of G. spinipes are
thickly clothed with very fine short hairs (Pl. XVI. fig. 12), but do
not bear any hairs much longer than the general average; the tarsi
of G. domesticus are smooth, but have a few straight hairs or
spines much longer than those of G. spinipes (fig. 5). The hairs
on the tarsi of G. spinipes are best seen in dry specimens using
an amplification of from 150 to 200 diameters ; they are not so
easily seen with lower powers nor in specimens mounted in bal-
sam, or even in fluid. Secondly, the tarsi of G. domesticus are

OF GLYCIPHAGUS DOMESTICUS AND G. SPINIPES. 289

considerably longer than those of G. spinipes. Thirdly, @.
spinipes has an obtusely conical shape anteriorly, and is suddenly
contracted between the second and third pair of legs; G. domes-
ticus has this form indicated but very slightly, being nearly as
wide posteriorly as in the middle of the body ; the difference can
only be well seen in living specimens. Fourthly, the bursa
copulatrix of the female projects considerably more in G. domes-
ticus (fig. 7) than in G. spinipes (fig. 15). Fifthly, G. domesticus
is a trifle the larger, but this cannot be relied upon. Sixthly,
the hairs of G. spinipes are more deeply serrated and stronger
than those of G. domesticus. Seventhly, the tibie of the first and
second pairs of legs in G. spinipes each bear two large and thick
curved, strongly-serrated hairs ; these are much less developed in
G. domesticus. Highthly, the second joints of the third pair of
legs in G. spinipes each bear on the outside a peculiar hair or
scale (fig. 14), shaped like a husk of corn, and placed with the
concavity inward; its outer surface is thickly clothed with fine
hairs or processes: this structure is most developed in the female ;
it has not, I think, been mentioned by previous writers ;
it is not found in G. domesticus. Ninthly, G. spinipes when
placed on its back, or otherwise disturbed, has a rapid struggling
or fighting motion with its first and second pairs of legs, a little
of which is seen in its ordinary walk ; this style of movement is
very characteristic of the species.

The Tyroglyphide, like most of the other Acari, have an inert
period previous to each ecdysis ; during this time the creature is
motionless and apparently dead, except that it does not shrivel
up; it is needless to say that a proper examination will disclose
that the development of the creature in its next stage is pro-
gressing inside the dead cuticle.

GLYCIPHAGUS DOMESTICUS.

During the month of May, 1885, when I was examining some
material which had been sent to me and which was swarming
with G. domesticus in various stages, I observed that there were
some which could only he classed as inert nymphs, but which had
not quite the ordinary appearance of the creature in that stage ;
the cuticle was more opaque, and seemed thicker and whiter, the
back was more arched, the empty skin of the legs was more
apt to be rubbed off, as if the inert stage had lasted longer than
usual, giving acase-like appearance; and I propose in this paper

290 MR. A. D. MICHAEL ON THE LIFE-HISTORIES

to use the expression “cases” to signify this phase in the life-
history ; it must, however, be understood that each case is only a
nymphal skin, but it is a nymphal skin under special conditions.
The finely granular marking or wrinkling of the cuticle of the
species was very apparent, and when a portion was examined with
a sufficient power this was resolved into the labyrinthine and vermi-
form wrinkling of which an illustration is given (Pl. XVI. fig. 3).
It was also noticeable that the contents of each case were more
drawn toward the anterior end than is usual in the nymph during
ecdysis. Of course the first step was to ascertain that these
cases were not simply dead creatures; for this purpose, in the
latter part of May 1885 I isolated three in a separate glass cell,
placing them under what experience had taught me would pro-
bably be favourable conditions of existence. On the 10th June,
1885, three immature Glyciphagus domesticus emerged from these
cases. The cases did not split irregularly like an ordinary
nymphal skin, but opened by the posterior end of the case,
which had been concave, being pushed out so as to become rather
convex, and separating from the lateral and ventral portions of
the case, while it remained attached to the dorsal ; thus when the
creature within had escaped, the posterior eud formed a sort of
lid attached to the dorsal surface (fig. 4). This I subsequently
found was the usual, although not the absolutely invariable, mode
of opening. I did not then examine the precise stage that the
creatures were in, but I noticed that the cases, although open, did
not appear to be entirely empty (I subsequently found that what
they contained was a cast skin).

On the 10th June, 1885, I placed thirteen more cases in a
second cell, and submitted them, as I had hoped, to conditions
similar to the first ; but either from some defect in the arrange-
ment of the cell, or from unfavourable weather, these specimens
gradually died, so that toward the end of July there were not any
alive. On the 30th July, 1885, I again isolated several cases in a
separate cell. In a week I found that two of them were open at
the posterior end and that there were two nymphs of Glyciphagus
domesticus in the cell; more subsequently emerged from the
other cases. I again observed that the cases from which they
had come did not appear to be absolutely empty. I now took
three of the nymphs which had emerged from the cases and
placed them in a separate cell; in about a week they had all
become inert; a few days later an adult female of G. domesticus

OF GLYCIPHAGUS DOMESTICUS AND G. SPINIPES. 291

escaped out of the cuticle of one of these inert nymphs, the
others soon followed. The cast skin from which the adult
emerged was thin and fine, very different in appearance from that
of the cases. One point at least had now been established, VIZ.
that the cases were a penultimate nymphal stage—i. e. that the
nymph which emerged from the case became adult at its first
ecdysis.

While these observations were in progress I also endeavoured
to obtain a knowledge of the matter by dissecting the cases and
their contents; but as, in order to avoid any chance of error, I
have lately repeated these dissections on a large number of speci-
mens, I will describe the results of both together in order to avoid
repetition.

_ At this time I was forced by other engagements to abandon
the investigation temporarily ; and I did not make any further
observations on the same species until the present year (1888).
On January 25, 1888, I placed in a cell four cases taken from
fresh material which I had obtained, but which contained very
few cases, some of which I reserved for dissection ; but in spite
of my providing them with gentle warmth and slight moisture,
such as would apparently form the most favourable conditions,
these cases still (May 3) remain in the same condition as when
placed in the cell*. On April 9, having found that a great many
cases had now formed in the same material from which the last
had been taken, I placed a number of these in three separate
cells. On 21st April I found an inert nymph of G. domesticus in
one of these cells which had escaped from a case, and must have
emerged and become inert since the cell had been last examined,
which happened to be two days; longerthan usual. On the 26th
April the adult emerged, and a second nymph had emerged and
become inert prior to the final ecdysis. On the 1st May an adult
female of G. domesticus emerged from this last-named nymph.
I carefully examined it and made certain of the species.

Between the commencement of January and the end of April
1888, 1 dissected a large number of these cases of G. domesticus ;
the results were entirely confirmatory of those which I had
obtained in 1885—viz., that inside each full case, and almost
filling up the anterior portion of it, but not the legs, which were
empty, was a protoplasmic mass which had a transparent, colour-
less, and almost structureless cuticle. This mass had a rounded

* They subsequently emerged on 15th July.
LINN. JOURN.— ZOOLOGY, VOL. xx. 23

292 MR. A. D. MICHAEL ON THE LIFE-HISTORIES

posterior and a pointed anterior end; was compressed dorso-
ventrally, particularly at the posterior margin; and had a more
or less plain sulcation round it, as though dividing cephalothorax
from abdomen (in some specimens this was very marked). The
mass had the general form of a Hypopus, but there was not, in any
instance, any trace of legs, mouth, or other external organs. In
many instances the protoplasm appeared to be divided into large
cells, ike an egg in an early stage of segmentation ; in others the
cell-division appeared to have gone further, the cells being much
smaller and finer, particularly in the posterior portion of the
creature, but some of the larger cells remaining; in others, pre-
sumably more advanced, the finer granulation was more uniform.
The mass was always motionless, but in one instance I did
find a living nymph within the case instead of the inert mass ;
this of course was ready to emerge. The cases from which the
occupant bad emerged almost invariably contained the cast
cuticle of the protoplasmic mass, which cuticle did not show a
trace of legs, mouth, or any other organs.

GLYCIPHAGUS SPINIPES.

At the end of 1885 I was at a farm-house for a short time, and
thought it a favourable opportunity to renew the investigation.
I found in the chaff-house, in the dust and chaff, and also attached
to the walls and beams, a number of cases which I at first
supposed to be similar to those I had before dealt with: I soon,
however, found that these were even more opaque than the former
specimens, and that they were coarsely reticulated instead of
being finely vermiform in markings; the empty cases also opened
differently, the posterior cuticle breaking away from the dorsal
and lateral, and remaining attached to the ventral, so that the
posterior end opened downward instead of upward, and was more
torn; the hinder part of the dorsal cuticle also was usually split
along the median line and the two sides somewhat separated. It
was therefore probable either that they were in a different stage
from those observed in June and July of the same year, or that
they were under different climatic or other conditions, or that
they belonged to a different species of Acarus. I finally found
that the last explanation was correct, and that these reticulated
cases were those of G. spinipes. Between the 28th December,
1885, and the 1st January, 1886, I found a considerable number

GLYCIPHAGUS DOMESTICUS AND G. SPINIPES. 298

of these cases. I placed some of them in a cell as before and
dissected others; these,in many instances, contained not the inert
legless mass found in the cases of G. domesticus, but a distinctly
formed living Hypopus, which had not assumed the usual brown
chitinous colour, and could not be called active, but still was fully
formed, and provided with short and stumpy, but thoroughly-deve-
loped legs, which it could move about, although only in a feeble
manner ; it was not capable of walking (ordinary Hypopi are very
active), and it was evidently not in a condition fitted for existence as
a free-living creature; but yet it was undoubtedly alive and fully
formed ; its total length was about ‘19 millim., its greatest width
about ‘15 miliim. Each tarsus of the first three pairs had an
exceedingly long, very slightly curved, blunt claw; the tarsi of
the fourth pair were devoid of claws and hairs. A figure of this
Hypopus, carefully drawn from a specimen dissected out of a case
in January 1888, is givenin Pl. XVI. fig. 9. The majority of the
Hypopi dissected out of the cases were not capable of any move-
ment ; it was only a few which were able to move their legs. I now
searched the chaff and material, and the dust &c. swept from the
wallsand beams of the chaff-house ,very carefully in hopes of finding
active Hypopi which had emerged and were capable of walking
about, but neither on this nor on any other occasion have I been able
to discover anything of the kind ; but I did find one or two inactive
Hypopi not capable of any movement, which seemed as if, from
some accident, the cases had been broken away from them. On 2nd
January, 1886, I took one of these, and one of the Hypopi dissected
out of a case and which could move its legs, and put them in a cell
by themselves. I watched them at frequent intervals, but they
did not die or shrivel up. On the 15th January I missed one of
the Hypopt; but close to where it ought to have been I saw a
nymph of Glyciphaqus spinipes, which had evidently just emerged
and had some thin membrane attached to it; on detaching this
membrane, and examining it with a higher power, I found that it
was the cast skin of the Hypopus. I put both the nymph and tne
east skin in glycerine for permanent preservation. A few days
later I dissected four more of the cases and took out four Hypopi
and placed them on a small piece of dried leaf in a separate cell;
the next day I found that two had collapsed, probably having
been injured during the dissection of their cases. The following
day I observed that one of the remaining two had moved, and on
touching it with a fine hair I saw that it was alive and able to

294 MR. A- D. MICHAEL ON THE 1L1FE-HISTORIES

move about very slowly; but the day after it had collapsed and
was dead. There was now only one Hypopus left; this did not
move nor show any sign of lite; but, on the other hand, it did
not collapse, and in this condition it remained for nearly four
months. I watched it every day without seeing any change
until the 24th May, 1886, on which day a healthy nymph of G.
spinipes emerged from the Hypopial skin.

While this experiment was progressing I had also watched the
cell in which I had placed the cases: one nymph of G. spinipes
had appeared in the cell and one case was open at the posterior
end; otherwise there was not any change. But on the 26th May
I found two more cases open and two Hypopi bad apparently
crawled out of them; on touching these Hypopi they appeared
inert and incapable of motion, but a few days after nymphs of
G. spinipes emerged from them.

In October 1887 I again obtained more material from the
same chaff-house, and also from another farm some fifty miles
distant from the first ; this latter contained G. domesticus. In both
samples the Glyciphagi were numerous and the larve and nymphs
were abundant, but I could not find any cases or any trace of
Hypopi in either.

In January 1888 I once more took up the investigation: I
obtained material and sweepings from the same chaff-house and
again found the reticulated cases of G. spinipes; most of them
were open at the posterior end, the occupants having emerged.
I put seven of the full cases in a separate cell; m one of these
cases I could easily distinguish a Hypopus moving its legs. I
also found one which had apparently come out of the case, and
which was capable of the same amount of motion; it was from
this specimen that fig. 9 was drawn. On February 6 I found
that one of the seven cases was open and that a nymph had
emerged from it. I isolated this in another cell, which I will
call cell 5. On February 8 another nymph emerged from a second
of the seven cases. I placed this nymph also in cell 5. On
February 9 an adult G. spinipes emerged from one of the nymphs
in cell 5, the second nymph also was inert. On February 26 an
adult of the same species emerged from the remaining inert nymph.

From this time up to the end of April, when I closed the
observations, nymphs of G. spinipes continued to emerge at inter-
vals from the remainder of the seven cases, and from other cases
which I had put in different cells, and adults continued to emerge

OF GLYCIPHAGUS DOMESTICUS AND G. SPINIPES. 295

from the nymphs, so that there could not be any doubt of the
species, or of the process being a regular one.

I also in January 1888, and again in April of the same year,
dissected numerous reticulated cases from which the nymph had
emerged; in almost every instance I found within the case an
unmistakable cast Hypopial skin having perfectly distinct legs.

CONCURRENT INVESTIGATIONS BY M. Méanrn.

It appears that while I was engaged on these observations
M. Mégnin was also investigating some points relative to the
Glyciphagi, and, inter alia, almost the same subject as my own *.
We did not either of us know of the other being so occupied.
Mégnin approached the matter from a standpoint different from
mine: he did not find an object which excited his curiosity to
discover its cause; but it struck him that as Tyroglyphus has
a Hypopial stage, so closely allied a genus as Glyciphagus ought
also to have it, and he deliberately set himself to search for
that stage ; but he tells us that he searched in vain; he tells us,
however, that he discovered an equally curious phenomenon, which
he says shows how prodigal nature is in processes for preserving the
lower creatures. He, believing that the change of a nymph of
Tyroglyphus into a Hypopus is caused by unfavourable surround-
ings, states that under similar conditions he found that those of
Glyciphagus became inert, that a liquefication of all the organs took
place “as in a change of skin,” and that the gelatinous substance
collectsin the cavity of the “ thorax’ in the form of a spherical
mass surrounded by a chitinous envelope and thus forming a cyst
very similar to those formed by some Infusoria previous to the
drying up of the water in which they are contained. Mégnin
suggests that in this condition the dried nymphal skins containing
the cysts would be bluwn about by the wind and would thus
finally arrive at some place where the conditions would be favour-
able and would then emerge, and that the species would be thus
distributed.

Mégnin says that his species were G. spinipes and G. cursor,
which latter is, as before stated, presumably the same as G. domes-
ticus ; but he does not distinguish between the life-histories of the
two, nor identify any particular observations with either species.

I do not intend here to discuss the vexed question of whether
the liquefication of the organs of an Acarus during ecdysis igs

* Comptes Rendus, ciii, (1886) pp. 1276-8.

296 MR. A. D. MICHAEL ON THE LIFE-HISTORIES

complete, involving a return of all the organs to an egg-like state,
or whether it is only partial: the former has long been Mégnin’s
view, and he cites Claparede as confirming it; but on reference
to the Swiss author’s writings this does not seem to be by any
means certain. Dr. Nalepa, who has lately investigated the subject
with regard to the Tyroglyphide, is strongly of opinion that it is
only partial*. I was formerly inclined to think, chiefly on
Mégnin’s authority, that it was complete; but more lately, having
paid some personal attention to the subject, I have come to a
conclusion with regard to the Oribatide + more in accordance with
Dr. Nalepa’s views. It is clear that Mégnin has been observing
a process either identical with or very similar to what I have seen
in G. domesticus, but that not having found the more fully de-
veloped Hypopus in G. spinipes he has naturally not recognized
that what he found was really a rudimentary Hypopus—the dif-
ference of the spherical form of the inert mass found by him in
his cysts from the more Hypopus-like form of the corresponding
mass found by me in G. domesticus may possibly be accounted
for by his specimens having been examined more shortly after
the change commenced than mine were, or by his observations
having been made on G. cursor, and that species not being really
identical with G. domesticus, although much resembling it, but
having the Hypopial stage even more rudimentary.

It is quite possible that Mégnin’s idea that the inert nymph
when containing the Hypopial form, or, as he calls it, the cyst,
would be blown about by the wind may be correct: I do not see
that it would be more liable to be so blown about during this
than during any other ecdysis, except in so far as this may last
longer; but undoubtedly the creature within, protected by its
Hypopial skin, would bear more exposure and rough treatment
than the Acari would during other ecdyses. It will probably,
however, be gathered from the foregoing pages that I cannot
agree with Mégnin that the change to the Hypopial, or as he
calls it encysted, condition is produced by desiccation or unfavour-
able surroundings. J have not conducted a series of experiments
expressly designed to determine this exact point, as I did with
the Tyroglyphi; but my material was not kept in any desiccated
or unfavourable condition; on the contrary it was naturally kept

* “Die Anatomie der Tyroglyphen,” Sitzungsb. k. Akad. Wien, i. Abth.
Juli-Heft, 1885, p. 151.
+ “British Oribatidw,” Ray Society, vol. ii. (London, 1888), p. 604.

OF GLYCIPHAGUS DOMESTICUS AND G. SPINIPES. 297

under the most favourable conditions I could devise, and the
Glyciphagi throve admirably; but the cases containing Még-
nin’s cysts were formed rapidly and numerously in it; from
which, aud from the general experience acquired during the
somewhat lengthy observations above described, I feel assured
that with the Glyciphagi, where the Hypopial stage is more or
less rudimentary, as with the Tyroglyphi, where it is an active
and functional one, the change to this stage is normal, although
not existing in the life-history of every individual, and is not
induced by desiccation or other unfavourable conditions ; but, on
the contrary, proceeds best and most rapidly when all conditions
are favourable. I do not deny that a creature may remain longer
in the Hypopial condition after it has been formed when the
surroundings are more suitable to that stage than to the ordi-
nary nymphal state, than it will when the converse is the case ;
I think this not improbable, although I have not seen any
evidence to prove that such is the fact.

CONCLUSIONS.

The results of the investigations detailed above may be sum-
marized as follows :—

1. There is a Hypopial stage in the life-history of Glyciphagus
just as there is in that of Tyroglyphus.

2. That this Hypopial stage is far less developed in Glyciphagus
than in Tyroglyphus, and is not, so far as is known at present, an
active stage.

3. That we do not at present know whether it occurs in all
species, but it certainly does not occur in the life of every indi-
vidual of a species.

4. That the stage is not the result of desiccation or other
unfavourable circumstances, but occurs as often under favourable
conditions.

5. That the stage, in the species investigated, occupies the period
between the penultimate ecdysis and that immediately previous.

6. That in G. spinipes the Hypopus is fully formed and capable
of moving its legs, but not of walking or other active movement ;
that it never becomes hard, or of the dark colour of the ordinary
chitin of active Hypop?. That, as a rule, it does not even leave
the skin of the young nymph in which it is formed; but that
the more adult nymph is formed within the Hypopus, and emerges

298 ON GLYCIPHAGUS DOMESTICUS AND G. SPINIPES.

from it, while the Hypopus is still within the young nymphal
skin; so that the Hypopus is not ever seen, except that, in a
few instances, it may crawl just outside the young nymphal skin
when the more advanced nymph is likely soon to emerge.

7. That in G. domesticus the Hypopial stage is even more rudi-
mentary ; what represents the Hypopus retaining only the general
form of that creature, but being without legs or other external
organs, and that it never emerges from the young nymphal skin.

8. That in both species the young nymphal skin within which
the Hypopus is developed thickens and forms a “ case”’ different
in appearance from the skin during ordinary ecdysis.

EXPLANATION OF PLATE XVI.

Fig. 1. Glyciphagus domesticus: adult female, underside; x 95.

2. Ditto: ‘‘case” containing the inert mass (very rudimentary Hypopial
condition); x 150.

3. Ditto: a portion of the cuticle of the “ case,” to show the vermiform
markings; xX 200.

4. Ditto: posterior end of “case” from which the nymph in its last stage
has emerged ; side view, to show the mode of opening.

5. Ditto: fourth left tarsus of adult female from above; x 200. At the
side is a line showing the extreme width of the abdomen; same
amplification.

6. Ditto: claw and caruncle from below, drawn from the living foot when
pressed by the creature against glass ; X 500.

7. Ditto: bursa copulatrix; x 200.

8. Glyciphagus spinipes : “ case” containing the Hypopus; x 150.

9. Ditto: Hypopus dissected out of the ‘ case”; x 180.

10. Ditto: a portion of the cuticle of the ‘‘case,” to show the reticulated

markings; x 200.

11. Ditto: posterior end of “case” from which the nymph in its last stage
has emerged ; seen from above, to show the mode of opening.

12. Ditto: fourth left tarsus of adult female from above; x 200. At the
side is a line showing the extreme width of the abdomen; same
amplification.

13. Ditto: part of the third left leg of adult female from the outer side ;
X 250. :

14. Ditto: hair from the second joint of the same leg; x 500.

15. Ditto: bursa copulatrix.

Linw. Soc, J OURN. Zoan. Vat.. Tee Tee ol

TORIES OF GLYCI PHAGUS DOMESTICUS AND G.SPINIPES.
‘ ik bP
a iy od »

v
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- (

STUDIES OF THE MACROOCHIRES. 299

Studies of the Macrochires, Morphological and otherwise, with
the view of indicating their Relationships and defining
their several Positions in the System. By R. W.SuHuFeEtpr,
M.D., C.M.Z.S8., Captain, Medical Corps, U.S. Army.
(Communicated by W. K. Parker, F.R.S., F.L.8.)

[Read 19th January, 1888. |
(Piates XVII.-XXIV.)

Ir may be remembered by those who are interested in the
structure and classification of birds that I published, in the
* Proceedings’ of the Zoological Society for 1885 (pp. 886-915),
a memoir entitled “ A Contribution to the Comparative Osteology
of the Lrochilide, Caprimulgide, and Cypselide.” 'That memoir
professed to be but little more than a mere introduction to
a subject which I will here enter upon more fully, although the
opinions there set forth are, in the main, substantially those that
I still hold, at least in the case of the Trochilide and Capri-
mulgide. Since the date of that paper, however, I have never
ceased in my endeavour to gather together the necessary material
for this, my second contribution on the subject; and, as will be
seen by the list of specimens in the subjoined Table, these efforts
have met with a very fair measure of success.

In the conclusions at the close of my former paper I contended
that ali the existing Caprimulgine birds of the world’s avifauna
should be grouped in one order, the Carrimusnat. In this
group, no doubt, would fall Myctibiws and Steatornis, and very
probably Podargus and Psalurus. Further, I proposed that the
Humming-birds should constitute another order, to be known as
the Trocui11. J made no final determinations in regard to the
Swifts, beyond that they should be separated from the Trochili ;
but these birds will be carefully studied in the present memoir,
and my opinions in regard to them stated in the conclusions
which close it.

Finally, I had something to say about certain apparent resem-
blances between the bones in the roof of the mouth of a Trogon
and the corresponding structures in a Humming-bird. But my
remarks were only drawn from a paper by W. A. Forbes published
in the ‘ Proceedings ’ of the Zoological Society three or four years

LINN. JOURN.—ZOOLOGY, VOL. XX. 24:

300 DR. R. W. SHUFELDT’S MORPHOLOGIOAL

previous to my quoting them*, and at the time I had not
had an opportunity of personally examining the skeleton of a
Trogon. In the present paper, however, the case will, in this
respect, be different ; for, thanks to the kindness of Dr. P. L.
Sclater, I have been enabled to study in this connection the
skeletons of two different species of Trogon, which he has
obligingly lent me for the purpose. For other material I am
under obligations to a number of friends, to whom I here desire
to express my sincere thanks; and I believe it will be found
that in the proper column of the Table below I make due
acknowledgments, by entering the names of the several donors
opposite the specimens they have been so good as to send me.
Indeed, had it not been for their kind and ready assistance, it
would have been impossible for me to have completed the pre-
sent work. Such material as I have been enabled to collect
myself is also set forth in the Table in question. My thanks
are further due to the Editors of ‘The Auk’ and of ‘ Forest
and Stream,’ for kindly inserting for me requests for specimens
of birds in alcohol to be used in the present connection.

Glancing over this list of material, it will be observed that, so
far as the ordinary forms of the American Caprimulgine birds
are concerned, it admits of giving a full account of their structure.
The skeleton in these also may be conveniently compared with
the skeleton in the two species of Trogons likewise represented ;
aud these latter with other types presented in the Table, as well
as with such a skeleton as is presented in Geococcyx cali-
fornianus, which I have elsewhere studied (Journ. Anat. and
Phys. Lond. vol. xx. 1886, pp. 244-266) and published an
account of its characters.

Similarly, we find the North-American CypsrEtrt very well
represented, the only important form not found among my mate-
rial being Cypseloides niger, and all my efforts to secure specimens
in alcohol of this interesting Swift have utterly failed +. In my
first contribution to the anatomy of these birds (P.Z.S. 1885,
p. 886), I advanced the opinion that they were but profoundly
modified Swallows, and should not be grouped with the Trochili
in our classification of birds. We now have the opportunity

* Forbes, W. A., “ Note on the Structure of the Palate in the Trogons
(Trogonide),” P.Z.8. 1881, p. 836.

+ I have since received alcoholic specimens of this form from my friend
Prof, A. Newton, F.R.S., who kindly procured them from Jamaica for me.

STUDIES OF THE MACROCHIRES. 301

of ascertaining how this opinion will stand the test of more
extensive researches into their structure, with the aid of a far
better supply of material than I was enabled to handle upon
the first occasion.

The Table also presents a very fine collection of skeletons
and specimens in alcohol of the Trochili; and the structure of
these, as I say, will in the following pages be thoroughly com-
pared with the corresponding structures to be found in the Swifts.

It is further my intention to compare the Swifts thoroughly
with the Swallows; and to this end I find that I have examples
of every genus of the latter birds, as they are represented in the
North-American or, rather, United States avifauna. Up to the
present time my efforts to secure specimens in alcohol of such
a form as Hemiprogne zonaris have not succeeded; but Mr. C. B.
Cory has kindly written for me to some collectors in the West
Indies, and I have taken the liberty to apply for some of these
birds to the governmental authorities in Jamaica. Sufficient
time has scarcely elapsed for me to have received replies, but I
am under the impression that when this type comes to be com-
pared with more typical Swifts, some light will be thrown upon
the present subject. ,

Finally, it is my intention in this memoir to review some of
the structural and other characteristics of the Passeres; not that
the majority of the ordinary ones are not already known to us,
but rather to have them arranged systematically at hand, for
comparison in the present connection. I have chosen for this
purpose a specimen of Ampelis garrulus, being influenced in my
choice more particularly by the fact that, although it happens
to be Passerine, it is not strictly and typically so: moreover,
some ornithologists apparently recognize in it certain Swallow-
like characters, more especially those which have a certain resem-
blance to such a bird as Progne subis, which it appears to approach
in the form of the bill. Others, and much earlier authorities, have
entertained the notion that the Waxwings belong rather to the
Clamatores, being probably related to the Zyrannide, and should
be placed near them. So that, on the whole, I trust that, in addi-
tion to meeting other ends in view, a glance at the structure of
this strictly American representative of the Ampeline will not
come amiss.

The order in which I propose to present the material to be
examined will be:—first, a sketch of the morphology of

24,*

DR. R. W. SHUFELDT’S MORPHOLOGICAL

302

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303

STUDIES OF THE MACROCHIRES.

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DR. R. W. SHUFELDT’S MORPHOLOGICAL

304

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305

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306 DR. R. W. SHUFELDT’S MORPHOLOGICAL

Ampelis cedrorum; secondly, a study of the osteology of Trogon
mexicanus and T. puella; thirdly, an examination of the struc-
ture of a number of the Caprimutel; fonrthly, an investigation
of the anatomy of the North-American Hirundinide; fitthly,
similar inquiries into the morphology of certain Cypsrii and
Trocui1t, including extensive comparisons with the facts brought
out in the first sections of the paper; and, lastly, a section
devoted to my final Comparisons and Conclusions.

Tur MorpHoLoGy OF AMPELIS CEDRORUM.

From an external examination of the subject, we find that the
following characters are presented :—

1. The soft feathers on the top of the head unite to form a
conspicuous crest.

2. The bill is broad at its base, rather short, and vertically
compressed; while both mandibles show a distinct notch at the
sides near the apices.

3. The cleft of the gape extends nearly as far back as the
anterior arc of the eyelids.

4, The nostrils are subelliptical and scaled.

5. The wings are ample, pointed by the 8rd primary of the 10
composing one of them, the 1st primary being rudimentary.

6. Peculiar wax-like prolongations of the shaft are found in
certain of the wing-feathers, and in some individuals in the
tail-feathers.

7. Tail very slightly rounded, and composed of 12 rectrices.

8. Lateral plates of tarsus subdivided, with the anterior portion
of this envelope composed of six distinct scutes.

9. Feet moderately strong, and characterized by having the
basal phalanges of middle and outer toes more or less united.

10. In form, the body is somewhat robust and full-chested.

To this we may add that the Cedar-bird builds its nest upon
trees, and lays spotted eggs, and that the young have a different
coloration of plumage from their parents, which, in this respect,
are alike.

On plucking the specimen the following additional characters
are revealed :—

11. The pterylosis agrees almost exactly in pattern with the
pteryloses of the majority of Passerine birds. A. cedrorum has,
however, a lateral and narrow tract running longitudinally down

STUDIES OF THE MACROCHIRES. 307

each side of the neck; these connect the humeral tracts with
the feathering of the head.

Otherwise both the dorsal and ventral tracts of the Cedar-
bird agree very well with the details of this important character
as seen in such a Passerine form, e. g., as Motacilla alba. This
fact may be better appreciated by comparing my drawings of
the former (Pl. XVII. fig. La & b) with Nitzsch’s figures of the
pterylosis of the latter *.

12. The oil-gland is found to be nude, and this gland has a
form such as is assumed among the great majority of the
Passeres. |

Upon removing the integument, one of the most convenient
anatomical points to be first examined is the method of insertion
of the muscles of the patagium. In the case of a small bird such
as we have in Ampelis, our present subject, I find an easy way to
do this is to seize the elbow of the plucked pectoral limb with
the thumb and index finger of the left hand, in such a manner
that the palmar surface of the index finger is applied to the
under surface of the patagium, and keeps it on the stretch. The
thumb is opposed to this, and firmly holds the elbow-joint, and
no more. We now make an incision, with the scalpel in our
right hand, through the skin, down the line of the triceps muscle,
and another at right angles to it, along the line of the ulna:
then carefully dissecting back the triangular flap of integument
thus outlined, the parts to be examined come nicely into
view.

13. In Ampelis the insertion of the tensor patagii longus and
brevis are typically Passerine in character, as may be seen in the
drawing here presented of these parts, which I made directly from
my dissection (Pl. XVII. fig. 2), and from Prof. Garrod’s excellent
description, in his memoir upon the anatomy of the group?, of this
method of insertion, as we find it in nearly all Passeres.

In A. cedrorum, however, we find another patagial muscle
present in all Passeres which I have examined, which else-
where I have named the dermo-tensor patagit, marking it dt.p. in

* Nitzsch’s ‘Pterylography :’ translated from the German by W. S. Dallas,
and edited by P. L. Sclater, for the Ray Society. London, 1867: Taf. iii.
figs. 1 & 2.

+ Garrod, A. H., ‘*On some Anatomical Characters which bear upon the
Major Divisions of the Passerine Birds,” P.Z.8. 1876, pp. 506-519 (read
June 6th, 1876).

308 DR. R. W. SHUFELDT’S MORPHOLOGICAL

Pl. XVII. fig. 2 of the present memoir. This muscle arises from
the inner surface of the skin at the lower part of the neck,
its fibres converging as they pass towards the shoulder to
terminate in a small and delicate tendon, which accompanies the
tendon of the tensor patagii longus in the free marginal fold
of the patagium, and merges with it about halfway between the
humerus and carpus. When I come to discuss this musele in
the Hirundinide I will enter more fully upon it, its relations,
and the birds wherein I have thus far detected its presence.

14. The musculature of the lower larynx of Ampelis cedrorum
is of a very perfect Acromyodian type; I distinctly make out
Jive pairs of intrinsic muscles inserted, as usual in Passeres, into
the ends of the three upper bronchial semi-rings ; and, in addition
to these, there is a well-developed pair of sterno-tracheales.

These lower laryngeal muscles are here not only firm and
fleshy, but easily individualized—a feat best accomplished with
a pair of dissecting-needles, under the 2-inch objective of a good
microscope.

Both in structure and position, then, the syrinx of Ampelis is
of a typical Passerine type.

15. Turning next to the pectoral muscles, I find all three—the
pectoralis major, the p. secundus, and p. tertiws—to be present
and fully developed. They also have their usual origins and
insertions.

16. In examining the muscles cf the thigh, with their vessels
and nerves, I find that the ambiens and accessory femoro-caudal
are both absent, while the femoro-caudal, the semimembranosus,
the semitendinosus, and accessory semitendinesus are all present.
The main artery is the sciatic; the main nerve the sciatic nerve;
and the main vein the femoral.

In all of these particulars Ampelis cedrorwm agrees with the
highest types of Passerine birds.

17. At the lower third and at the back of the tarso-metatarsus
I find in both feet that the tendons of the flexor longus hallucis
and the flexor profundus digitorum are not connected by a fibrous
vinculum. This is also another Passerine character.

18. In examining the heart and great vessels, I find but one
carotid artery—the left one—passing up in front of the vertebrae
in the neck.

19. I find the nostrils in Ampelis but very feebly partitioned
from each other by a median membro-cartilaginous septum.

STUDIES OF THE MACROCHIRES. 309

20. The alimentary canal agrees in general with the Passerine
birds, and a small pair of ceca coli are present.

The Skeleton of Ampelis.

To any one who has examined series of skeletons of Passerine
birds, it is a well-known fact that, as we pass from one specific
form to another, from the higher to the lower types of orga-
nization, or vice versd, we are impressed with the very few and
inconspicuous structural modifications that we encounter; as we
serially investigate the allied species of a group or groups, the
shading of nearly all of the morphological characters of one
species into those of another, and of the latter into the next
nearest related form, is, as a rule, extremely gradual, and almost
beyond the power of the pen to adequately describe. Yet, if we
choose birds from the extremes of the Passerine order, very
excellent taxonomic characters are met with, and if closely
studied often point to affinities otherwise obscure or very
puzzling.

As shown in many of the characters of its skeleton, Ampelis
cedrorum seems to hold a sort of middle position here; and I
think we shall find that, in some particulars, it is not to be distin-
guished from the highest types of the Oscines, while in others it
possesses marked affinities with the Clamatorial plan of skeletal
structure.

21. Of the Skull (Pl. XVII. figs. 4 & 6).— We find the superior
osseous mandible of Ampelis to be somewhat flattened or com-
pressed from above downwards; the narial apertures are large
and subelliptical in outline. This part of the skull is broad
at its base, tapering rather gradually to its apex ; while above, its
naso-frontal regions gradually merge into each other, there being
no well-defined transverse line dividing them. The median
rhinal partition, or nasal septum, is wholly in cartilage in this
bird (Pl. XVII. fig. 4)—a character wherein it agrees with such
forms as Hesperocichla (Pl. XVII. fig. 5), Merula, and probably
all the higher Thrushes, as well as with such a type as Otocovis*.
On the other hand, a very well-developed nasal septum is to be
found in the skulls of the American Tyrannide, as, for example,
in such a Flycatcher as Tyrannus verticalis (Pl. XVII. fig. 3). So

* Shufeldt, R. W., “Osteology of Hremophila alpestris [ Otocoris alpestris

arenicola],” Twelfth Annual Report U.S. Geol. and Geog. Surv. of the Territories,
1882, p. 652, pl. iv. fig. 22.

310 DR. R. W. SHUFELDT’S MORPHOLOGICAL

far as this character goes, then, Ampelis agrees, as it does in so
many other particulars already defined above, with the Oscines
rather than with the Clamatores.

We also notice that upon the underside of the superior osseous
mandible in the Tyrant Flycatchers the greater portion of that
space which occurs between the anterior limbs of the palatines is
filled in by a continuous plate of bone, which joins them, while
in Ampelis (Pl. XVII. fig. 6) this space is open, as we likewise find
it in most (or perhaps all) Oscines. In the Tyrants the osseous
nasal septum unites with this bony inter-palatine plate above and
along its median longitudinal line.

There is but little to detain us upon the superior aspect of the
skull; all the three specimens shown in Pl. XVII. figs. 3-5 are
rather broad in this region, between the orbital margins, while a
moderately well-marked median furrow is to be seen, which is
best exemplified in the Thrush, and least so in the Tyrant Fly-
catcher. All the higher Thrushes have the facio-cranial line
fairly well defined on this aspect. I have already remarked that
it is barely traceable in Ampelis. .

Upon the lateral view of the skull of this Chatterer (Pl. XVII.
fic. 4) there are two or three points presented of more
or less interest. The “ pars plana,” or antorbital plate, is found
to be ample, and completely divides the orbit from the rhinal
chamber. In front of this, on either side, is to be found
a freely articulated little ossicle which I take to represent the
lacrymal. It shows a constriction at its middle, giving rise to
enlarged superior and inferior extremities. The superior end
is wedged in between the antorbital plate and the postero-
superior edge of the corresponding nasal. Below, its inferior
dilatation rests upon the maxillary bar. Behind, it is applied
against the anterior surface of the pars plana, while in front it
is only in contact with the nasal superiorly, as already described.
I find this bone present in all of the North-American Tyrannide,
in the true Corvide, in the genus Sturnella, but absent in the
Icterid@, in the true Thrushes (Merula, Hesperocichla, Pl. XVII.
fig. 5), in Otocoris, and others. I am not prepared, as yet, to
say of what value it will eventually prove to be as a taxonomic
character; but in the present instance it is just as well to note
that this free lacrymal bone is found in Ampelis and the
Clamatores (ZLyrannide), and not in the true Thrushes.

STUDIES OF THE MACROCHIRES. 3l1

The interorbital septum, in the specimen of Ampelis betore
me, is entire, although encroached upon above and behind by
the great deficiency or vacuity in that part of the brain-case from
which the first pair of nerves make their exit. The foramen for
the exit of the second pair is no larger here than the size of the
nerve demands. In Hesperocichla both of these foramina are
considerably larger than is required for the passage of the
nerves, while in addition a vacuity occurs in the interorbital
septum of the Thrush.

Among the higher types of Thrushes, as in the one just re-
ferred to, the bony entrance to the ear looks directly forwards,
while in the Zyrannide this aperture looks downwards, forwards,
and outwards; in the case of Ampelis it is more as we find it in
Hesperocichia, and in both of these latter forms a conspicuous
shell-like covering of bone is developed from behind forwards,
which protects this important orifice.

In all the true Passerine birds that I have examined a quad-
vate bone has a well-pronounced orbital process; two facets
upon its mandibular foot, the outer being placed transversely,
the inner obliquely ; while the long diameter of its mastoidal
head is also transverse, and supports two articular facets in
Ampelis and the Oscines, which facets in the Zyrannide almost
completely merge into one.

Turning now to the under view of the skull as shown in P]. XVII.
fig. 6, we observe that the anterior extremities of the palatines
(pl.) are very slender and wide apart. I have already pointed
out how the space between them beneath the superior osseous
mandible is filled in by a plate of bone in the Zyrannide. The
postero-external angles of the palatines in Ampelis are rounded
and projecting, much as we find them in some Swallows, and
not very unlike the form they assume in some Swifts.

In this Chatterer the hinder portions of the palatines are
considerably extended laterally, and lie principally in the hori-
zontal plane; whereas in such a bird as Hesperocichla they are
less noticeable for this, while on the other hand their internal and
external “lamine” are better developed in this Thrush.

So far as I have examined, in all Passerine birds these bones
meet for nearly the entire length down the middle line to the
articulation of the pterygoidal heads, and only diverge slightly
in front to meet the backward-extending forks of the vomer.

312 DR. R. W. SHUFELDT’S MORPHOLOGICAL

The mavillo-palatines (Pl. XVII. fig. 6, mxp.) are markedly
long and slender in Ampelis, and mesially separated by an unusual
interval. They slightly exaggerate, however, the conditions
in which we commonly find them among the Passeres generally.
This remark applies also to the vomer (v.), which, too, is rather
broad, though it bears out its Passerine character in being ante-
riorly truncate with somewhat produced lateral tips at that
extremity to meet the cartilaginous wings of the nasal septum.

Among the Zyrannide, the palatine heads of the pterygoids
meet each other beneath the rostrum of the sphenoid; these
parts are separated by quite an interval in Ampelis (Pl. XVII.
fig. 6, pt.), as they are also, though to a less extent, in the
Thrushes (e. g., H. nevia). Here, again, the Chatterer agrees
with the typical Oscinine bird. In general form the pterygoids of
Ampelis agree with those bones as we find them in other Passeres.

In view of the fact that the general characters of the skull of
a Passerine bird are well known, and as I believe that I have
now thoroughly directed attention to the most important
departures therefrom, or similarities therewith, in the fore-
going paragraphs, there is nothing else that remains worthy of
special record in this skull of Ampelis, and the notes thus far
made will meet all requirements of reference for what has to
follow.

The mandible of Ampelis is but feebly developed when com-
pared with that bone as we find it in others of this great group
of birds. It has more the appearance of a Swallow’s jaw than
that of either a Thrush or a typical Flycatcher. In general
form, however, it is essentially passerine, having an outline not
very unlike the mandible of Otocoris alpestris (fig. 29 of my
memoir upon the Osteology of that Lark). A minute ramal
vacuity is present, which is found to be larger in the Zyrannide,
and of a still greater size in Hesperocichla. As might be ex-
pected, the mandible in the Flycatchers is a much stronger and
comparatively heavier bone than it is in these Oscinine types.

22. The hyotd arches in our subject are likewise typically
Passerine in character, having their several elements delicately
constructed, with first and second basibranchial in one piece ;
with a cartilaginous glosso-hyal, and with comparatively large
and free cerato-hyals. Lantus is a bird that shows very well the

STUDIES OF THE MACROCHIRES. 313

Passerine pattern for the hyoid arches, and I have elsewhere
figured them for that form *.

The sclerotal plates of the eye are composed of separate little
pieces of bone in all of these birds of which we have been speaking ;
but in old specimens in some of the Zyrannide I have often
noticed their tendency to anchylose together, notably in the
genus Sayornis.

Passeres usually have a small sesamoid bone at the angle of
the jaw. I have failed to find it in Ampelis, but would prefer
examining more specimens before positively announcing its
absence. It seems to be invariably present where indicated in
all of the higher Thrushes.

23. Of the remainder of the Axial Skeleton.—It will be pro-
bably a very long time before a sufficient number of the vertebral
columns of existing birds have been examined, to ascertain with
certainty whether the vertebre composing them are constant
for the several divisions and for the species. When this has
been done, and the exact averages ascertained, the result will
be no doubt important in assisting to decide doubtful points
in classification

The figures recorded in the subjoined Table have in every case
only been set down after carefully counting the segments two
or three times. With respect to the number of vertebre in
the consolidated sacrum of the pelvis, they are here counted
in the adult bird, and consequently must be taken cwm grano
salis, though I believe them to be quite correct.

I am inclined to think at present that the number of free
coceygeal vertebre are liable to vary even in representatives
of the same species.

This Table, of course, could easily have been made far more
extensive, but in that case would have overstepped the object at
present in view. I trust, however, that those who at any time
may be engaged in examining the structure of birds will record
such data as I have attempted to do here, and only after the most
careful counting. To do this, reliance can be placed only upon
spirit specimens, or recently killed birds, where there has been
no possibility of any of the vertebra having been lost.

* Shufeldt, R. W., ‘ Osteology of Lanius ludovicianus excubitorides,’ Bull.
U.S. Geol. and Geog. Surv. of the Territories, vol. vi. pp. 351-359, pl. xiv.

fig. 101. Also same plate and figure in the author's collected memoirs
‘Contributions to the Anatomy of Birds.’

314 DR. R. W. SHUFELDT’S MORPHOLOGICAL

Species. reachingthe} occur on | sacrum of |division, in-
sacrum. the pelvis. cluding
pygostyle.
Ampelis cedrorwin ......+4. 18 12th. 10 9
Hesperocichla nevia...... 18 12th. 12 a
Tyrannus verticalis ...... 19 13th. 10 8
Otocoris alpestris ......... 18 13th. 1 ‘i

Of this kind of data I think the greatest reliance is to be
placed in the number of free vertebre in the cervical + the
dorsal region, as shown in the first column, far more than that
shown in the three remaining columns. If we take immature
birds, however, chosen just at that point in their development
when we can count with certainty exactly how many of these
segments will enter into the pelvic sacrum, and exactly how
many will be appropriated by the pygostyle, then the total count
of all the vertebre in the column, irrespective of its divisions,
will be of value.

24. In both Ampelis and Hesperocichla five pairs of dorsal ribs
possess costal ribs that meet the sternum, and each of these
birds have a pair of ribs that spring from the first sacral ver-
tebra, which articulate below with a pair of “floating” costal
ribs. This condition also obtains in my specimen of Tyrannus
verticalis, but in this species there are only four pairs of
costal ribs that articulate with the sternum. These “ sacral
ribs’ are without epipleural appendages, though these processes
are found freely articulated at their usual sites upon all of the
true vertebral ribs in the several birds just alluded to.

25. It has been always a matter of some surprise to me that
the pelvis of a bird has not had that importance attached to it
among skeletal characters which has been so universally awarded
to the sternum. In many respects its form in some species is far
more distinctive of the bird to which it belongs than the ster-
num can be,and is consequently more reliable. For instance, the
sternum of Passerine birds varies but very little throughout the
group, and in many cases it would be very difficult to designate
the species by simply examiuing the sternum alone; moreover the

STUDIES OF THE MACROCHIRES. 315

sternum may sometimes assume a different form even for the
same species *, which 1 have never found to be the case in the
pelvis.

In Pl. XVIII. fig. 7 is represented the superior or dorsal
view (double the size of life) of the pelvis of Ampelis cedrorum ;
it shows very well indeed the general form and characters of this
bone as it occurs among the Passeres. Considerable interest
attaches, however, to an examination of a transition series of
pelves through the Passerine group of birds into other orders
wherein marked differences are to be found.

Now in such a bird as Harporhynchus rufus, for instance, or
any of its genus, the pelvis, when viewed from above, has pretty
much the same form as it has in Ampelis; but all the processes
are more prominent, and all the ridges and crests more con-
spicuous and defined. This lends to the bone quite a striking
appearance in these higher Thrushes. But as we pass through
the members of the Oscinine group and into the Clamatores, this
bone, although it retains its general pattern, gradually loses
this peculiar angularity, and gains in breadth while it becomes
comparatively shorter in the longitudinal direction. My meaning
will be made clearer when we come to examine, further on, the
pelves of the Swallows and Swifts.

As to the characters of the pelvis in Ampelis, we are to note
that, anteriorly on its dorsal side, the inner margins of the ilia
are widely separated from the crista of the sacrum; that the pre-
and post-acetabular areas are about of equal dimensions, that the
former are concave outwards, while the reverse condition obtains
with the latter; that the “sacrum” upon this view is roughly
lozenge-shaped, and that interapophysial foramina of varying
sizes may be found to exist in it.

Upon the lateral aspect it is to be observed, that not only is
the acetabulum (as it invariably is in birds, I believe) completely
surrounded by bone, but the ischiadic, the obturator, and the
obturator space are true foramina, or at least are entirely en-
circled by bone. In TZyrannus verticalis the ischium fails to
meet the post-pubis between the obturator foramen and obturator

* For examples of this see my remarks upon the different forms of sternum
in the Cathartide, ‘ Osteology of the Cathartidi,” in Contributions to the
Anat. of Birds, from U.S. Geol. and Geog. Survey, 1882 (Hayden’s 12th
Annual), pp. 771, 772, where four figs. of sternum of C. ura are given.

LINN. JOURN.—ZOOLOGY, VOL. Xx. 25

316 DR. R. W. SHUFELDT’S MORPHOLOGICAL

space, aud thus convert these openings into veritable vacuities, as
is the case in all of the Oscines that I have alluded to elsewhere.

Otocoris agrees in this respect with the Oscines, as may be
seen in my side-view figure of its pelvis (Contrib. Anat. Birds,
pl. iv. fig. 22).

The post-pubis in Ampelis extends but slightly beyond the
hinder extremity of the ischium, which latter meets it behind
in a broad foot-like process. These parts in the dried skeleton
are very apt to curl outwards, and deceive us as to the true
shape of this part of the pelvis during life; so that it is only in
fresh specimens that we can gain a correct notion of this bone
in most Passeres.

This post-preparatory deformity of this part of the skeleton
has been taken into consideration in figure 7, and duly corrected.

The cocecygeal vertebre and pygostyle in Ampelis require no
special description, for they agree in all essential particulars with
the parts as found among the Oscines generally. They are very
well shown in my figure of the skeleton of Otocoris, alluded to
above.

In Tyrannus the coceygeal vertebre are comparatively very
large and their diapophyses very broad.

26. As we would naturally be led to suspect, the sternum of
Ampelis is, of course, a thoroughly Passerine one, having the
characteristic bifurcation of the manubrium; the lofty costal
processes, the well-developed and deep carina, the cordate-
shaped notch on either side of the xiphoidal extremity, and the
jive facets upon either of its costal borders.

Among the American Zyrannide the sternum has essentially
the same shape, but it has only four facets for the hemapophyses
upon each of its costal borders.

A fuller description of this bone will not be required here. I
have already published a pectoral view of a typical Passerine
sternum elsewhere (Coues’s ‘ Key,’ 2nd ed. fig. 58), and other
forms of it may be seen in my figures of the sterna of Lanius,
Otocoris, and others, in memoirs already cited.

This bone will be taken into consideration again, further on,
when we come to treat of the sterna of the Swallows, Swifts, and
Humming-birds.

27. The elements of the shoulder-girdle in Ampelis more
closely resemble those parts in the typical Oscines than in
the Clamatores. In form and arrangement they make scarcely

STUDIES OF TIE MACROCHTRES. 317

any perceptible departures from such structures as we find them
in Lanius (see Contrib. Anat. Birds, pl. xiv. figs. 93, 94, 95, and
100).

The blade of a scapula is sabre-shaped, long and narrow, and
anteriorly abuts against the laterally compressed and expanded
end of the furcula of the corresponding side. The shaft of a
coracoid is long, slender, and subcylindrical in form. Its clavi-
cular head is tuberous, being hooked forwards and inwards.

The furcula typifies the U-shaped pattern of this bone, and ]
would especially call attention here to the form of its hypo-
cleidium in Ampelis. This process is a long, suboval, laterally
compressed lamina of bone, directed upwards and backwards
towards the manubrium, when the girdle is articulated in situ.
My figures, already referred to above, of Lanius and Otocoris
show very well this form of the hypocleidium of the furcula.

Now in Tyrannus verticalis, taken as representing the Clama-
torial group of birds, this process of the furcula is nearly circular
in outline, and decidedly smaller. Little points of this kind,
when they are found to be constant, should be borne in mind
here, for they will surely arise again, when we come to see how
such characters are exhibited among the Macrochires.

28. Of the Appendicular Skeleton.—Not only in the case of
Ampelis, but with the Passerine birds generally, the composi-
tion, forms, and structure of the skeleton of the limbs are so
well known, that I will not here attempt to add anything to this
part of my subject. So far as the bones are concerned, I fail to
find, even upon close scrutiny, any reliable set of characters
that one could use with certainty in deciding in any case whether
the skeleton of a leg or a wing belonged to an Oscinine or a
Clamatorial bird.

When we come to deal further on with the skeletal limbs of
the Macrochires and others, where such characters as are present
in these parts in the Passeres can be, if ever, usefully compared,
it will then be ample time to bring them forward for comparison,
and decide whether these structures afford anything helpful in
determining affinities. To recapitulate here the well-known points
in the skeleton of the limbs in a Passerine bird would, 1 am
sure, avail us nothing.

Suffice it to observe that in its organization Ampelis is by the

majority of its structural characters an Oscinine bird, though
25*

318 DR. R. W. SHUFELDT’S MORPHOLOGICAL

it also presents here and there in its economy traces of a Cla-
matorial type, such as is shown in its free lacrymal bone and a
few other minor points. I fail to find anything in its mor-
phology that especially connects it with the Hirundinide. As
I anticipated, however, the brief synopsis of its structural cha-
racters goes to show, in support of the views already elsewhere
expressed by Garrod, that Ampelis has an organization of an
average Oscinine bird, by which I mean not typically so, and
consequently will be of service here as an aid to comparison as
we proceed.

OsTEOLOGY OF TROGON MEXICANUS AND TROGON PUELLA.

As will be seen by my tabulated list of material, 1 am indebted
to Dr. Sclater for two excellent skeletons of these Trogons.
He kindly had them prepared for me, and sent by post in ample
time to use in the preparation of the present memoir.

In my first paper upon the Macrochires, I stated that I
wished to compare the skeleton of a Trogon with one of a
Humming-bird (P. Z.S. 1885), as Forbes had found in them a
peculiar structure of the palate, which led me to believe that
there might be other points in the skeleton of one of these birds
which would indicate some remote affinity perhaps with the Trochili.
Hence it was with no little interest that 1 opened the package
that Dr. Sclater had sent ; but my eyes had no sooner fallen upon
the two neatly prepared specimens it contained, than the excla-
mation was forced from me, “ Why, they are Caprimulgine
Cuckoos! ”

How well this first impression will be supported by a careful
examination of the skeleton in detail we will now see.

In figuring the osteology of the Trogons, based upon these
two specimens, I chose the skeleton of 7. mexicanus for all my
drawings. This I did as it no doubt agrees in all particuiars
with the skeleton of our own Trogon, T. ambiguus, and I have
long been desirous of contributing to the knowledge of the
osteology of that lone species of the genus in the United States
avifauna.

Practically, however, a description of the skeleton of Trogon
mexicanus will answer very well for the skeleton of T. puella, for
there is but very little difference between them. I must not,
however, be understood to say that the difference in the skeletons

STUDIES OF THE MACROCHIRES. 319

of these two species is of so slight a nature that we cannot
readily distinguish them; for if a series of each were mixed up
before me, there would be no difficulty whatever in rapidly
selecting the skeletons which belong to either of these two
species. As I proceed with my detailed descriptions, I will
point out the more marked of these differences, while my obser-
vations must be considered as applying directly to the skeleton
of T. mexicanus, unless otherwise specified.

Of the Skull (Pl. XVIII. fig. 8, and Pl. XIX.).—Viewing this
part of the skeleton from above, we find that a strong transverse
line separates the superior osseous mandible from the fronto-
lacrymal region, which latter arches upwards and backwards from
it. This superior mandibular portion of the bony beak is rather
low from above downwards, broad at the base, tapering rapidly
to the tip, while its osseous tomia are sharp, and the culmen is a
graceful, rounded, and unbroken arch from the frontal region to
the tip of the beak. The narial apertures at its sides face
upwards and outwards; they are large, of an elliptical outline,
and with smooth, rounded edges.

Each lacrymal bone is a somewhat slender, peculiarly-shaped
element of an f-like form, which freely articulates by its supero-
mesial surface or moiety with a vertical facet offered by the cor-
responding frontal. Its upper end rises above the surrounding
surface of the skull, while its lower extremity fails to reach the
quadrato-jugal bar below.

The region between the orbital peripheries on the superior
aspect of this skull is very narrow, and presents a shallow longi-
tudinal furrow. Posterior to this part the parietal region is
characterized by two laterally placed roundly convexed eminences,
with a rather better marked median longitudinal track between
them, being the continuation of the furrow just alluded to in the
interorbital region.

Viewing the skull of this Trogon upon its lateral aspect, this
marked rounding of the parietal region lends to it rather an
unusual configuration for the cranium among birds.

Upon this view, too, we must note the rudimentary condition
of the sphenotie process, while the squamosal apophysis below it is
large and conspicuous. The interorbital septum, per se, is entire
though very thin, while, on the other hand, the foramina at the
posterior aspect of the orbital cavity are far larger than is required
for the nerves to which they severally give passage; especially

320 DR. R. W. SHUFELD1T’S MORPHOLOGICAL

is this true of the superior vacuity. In both of these Trogons,
however, although these foramina are very large, they still retain
their individuality, and do not merge with each other.

At the upper part of the interorbital septum we observe,
plainly marked, the track for the first pair of nerves. It does
not in either of these specimens communicate, through a thinning
of the walls, with a similar track in the opposite orbital cavity,
as we often see in other birds.

Anteriorly the partition between the orbit and rhinal chamber
is composed of an exceedingly small pars plana, and of a thin
membrane which stretches from it to the already-described lacry-
mal bone. In the dried skeleton, after the membrane has been
removed, the communication between these two cavities is un-
usually free, more so than in any other bird that I happen to
recall at the moment. The periphery of the orbit is sharp for
its anterior two thirds, but becomes rounded off as it passes
backwards over the anterior wall of the brain-case in line with
the lateral processes of the skull.

Sutural traces among the original elements composing the
quadrato-jugal bar are hardly perceptible, and this osseous rod is
quite straight from one end to the other, its tip only being bent
up as it articulates with the quadrate.

Each nasal bone, in this skull, has completely merged with the
premaxillary and frontal of the same side, so that all traces of its
- original borders are obliterated, and 1t has become thoroughly
incorporated in the conformation of the osseous superior mandible.

We will now turn to the underside of the skull of Trogon meai-
canus, where we find many points of interest presented for exami-
nation. Approximately speaking, the majority of structures
observable upon this aspect are found in the same horizontal
plane, which plane nearly includes the entire foramen magnum,
and the anterior moiety of this opeuing may be said to lie
in it.

The roof of the anterior half of the mouth is composed of a
continuous plate of bone contributed by that portion of the pre-
maxillaries. Into the hinder border of this plate we find the
auterior and dilated ends of the palatines merging. At this
point these latter bones are quite close together; posterior to it,
however, an oval, longitudinal interpalatine space occurs through
which we can see the osseous nasal septum and the vomer.

STUDIES OF THE MACROCHIRES. $2)

These latter structures will be alluded to again after completing
the description of a palatine bone.

The anterior half ofa palatine is a narrow ribbon of bone placed
horizontally, dilated at its further extremity, which, as has already
been said, blends with the horizontal portion of the corresponding
premaxillary. The posterior division of the bone feebly develops
an inner and an outer carination, the “ postero-external’’ angle
being completely rounded off.

In the median line, beneath the basisphenoidal rostrum, these
palatine bones meet each other, and in front the vomer, for their
entire lengths, a union which, in both 7. mexicanus and T. puella,
seems to amount to an absolute anchylosis.

if this prove to be universally the case in the skulls of fully
adult Trogons, it need not surprise us, for when we come to the
Humming-birds there will be another peculiar anchylosis to be
described that is occasionally to be found in their skulls.

Now the vomer (fig. 8, v.) in both of these Trogons is a rather
short subcylindrical rod with a bluntish point. This point rests
directly upon the posterior free edge of the osseous nasal septum
(1.8.).

This intimate relationship between the vomer and nasal septum
in the Trogons led the late W. A. Forbes into an error, which will
be at once evident upon an examination of his drawing of these
parts in Pharomacrus mocinno (P. Z.S. 1881, p. 887). At least
it does not hold quite true in the species at hand; and I suspect
that Forbes, in examining alcoholic specimens, included this thin
posterior edge of the nasal septum with the anterior tip of the
vomer, giving it that “thin and filiform” appearance to which
he alludes. This slight error might easily be made by that kind
of an examination, when in a dried skull, sich as I have before
me, these parts would be better distinguished. His description,
however, in the contribution above referred to, is a marked
improvement upon that by Professor Huxley, which it was
written to correct; and the former writer was fully aware of the
fact that the nasal septum in the Trogons ossified. In my
specimen of 7. mewicanus this plate has a large vacuity in its
centre, while in 7. pwella it is entire.

The tips of the mawillo-palatines do not show in the interpala-
tine median space, upon this view of the skull, as they are said
to do in P. mocinno by Forbes ia the paper just quoted. (Com-
pare figure in P. Z. 8. 1881, p. 837, and fig. 8 of the present

322 DR. R. W. SHUFELDT’S MORPHOLOGICAL

paper.) Indeed, I can fully verify Forbes’s statement that
the Trogons are not desmognathous birds, but very decidedly
schizognathous ; and, furthermore, I am inclined to believe that
there is a fair measure of truth in the words of this anatomist
when he stated “ that the structure of the palate has not that
unique and peculiar significance that has been claimed for it in the
classification of birds.” My eyes were opened to this fact more
thoroughly than ever when I came to find such a bird as Chor-
deiles completely desmognathous, while certain forms of Capri-
mulgus and Phalenoptilus were egithognathous, as defined by
Professor Huxley in his well-known “Contribution to the
Classification of Birds’ (P. Z. 8. 1867, p. 468).

Large basipterygoid processes are developed both in 7. mexicanus
and 7. puella, which in each case are articulated with ample facets
upon the pterygoids themselves.

The palatine heads of the pterygoids are widely separated in
the median line; while, as I have already stated, the palatines
opposite their place of meeting them are in contact to their very
ends. The outer edge of a pterygoid is quite sharp, while this
bone is compressed from above downwards and articulates in the
most usual manner by a ball-and-socket joint with the corre-
sponding quadrate.

A quadrate develops a long orbital process with dilated tip.
Its mandibular foot supports two facets upon it, which are sepa-
rated from each other by an intervening valley. There are also
two such facets upon the squamosal head of the quadrate.
Otherwise this element is notable for the unusually long apophysis
it offers laterally to meet and articulate with the posterior end
of the quadrato-jugal bar.

The underside of the basisphenoidal rostrum is much thickened
and rounded for its entire length, thus affording a broad sliding
surface for the articulation of the pterygo-palatine ends. Barely
an apology for a bony scale overhangs the entrances to the Eu-
stachian tubes, while immediately behind them the base of the
cranium is very broad between the aural apertures. The occipital
condyle is comparatively very small indeed, though the subcircular
foramen magnum is fully up to the average in point of size. Upon
either side of it, we find the usual group of foramina for the
passage of nerves and vessels (Pl. X VIII. fig. 8).

I have made no special examination of the interior of the
cranial casket in these birds, but we are to note the great delicacy

STUDIES OF THE MACROCHIRES. 320

of the walls of this part of the cranium and how thin they are.
Air seems to gain access to the major portion of the skull in both
of these specimens, including the quadrates and perhaps the
pterygoids. 7

I am reminded in my examination of the mandible of Trogon
of the form this bone assumes in some of the smaller American
Owls, as Speotyto for example. Its articular ends are rather
large, being bluntly pointed behind, and having long, sharp, in-
turned mesial tips.

The borders of the rami are rounded off, while their height
remains quite uniform for the entire length of the jaw. Upon
their outer aspects, for the posterior moiety of each, an excavation
occurs, at the middle of which, on either limb, is seen a small
ramal vacuity.

The symphysis is deeper by half again than either ramus, and
the superior border above it issharpened. In generai outline the
mandible of a Trogon is broadly V-shaped, and this bone is partially
pneumatic.

So far as these two specimens are concerned, I find that 7
puella differs from 7. mexicanus in its skull in having an entire
osseous nasal septum, a rather wider frontal space on the superior
aspect of the skull between the orbital margins, the parietal
eminences are not so lofty, and a better developed osseous lip
protects the entrance to the Eustachian tubes. Their mandibles
are essentially similar.

Of the Hyoid Arches.—As might be expected, these practically
present little or no difference in the two species of Trogons
before me. The hyoidarches in 7. mexicanus are small as com-
pared with the size of the skull of the bird, the thyrohyals barely
curving up behind at all. The apparatus as a whole reminds me
not a little of the hyoid arches in some of the smaller American
Owls (Glaucidium).

The glossohyal is formed entirely of cartilage, while the
ceratohyais have ossified. In this adult bird the first and second
basibranchials are joined in one piece by anchylosis, the cerato-
branchial of the thyrohyals apparently articulating in the lateral
sockets at their point of union.

Cartilaginous tips finish off the hinder ends of the epibranchials,
and these elements of the ‘‘ greater cornua” are nearly straight
longitudinally, nor are they notably curved in the direction of
the median plane of the body.

824, DR. R. W. SHUFELDT’S MORPHOLOGICAL

Of the Remainder of the Axial Skeleton.—From a careful ex-
amination of the vertebral column of both of these Trogons, I am
enabled to present the subjoined table of data in regard to this
part of their skeletons and in which both these species agree :—

Number of cervical vertebrx without ribs .......... 12
The first cervical vertebra supporting a pair of free ribs =13th.
The first vertebra of the column wherein the ribs articu-

late with the sternum by means of costal ribsis the 14th.

Number of true ribs thus articulated .............. 4
Number of hemapophysial facets on sternum........ 4
Number of true-dorsal vertebra ...... 0a. ae 5
Number of vertebre consolidated with pelvis........ 12
Number of caudal vertebree (free) including the pygo-
style are... 008405 ee ue oes + +o ae

It will be noted from this table, of course, that the pair of
vertebral ribs attached to that vertebra here called the fifth dorsal
have costal ribs that do not meet the sternum, but articulate
with the hinder borders of the last pair of costal ribs that do.

There is also a pair of ribs that spring from beneath the fore
part of the sacrum, articulating with the first or anterior vertebra,
that becomes consolidated with the pelvis. These ribs also meet
short and free hemapophyses below, which in their turn articulate
with the posterior margins of the pair of first or leading “ floating
ribs”’ referred to in the foregoing paragraph.

Neither the last pair of dorsal vertebral ribs nor the sacral
vertebral ribs possess epipleural appendages ; consequently we
have but five pairs of true vertebral ribs that do support these
processes.

And, further, we ascertain from the table that the total number
of free vertebre in the column before we arrive at the first one
that becomes consolidated with the pelvis is 18.

Among the principal characteristics of the first twelve vertebrae
of the column we are to note that the usual outstanding processes,
such as the neural spines, parapophyses, and hyapophyses, are
but feebly developed, being for the most part short and incon-
spicuous. ‘lhe superior spines of these vertebre are most pro-
minent in the axis and next following three or four segments.

In neither of the specimens examined does a perforation exist
in the cup of the atlas, while the neural arch of this verte bua is
comparatively narrow.

STUDIES OF THE MACROCHIRES. 325

The hypapophysial processes of the fifth, sixth, seventh, and
eighth cervical vertebre part mesially in order to form a canal
for the protection of the left carotid artery, the only one present
in these birds (Garrod), as it passes to the head.

Upon the last few cervicals, and upon the first two or three
dorsals, very well-developed hy papophyses are to be found, which
may become tricornuted towards the latter end of the segments
specified (Z. puella).

Diapophyses of the dorsal vertebree are eoebusls broad, with
their outer extremities rounded, being considerably wider even
here than the bodies of the ribs that articulate beneath them (7.
mMexicanus).

Below, the centra of these dorsal vertebra are deep and much
compressed laterally; their median, longitudinal, inferior lines
being almost cultrate in character.

The dorsal neural spines are all nearly of the same height, but,
on the whole, rather low in point of altitude. Their superior
rims are thickened, and interlock at their anterior and posterior
ends. These five dorsal vertebra, although freely articulated, are
very closely moulded upon each other, and consequently the mo-
bility of this division of the column is somewhat limited.

Facets for the vertebral ribs are here, as usual, found for each
pair just within and posterior to the anterior margins of the
centra of the respective vertebre at their lateral aspects.

Thus far in the column of these Trogons the plan of articu-
lation seems to be truly “ heteroccelous,” 7. e. the ends of the
centra present saddle-shaped facets, which in turn lock with
a counter-placed one on the opposed face of the vertebra next
behind.

Of the Sternum.—Leaving the vertebral column for the moment,
we will turn our attention to this bone. There is a good deal in
the sternum of Zvrogon to remind us of the same part of the
skeleton in Geococcyx ; of which we may at once satisfy ourselves
by comparing the figures of the bone as it is found in these two
forms and shown in my plates. Figures of the sternum of Trogon
mexicanus illustrate this memoir (Pl. XIX. figs. 12,13), while
corresponding views of it for Geococcyx are to be found in the
plates of my contribution upon the osteology of G. californianus
in the ‘Journal of Anatomy and Physiology’ for 1886.

Among the Trogons the sternum is short, and, when taken in
comparison with the size of these birds, rather expansive behind,

326 DR. R. W. SHUFELDT’S MORPHOLOGICAL

where it shows two deep rounded notches on either side of the
carina. The outer pair of lateral processes which are thus pro-
duced have expanded posterior ends. As already stated, each
costal border supports four small transverse facets for the hema-
pophysial ribs, which are crowded pretty close together. Beyond
these the costal processes are of a quadrate outline on either side,
and are directed upwards and very slightly forwards as conspicuous
projections.

Tribedral in form and rather inclined to bifureate at its summit,
the well-developed manubrium is cultrate mesially in front and
this prominent apophysis rears itself upwards and forwards from
its usual site, as seen in birds generally where it is present.
Immediately behind it are the coracoidal grooves, which, though
narrow from above downwards, extend far out laterally and meet,
or very nearly meet, at the middle point posterior to the manubrial
base.

The keel of this bone is comparatively deep and extends the
entire length of the sternal body, while its inferior border is
gently convex for its entire length forwards, and its anterior one
considerably concave. At their intersection in front the carinal
angle is bluntly rounded off.

From the fact that the xiphoidal processes are spreading in
character and the sides of the sternal body uniformly raised,
not an inconsiderable concavity is enclosed on the thoracic
aspect of this sternum ; and, as is usually the case in birds of
this form, the pelvis above is seen to be correspondingly wide-
spread, indicating roomy abdominal and thoracic cavities within
their enclosure. |

Pneumaticity is not a prominent feature of the sternum among
the Trogons, although a few insignificant foramina do admit air
to this bone where such openings ordinarily occur.

Of the Shoulder-Girdle (Pl. XIX. fig. 13)—We find the os
furcula to be of the U-shaped variety, with a well-formed hypo-
cleidium of a rounded outline at its clavicular junction beneath.
The limbs of this bone are slender, while the heads are somewhat
expanded and much compressed laterally. They rest against the
mesial aspects of the superior tuberosities of the coracoids, and
on either side extend backwards to the scapula, which they over-
lap to some extent, resting upon the upper side of the clavicular
process of the bone.

A scapula here makes the usual articulation with a coracoid

STUDIES OF THE MACROCHIRES. 327

and offers the proportional amount of articulatory surface to the
glenoid cavity. Its blade is narrow, rather long, of an equal
width throughout, and compressed from above downwards. Pos-
teriorly its extremity is obliquely truncate from within outwards,
while the end itself is slightly curved in the same direction.

Hither coracoid is characterized by a very extensively expanded
sternal extremity of a quadrilateral outline, and of no great thick-
ness in the antero-posterior direction. The shaft of the bone
above this dilated end is rather slender, subcylindrical, being
compressed from before backwards, and is evidently hollow. Its
summit is not conspicuously enlarged, though it is rather more
tuberous than we find it in such a group, for instance, as the
Passeres. The head is directed in the articulated skeleton upwards,
forwards, and inwards. Its scapular process is not very wide, for
the scapula projects over it a little, both mesially and to its outer
side ; while in the former direction it stands between its superior
articulating edge and the corresponding head of the clavicle, 7. e.
the scapula does. Air seems to gain access to the shafts of the
coracoids, and perhaps to some extent to the extremities of these
bones; but, so far as I have been able to discover, neither the os
Surcula nor the scapule possess any pneumaticity.

Neither of these Trogons possess, upon either side, the little
ossicle at the shoulder-joit known as the os humero-scapulare,
though it is just possible that it may in every case have been
removed by accident during the preparation of the specimens.

Of the Pelvis and the Coccygeal Vertebre.—No marked differ-
ences distinguish the pelves of these two species of Trogons.
There is some general resemblance between the pelvis of Z. mewi-
canus and the bone as we find it in certain Caprimulgine birds,
though when we come to the details in such a comparison the
divergence is sufficiently marked.

Viewing the pelvis of Trogon mexicanus from above, we observe
that the preacetabular area is considerably more extensive than
is the postacetabular (Pl. XIX. fig. 14). The outline of this
upper surface is somewhat quadrilateral, its average width being
about equal to its average length. In this specimen there are no
existing vacuities among the diapophyses of the sacral vertebra.
One or two extremely minute ones are found in these positions in
the specimen of 7. puwella among the ultimate vertebrae.

Marked lateral extension characterizes the transverse pro-
cesses of the sacral vertebra, more especially those three which

328 DR. R. W. SHUFELDT’S MORPHOLOGICAL

are about opposite the acetabule; those in front and behind
these are proportionately graduated off; and although, as I have
observed, no foramina exist among them, the overspanning bone
is in some places exceedingly thin.

The anterior margins of the ilia are obliquely truncate from a
direction within outwards and backwards; and these borders
have a delicately thickened rim. Internally, the margins of
these bones fail, on either side, to meet the consolidated neural
crest of the leading sacral vertebre, there being short “ilio-
neural canals’’ present in the pelves of these Trogons, of some
two millimetres in width.

As to the superficial form of the several areas of this pelvic
roof, we find the anterior moieties of the ilia to be concave ;
the posterior and smaller ones convex; and the middle area,
formed by the sacrum, is an ample lozenge in its general
outline.

Turning now to the lateral aspect of the pelvis of this Trogon
(Pl. XIX. fig. 13), we are to observe that the anterior or pre-
acetabular division of the bone lies in the same general plane
with the longitudinal axis of the dorsal vertebre, while there is a
gradually increasing droop of all the hinder division of the bone,
until we arrive at the slender post-pubic element, the posterior
extremity of which turns slightly inwards and upwards.

Comparatively speaking, the acetabulum is rather small, and
its base is deficient in bone, being so rendered. by the usual
circular vacuity there. The antitrochanter occupies here its
common site, above the acetabulum, and faces forwards, down-
wards, and a little outwards. Behind it again we find an ample
and subelliptical ischiadic foramen, situated thoroughly within
the borders of the surrounding bone. The obturator space and
the obturator foramen have so merged with each other that
scarcely a distinguishing trace of separation can be detected
between them.

A long subelliptical foramen is thus formed, the lower margin
of which is bounded, as usual, by the narrow bar of the post-
pubic element, as it sweeps by to the rear. This foramen is
closed in behind by the foot-like process of the ischium which
descends to meet the post-pubis, the latter extending for some
three or four millimetres behind it, and thereafter taking a
direction already described in a foregoing paragraph.

On the underside of this pelvis we find its “basin” to be

STUDIES OF THE MACROCHIRES. 329

wide and capacious, though not at any point correspondingly
deep. The first three or four sacral vertebre throw out their
lateral processes to abut against the nether surface of the ilia,
on either side, their extremities co-ossifying with the same;
while the leading vertebra of all of the sacrum, as I have already
said above, supports a pair of freely articulating ribs. A strong
pair of lateral processes come off from the sacrum opposite the
acetabule, and by abutting against the inner pelvic walls just
above these circular apertures, they form strong braces to this
part of the pelvis. Other members of the class frequently pos-
sess this feature. Now the posterior border on either side of
this pelvis shows scarcely any mark to distinguish the union of
the ilium and ischium, beyond a slight elevation at the usual
point; in some birds, as we are aware, a notch defines the
place.

Among the coccygeal vertebre of the tail the transverse pro-
cesses are all long and spreading, the last three being con-
spicuously so. Their neural spines are low and not prominent ;
while only the ultimate vertebra supports a bifid hypapophysis
beneath its centrum.

To complete my account of the axial skeleton of a Trogon, it
may be noted that the pygostyle is of arhomboidal outline, with a
considerably thickened base, and a perforation in its plate-like
part near the supero-anterior angle.

Of the Appendicular skeleton ; the Pectoral Limb (Pl. XIX.
fies. 12-14).—No very striking feature distinguishes the hwmerus
of one of these birds from the same bone as we find it in a
considerable number of the Passeres. It seems to be thoroughly
pneumatic, and the fossa that harbours the foramina occupies its
usual site, and is surrounded in the usual manner by the ulnar
crest at the proximal extremity of the bone.

The radial crest is rather low ; its free border being long and
convex, and the plate itself being bent palmad, as we so commonly
find it among Passerine birds.

Coming to the shaft, we find it to be of a subcylindrical form,
somewhat compressed laterally, and possessing the usual sigmoid
curve, only in a moderate degree.

Nothing worthy of special record marks the distal extremity
of this humerus, it being tuberous only to an extent in harmony
with the general size of the bone; and upon its palmar aspect

330 DR. R. W. SHUFELDT’S MORPHOLOGICAL

are to be found the two usual tuberosities for articulation with
the bones of the antibrachium. |

In neither of the Trogon skeletons at hand do I find any
sesamoid bones present at the elbow-joint ; but it is just possible
that these birds may possess them, and that in the present
instance they have been lost in the preparation of the skeleton ;
I am inclined to believe, however, from the general appearance
of the dried ligaments and other structures that have been
retained in one specimen, that the Trogons do not have these
ossicles at the elbow-joint.

Turning to the bones of the forearm, we observe that the shaft
of the radius is very slender and nearly straight. Its extre-
mities are comparatively but slightly expanded, and on the
whole this bone is not so powerfully developed as we find it in
many birds of the same size. The interosseous space between it
and the ulna is ample, but is largely due to the curvature in the
shaft of the latter bone. The ulna has the usual form as we
find it in some of the Passeres. Its olecranon process, however,
is not conspicuous, while the shaft is smooth, subcylindrical, and
devoid of the row of papille for the insertion of the quill-butts
of the secondaries of the wing, so prominent in some birds, as,
for instauce, many of the Picide.

Radial and ulnar ossicles compose the carpal joint, and make
the usual articulations with the surrounding bones. Jam unable
to discover any sesamoids about this joint, and am of opinion
that none exist.

Extending our observations to the hand, we may note the
peculiar form of its main bone, the carpo-metacarpal. This
peculiarity does not consist in any radical change of its form as
it is found to be in most birds, but of the unusual width it as-
sumes at its distal end, in the antero-posterior direction (Pl. XIX.
fig. 13). The increase of surface thus gained is for the accom-
modation of the articular facet for the single and terminal
phalanx of mid-metacarpal digit, here disproportionately large
as compared with the bones of the other two phalanges.

Pollex phalanx is small, slender, and trihedral in form; it is
not provided with a claw at its distal extremity.

A very similar joint is the distal phalanx of index digit, and
this, too, is without a terminal claw.

The proximal bone of this finger has the general form it as-
sumes among birds, but in the present instance the expanded

[
{

STUDIES OF THE MACROCHIRES. 33al

_ blade-like portion behind is very thin, its surrounding margin
_ only being thickened to lend it the proper support.

‘When we come to measure the lengths of the bones composing
the pectoral limb in this Trogon, we find that the humerus is
2F1 centimetres long, the ulna 3°6; and the skeleton of manus,

; measuring from the summit of the carpo-metacarpus to the distal

“4pex of the last phalanx of index digit, 3:0 centimetres long.
| Of the Pelvic Limb (Pl. XIX. fig. 13)—Trogons have a
thoroughly pneumatic femur, and a large foramen or two to

admit the air are found upon the anterior aspect, near the sum-

mit of the bone, between the trochanter and the head. I am not
quite certain but that the tibia also possesses a moderate amount
of pneumaticity, as the shaft is completely hollow and the bone
has the general appearance of a pneumatic one. However, I
have failed to discover the presence of the foramina in this part
of the skeleton of the leg. As for the fibula and the remaining
skeletal parts of this limb, they are entirely devoid of this
feature.

Returning to our examination of the femur, it is to be noted
that the trochanter is but feebly pronounced, and does not rise
above the summit of the bone. The caput femoris is globular
and quite sessile with the shaft. We can scarcely discern any
pit whatever upon its superior surface to lodge the attachment
of the ligamentum teres. Descending to the shaft we find this
part of the bone nearly cylindrical in form, very smooth, and
quite straight. At its distal extremity the condyles are rather
small comparatively ; the external one being situated lower, and
at the same time somewhat more prominent than the internal
one.

Trogons possess a very well-developed patella of a subcordate
form, it being fully twice as wide as it is deep.

Regarding the fibula, we find that it presents little or nothing
worthy of special note. Feebleness of development charac-
terizes this bone in the Trogons throughout. Its head is
small and the shaft slender, the lower end of the latter being
free from the tibia, and descending to a point about opposite the
junction of the middle and lower thirds of its shaft. Scarcely
any evidence exists of the presence of the tubercle for the inser-
tion of the tendon of the biceps muscle, a feature which is
quite prominent in some birds.

LINN. JOURN.—ZOOLOGY, VOL. xx. 26

332 DR. Rs W. SHUFELDT’S MORPHOLOGICAL

Next, passing to the consideration of the tibia, we find its
shaft to be nearly straight, being but slightly convex forwards;
while for the greater portion of its midcontinuity it is of a sub-
cylindrical form, changing only as it approaches its enlarged
extremities. At the proximal end of the bone the pro- and
ectocnemial ridges on its anterior aspect are considerably sup-
pressed, and soon merge into the shaft below; they are nearly
of equal size, and the cneniial crest above them does not rise
above the tibial summit.

Although the condyles at the distal end of the bone are very
similar to these protuberances as they are commonly found in
the majority of small birds, they are yet peculiar in having
between them, below and behind, a mid- and well-marked longi-
tudinal ridge, constituting a feature that at this moment I do not
remember to have noticed among the tibie of the class.

On the anterior aspect of this tibia, just above the condyles,
we notice the usual longitudinal tendinal groove, spanned at its
lower part by an osseous bridgelet thrown directly across it.

The tarso-metatarsus has a length equal to rather more than half
the length of the tibia, while the calibre of its shaft is about one
third less than that of the latter bone. This tarso-metatarsal
shaft presents three plane and thgrooved surfaces, an anterior
one and two lateral, or rather postero-lateral, ones. The summit
of the bone is moulded in the usual manner for articulation with
the tibial condyles. Behind the proximal extremity of the bone
we find a fairly well-developed hypotarsus, vertically pierced by
two tendinal canals placed side by side.

Passing to the distal end of the tarso-metatarsus, we find the
trochlez so disposed as to accommodate themselves to the zygo-
dactyle condition of the podal digits, which consists, as we know,
in Trogons of a permanent reversion of the second toe.

These digits have their bony phalanges arranged upon the
most usual plan as we find it in the vast majority of living birds,
2. e. 2, 3,4, and 5 joints for the first, second, third, and fourth
toes respectively.

Considered as a whole, although the skeleton of the foot of
this bird is in due proportion with the rest of the limb, it never-
theless strikes us as being rather a delicately formed structure.
The accessory metatarsal possesses a shape usually assumed by
it among birds, but in the present instance makes a very close

STUDIES OF THE MACROCHIRES. 333

articulation with the shaft of the tarso-metatarsus at its most
usual site.

Measuring the lengths of the several bones of this pelvic limb,
as we did in the case of the pectoral one, we find that the femur
is 2°3, the fibula 2°1, the tibia 3:3, and the tarso-metatarsus 1°6
centimetres long.

Without measuring the several lengths of the joints of the
pedal digits of a Trogon, I am enabled to say that they are quite
as harmoniously proportioned as are the corresponding phalanges
of the average foot of any Passerine bird that I have ever
examined.

This completes my description of the skeleton of 7. mexicanus,
and, as observed, it will apply with almost equal exactness to the
skeleton of 7. puella. In proceeding with my account I have
been careful, I believe, in every instance to point out any con-
stant character that seems to distinguish them; and no doubt
my description will practically answer for other nearly related
species of this handsome group of birds.

It seems scarcely necessary to tabulate the salient features of
the osteology of this Trogon here, as my brief account presents
but little more than an enumeration of the essential charac-
teristics. It will therefore be omitted, in the belief that the
several figures illustrating my text and the description will be
amply sufficient even for convenience of reference.

Comparing these osteological characters of Trogon with the
corresponding ones of such a Humming-bird as 7’. Alexandri, as I
presented the latter in my former memoir, P. Z. 8S. 1885, it will
at once become evident to us that, so far as the skeletology of
the two forms is concerned, there is absolutely little or nothing
that mutually characterizes them.

So much for the comparative osteology of Humming-birds and
Trogons, but this will not exactly apply to some other groups of
birds, such, for example, as the Cuckoos and Nightjars; and I
will now proceed to draw a few comparisons among some of
them.

I regret to say that the only Cuckoo-like bird I have at hand
is Geococcyx californianus, and, as stated above, I have already
published an account of its osteology in the ‘ Journal of Ana-
tomy. JI did have, not long ago, a fairly good skeleton of
the Yellow-billed Cuckoo (Coccyzuws americanus); but Prof.
Parker was at that time in search of all the Cuculine birds

26*

334 DR. R. W. SHUFELDI’S MORPHOLOGICAL

he could procure, and it gave me great pleasure to forward it to
him along with a few others that I had collected, in response to
his request for such material.

The characters of that skeleton have escaped me, but the
reader can easily compare such forms as he may have at hand
with what follows.

My former memoir (P. Z.S. 1885) contains an extensive
account of the osteology of Chordeiles and Phalenoptilus; so in
the present connection I may point out what has been already
ascertained in regard to a comparison of these Caprimulgine
forms and Geococcyx with the Trogons. Thanks to my friend
Mr. Sage I have before me a fine alcoholic specimen of our
American Whip-poor-will; but I do not intend to dissect that
until we enter upon the next section of this memoir, wherein
it will constitute my type for the general anatomy of a Capri-
mulgine bird.

A comparison of the skulls of Trogon, Chordeiles, and Geo
coccyx need not detain us long, for they have but very few
characters in common. With respect to the skulls of Trogon
and Gteococcya# they may be dismissed by stating that they differ
from each other in every essential particular, beyond the fact
that they are both skulls of birds.

This difference is quite as great when we come to compare the
skulls of Chordeiles and Geococcyx, for here, too, it would be
very difficult, if not quite impossible, to pick out a single feature
in the one that would in any way be comparable to the corre-
sponding one in the other.

Except for the fact, as stated, that they are both skulls of birds,
they are totally unlike.

Not nearly so much so is this the case with Zrogon and
Chordeiles ; for, different as the skulls of these two forms really
are, I think I can see a certain resemblance between them,
slight as it is.

Still even here, at the best, it is little more than a superficial
likeness; they have, however, in common the basipterygoidal
processes, if nothing beyond that. Their mandibles, as we know,
are entirely dissimilar. Notwithstanding this, it would be far
easier for us to conceive that a Trogon’s skull was a very much
modified Caprimulgine one than it would be to picture any
relation between it and the skull of Geococcyz.

With these facts before us we are not surprised to find, what

STUDIES OF THE MACROCHIRES.

is really the case, that the osseous hyoid arches of

335

these several

types are also of very different patterns, and do not suggest te

us any special relationship of their owners.

To sum up then, so far as the skulls of these birds are con-
cerned. Had we no other part of their structure to guide us,
I think we should be fully justified in placing them in separate

orders of birds.

But let us still further compare the charac-

ters of their osseous structures and pass to a consideration of

the remainder of their axial skeletons.

First let us take a glance at the number of vertebre in the
spinal columns of the several birds in question, irrespective of
any special characteristics these vertebrae may possess in them-

selves. For I wish it to be distinctly understood that although
Number of verte- ‘
bree in the cervi-| The first vertebra | ee ane ee
Species. eal portion of the | that bears a pair | Aen with these.

column that are | of free ribs.
without free ribs.

13th
(without unciform
processes).

12
Trogon mexicanus

eeeeeeeee

12th
(with unciform
processes).

11
Chordeiles texensis

«ee eeetee

Geococcyx californianus ... 12 13th

num by costal ribs.

14th

13th
(14th in
P. Nuttalli).

15th

Number of dorsal

vertebrae con-
necting with ster-
num.

+
(and one more
dorsal vert. that
does not so con-
nect, making 5
dorsals).

No. of vert. in
sacrum and the

Species. ;
P sacral ribs.

12
There is one pair
of sacral ribs.

3 10
sacral ribs

sternum,

eeeee

join

3 id
(and one more
dorsal vert. that
does not so con-

nect).

Geococcyx californianus

No. of free tail
vertebra, includ-
ing pygostyle.

(

6
(5 in the Whip-
poor-will)

336 DR. R. W. SHUFELDT’S MORPHOLOGICAL

I place the just amount of weight that should attach to the
number of these segments in the spinal column of any bird, IL
think it should be borne in mind that these vertebre are as
much entitled to be considered in the light of the special form
each or any of them may assume, as is any other part of the
skeleton.

The day may yet come when the question of the exact affinity
of avian forms (or any other class of vertebrates for that matter)
will have arrived at such a point of refinement as to require
that even the morphology of each vertebra shall be known, to
assist us in correct decisions. In the table which I here
introduce (p. 335) the number of ribs and some few other points
which I deem it well to compare have been entered.

So far as we are able to judge by a comparison of these, it
would seem that, taking into consideration the kind of data pre-
sented, Trogon comes nearer to G'eococcysx in its vertebral column
than it does to any of the Caprimulgi. But it must be remem-
bered that it is really very difficult to discern any truly striking
resemblances among the vertebral columns of the several birds
under consideration.

Turning to the pelves, we find on comparing the pelvis of
Trogon with that bone as we find it in some of the Nightjars and
Whip-poor-wills, that there is a certain superficial likeness which
strikes us; but when we descend to the comparison of details, we
are again met by the fact that these resemblances are purely
superficial Of course neither the pelvis of Chordeiles nor
Trogon reminds us in the least of the unique pelvis which so
conspicuously characterizes the skeleton of Geococeyx. How
they would compare with certain other Cuckoo-like birds I am
unabie at present to say, from lack of proper material on which
to form an opinion,

Passing to the sternum (and I have figured this bone for both
Chordeiles and Geococcyx in my memoirs above referred to,
and for Trogon in the present paper), we are at once struck by
the resemblance between the sterna of Trogon and Gleococcyz ;
the bones here are really very much alike, and both are
essentially different from the single-notched sternum of Chor-
dewles.

Coming next to the shoulder-girdles, we are once more at
sea, for these parts not only have no special likeness to each
other, so far as Trogon and the Caprimulgi are concerned, but

STUDIES OF THE MACROCHIIRES. 337

both, on the other hand, are extremely unlike the pectoral girdle
as we find it in Gleococcyx.

These remarks apply with equal truth and force to the pectoral
and pelvic limbs of the several forms under examination; and
even in the case of the reversion of the toes in Trogon and the
Ground-Cuckoo, we are all aware that in the former bird the
second toe is turned back, while in the latter it is the fourth
one that is reversed.

I agree entirely with Professor W. K. Parker when he
states that “the familiar term ‘ zygodactyle’ for birds with a
certain form of foot has been very useful; and yet how much
ignorance it may be made to hide! It seems to be something
when one knows that a certain bird belongs to that group; and
yet a Cuckoo, a Parrot, and a Woodpecker come none the nearer
each other zoologically by the possession of that kind of foot ’’*,

To recapitulate then, and judging from the skeletons alone, we
must see that such a form as Geococcyx californianus is more or
less remotely related to such birds as <Alcedo and Dacelo,
perhaps much nearer them than it is to the true Tree-Cuckoos.
In saying this J am aware that in a paper recently read for me
before the Zoological Society I was still inclined to support
the classification of Garrod, who divided the Cuculide into two
subfamilies, viz. the Ground-Cuckoos (Centropodine) and the
true Cuckoos (Cuculine) (P. Z. 8. 1874, p. 121); and this paper
of mine referred to the anatomy of G. californianus, but at the
time I had no specimens of true Cuckoos to compare it with.
Still I am inclined to adhere to that opinion until I have had an
opportunity of making further researches into the structure of
many other types more or less nearly related. On the face of it
I should be disposed to think that Geococcys, so far as its skele-
ton is concerned, came nearer to such a form as Dacelo gigantea
than to Cuculus canorus, for instance, notwithstanding the
structure of the foot. But many of these interrelated groups
are exceedingly puzzling, and still require a considerable amount
of original investigation of their morphology.

Of the Caprimulgi, of course, I shall have more to say further
on; it is very evident, so far as their osteology indicates, that
they are very widely separated from the Trogons.

And now as to the Trogons themselves, still being guided by

* Parker, W. K., “On the Morphology of the Skull in the Woodpeckers
and Wrynecks,” Trans, Linn, Soc., Zool. 2nd ser. vol, i. pp. 1-22.

338 DR. R. W. SHUFELDT’S MORPHOLOGICAL

the skeleton alone, it is very plain that they have no special
affinity with the Trochili.

When my eyes first fell upon the skeleton of one of these
Trogons, as I have already stated, the remark was forced from
me that they were “ Caprimulgine Cuckoos.” First impressions,
however, are not always to be relied upon, for, apart from the
general likenesses of their sterna, and having the same number
of cervical vertebre without ribs, from skull to pygostyle, and
from pinion to pes, so far as the skeleton goes, Trogon mexicanus
has nothing whatever to do with Geococcyx californianus, and, as
stated, I have no true Cuckoos to compare it with. I dare
say that if related to the Cuckoos at all, it is more than likely
to be nearer these latter in its osteology. I have not had the
opportunity, as yet, to examine the skeletons of either Croto-
phaga or Scythrops.

Beyond a few apparent resemblances I find nothing in the
skeleton of the Trogons that in any way points to their being re-
lated, even remotely, to the order Caprimulgi; and it seems to
me that there must be a considerable gap between the Trogons
and Kingfishers.

Tur ANATOMY OF CERTAIN CAPRIMULGI.

Glancing at our list of material, we find that we have but a
few American forms to illustrate the structure of this highly
varied group of birds. There is an alcoholic specimen of the
common Whip-poor-will, a couple of skeletons of the Nuttall
Poor-will, and several species of Chordeiles, both in alcohol and
skeletons.

In my first contribution in the P. Z.S., relative to the structure
of the Caprimulgine types, I gave a very full account of the
osteology of Chordeiles, with a number of figures, as well as a
description of the skeleton of Nuttall’s Poor-will. I am con-
vinced that when the hundred or more species composing this
order come to be carefully examined as regards their structure,
there will be not only some very good characters brought to
light, but considerable difference found in the economy of the
forms composing the group.

Among the notable departures it will be remembered that in
my first contribution to their structure I found the arrangement
of the bones of the palate entirely different in the Nightjars and
true Whip-poor-wills.

STUDIES OF THE MACROCHIRES. 339

As the external characters of these commoner American forms
are well known, and are fully set forth in general works upon
ornithology, I need not introduce them here.

Suffice it to say that these characters fully rank as ordinal
ones in so far as they distinguish these birds from either the
Swifts or the Humming-birds.

When I say ordinal ones I mean as pertaining to an order in
the sense which that division holds as applied to Avian taxonony,
‘and not to other vertebrate classes, where, as we know, structural
differences are far greater than are to be found even among
the extremes in the class Aves.

Having gone carefully over all the literature and material now
available that bears in any way upon the present group, I find
no reason to change my opinion as originally set forth in my
memoir published in the Proc. Zool. Soc. 1885, where I proposed
(p. 914) that all the Caprimulgine birds should be considered
as constituting an order—the order Caprimutcr. I men-
tioned a number of the more doubtful forms that should be
admitted to this order, as Nyctibius, Steatornis, Podargus, and
others. Scarcely a doubt exists now, I think, in regard to the
relation these birds bear to the Owls, through Steatornis, and,
further, they have no particular affinity either with the Humming-
birds nor the Switts.

Their morphology is full of interest, and will repay very careful
research in the future.

In the present connection it is my intention to lead off with a
full description, if the one fine specimen in my possession will
admit of it, of the anatomy of our common American Whip-
poor-will (Antrostomus vociferus), making it comparative with
the more aberrant genus Chordeiles, and then add something
further in regard to the skeleton of Phalenoptilus Nuttalli.

On the Pterylographical tracts of Antrostomus and Chordeiles
(omitting the remiges and rectrices).

Having carefully plucked my specimen of Antrostomus voci-
ferus and one of Chordeiles texensis, and opened before me my
copy of Sclater’s edition of Nitzsch’s ‘Pterylography’ at the
proper page and plate (p. 87, pl. iv. figs. 1 & 2), I am prepared
to present a few remarks upon the pterylosis of the Caprimulgine
birds in my hands.

340 DR. R. W. SHUFELDT’S MORPHOLOGICAL

Commencing with the pterylosis of the head, I find in A. voci-
ferus the same character which Nitzsch points out for the
European Whip-poor-will (see Pl. XVIII. figs. 9 and 10 of this
paper), and that is, on its superior aspect there is a triangular
patch filling in the space just posterior to the superior mandible.
Behind this the feathers form a median longitudinal tract of
some width, which, extending down the back of the neck, as the
dorso-cervical tract, forks between the shoulder-blades. Between
this median tract on the top of the head and within the superior
eyelid, on either side, we find a double longitudinal tract of
contour feathers which join those in front, and posteriorly unite
with the pterylosis of the inferior aspect of the head or the
throat. Apteria occupy the interspaces among these supra-
capital, longitudinal pteryle on the head of this Whip-poor-will,
and as a distinctive feature it is even better marked in our speci-
men of Chordeiles.

It will be remembered that Nitzsch figured this character for
Caprimulgus europeus and Nyctornis grandis, while he states in
the text that he compared these two forms with C. longipennis,
C. forcipatus, and C. psalurus. He also examined pterygraphi-
cally Afigotheles Nova-Hollandie, Podargus gigas, and Nyctornis
ethereus.

On the throat of the Common American Whip-poor-will the
feathers are arranged in fairly well-defined, longitudinal rows,
and Nitzsch found this to be characteristic also of the European
bird; but in Chordeiles these rows are not very easily made out,
if the throat-feathers are inserted upon any definite arrangement,
and I am inclined to believe that in this latter form this is not
the case.

Anteriorly the cervical region is densely feathered in both
Antrostomus and Chordeiles, the tract extending to the points
opposite the clavicular heads of the os furcula, laterally ; while
mesially an aptera occurs of no great extent between the forks
of the bone just alluded to (fig. 9).

Nitzsch found a different state of things in this region in the
European Nightjar, for he draws the entire antero-cervical space
without feathers, which reduces the neck-tracts to two longi-
tudinal, Jateral pteryle*, as shown in his figure of that bird.

The superior mandibular bristles in the Whip-poor-will before
me are conspicuously long, and are deeply inserted as a single

* “ Reather-tracts,” from wrepdy and vAn.

,
————— a F

STUDIES OF THE MACROCHIRES. 341

row just within the margin of the gape. These bristles gradually
increase in length from before backwards, the posterior one
being nearly 4 centimetres long. A few short and straggling
ones are also found in the gular space beneath. In the Night-
jar these bristles are very short, both above and below, and are
by no means a striking feature in this bird, as they certainly are
in the Antrostomus.

Returning to the dorso-spinal tract in the last-named specimen,
we find the extremities of the forks between the shoulder-blades,
already alluded to above, joined by the ends. of a similar but
counter-disposed fork, coming, as it were, up from the lumbar
region. From the apex of this latter the spinal tract appears
to be more or less distinctly divided into two parallel rows, the
median space between them being filled in with less regularly
arranged feathers. Posteriorly the oil-gland stands between
these rows, which slightly diverge as they reach it. This
course of the spinal tract evidently creates a lozenge-shaped
pteryla between the shoulder-blades, and this is even better
marked in Chordeiles. The apteria or “ featherless spaces” on the
dorsal aspect of these birds are very sparsely covered with feathers
to the extent shown in figure 10.

Now Nitzsch found quite a different arrangement of the spinal
tract from this in the European Nightjar, as may be seen from
his figure, and the words of his text, where he says, “ spinal
tract at first broad, forked between the shoulder-blades, each
branch united to the broad rump-band by a single row of con-
tour-feathers.”’

We must, however, recollect that this eminent naturalist also
stated that these tracts differed “‘in the various genera.”

A curious departure is seen in Chordeiles texensis, where, on
either side, a broad tract joins the hinder apex of the lozenge-
shaped dilatation of the spinal tract with the posterior extre-
mity of the ventral band of the corresponding side. The course
of this broad connecting band is directly beneath the “arm-
pit.”

Speaking of the “ oil-gland” in these birds, Nitzsch says it “is
remarkably small, probably the smallest in proportion that
occurs in the whole class of birds; itis of an elongated oval form,
without a circlet of feathers at the tips” (op. cit. p. 87).
This description applies in every particular to the two American
forms of Caprimulgi before me.

342 DR. R. W. SHUFELDT’S MORPHOLOGICAL

As shown in Pl. XVIII. fig. 10, the “femoral tracts” are very
definitely marked in Antrostomus ; they are broader and more
diffuse in Chordeiles. The feathering of the integuments of the
pelvic limbs of either of these genera is somewhat sparse and
scattered, and without definite pattern.

Turning now to the ventral aspect of the body, we find, in
common with what Nitzsch found in Caprimulgus, that in these
American forms the anconal surface of the wings are very thickly
feathered all over; the feathering becomes scattered as we pass
on to the patagium; but the free anterior edge of this latter
membrane has a narrow and dense row of small feathers inserted
along its entire length.

I have already described above the pterylosis of the anterior
cervical region; this leaves us to consider the feathering of the
pectoral aspect of the body. Here we find that both Antrostomus
and Chordeiles agree very closely with the European Night-
jar, as the pterylosis of that bird has been deseribed by Nitzsch.

The ventral tracts in all are broad, broader than the well-
defined humeral tracts of the dorsal aspect, being rather widely
separated in front, and blending somewhat with the aforesaid
humeral tracts at the summit of either shoulder.

The median apterium of the chest (Gf we may apply this term to
it here) is uniformly, though thinly, feathered in both Antrosto-
mus and Chordeiles, which sparse feathering is extended over the
abdomen below. This condition is not taken into consideration
by Nitzsch in his figure of these parts in Caprimulqus. From the
posterior extremity of the broad ventral tract on either side in
the Whip-poor-will and Nightjar we find a narrow contour
abdominal tract running backwards on a curved line to join the
fellow tract of the opposite side behind the vent.

The pelvic limbs are fairly well covered with feathers upon
this aspect, though not so much so as they are upon the reverse
sides.

So much, then, for the pterylography of the Caprimulgine
birds now under consideration. After the detailed way in
which I have attempted to describe it in the above paragraphs,
it will be hardly necessary to present synoptical tables of differ-
ences or similarities to be found in the two American genera
examined. The principal facts to bear in mind are that the ptery-
losis of the American Antrostomus vociferus differs from the

STUDIES OF THE MACROCHIRES. 343

pterylosis of the European bird as described by Nitzsch in a few
well-marked particulars, and that both of these forms again differ
in this particular from Chordeiles, to say nothing of the
further departures which we find when we come to compare
both Whip-poor-wills and Nightjars with such types as Nyctornis
grandis and some others.

No doubt further on we shall find that still more striking dif-
ferences in pterylosis exist among the Caprimulgi and the Swifts
and Humming-birds, to say nothing of what may be discovered
between the last two groups in this regard.

Before concluding what I have to say about this character
in the Caprimulgi, it should be observed that although they
differ among themselves in their pterylography, there is a certain
general similarity of pattern in them all, the fundamental cha-
racters of which are probably well exemplified in our Antro-
stomus, as shown in figs. 9 and 10; while the departures from it
may be easily made clear and apparent by the most superficial
comparison of the several genera, as I have attempted to point
them out or directed attention to those already described by
Nitzsch.

Observations on the Anatomy of Antrostomus apart from the
Skeleton.

(Comparisons with Chordeiles. )

Thanks to the labours of Huxley, Miller, Nitzsch, Macgillivray,

. Cuvier, Garrod, and Forbes, and to an admirable paper by

Mr. Frank E. Beddard, the present Prosector to the Zoological
Society of London (P. Z. 8. 1886, p. 147), much is already known
of the anatomy of the most prominent representatives of the
order Caprimulgi.

In the present connection I shall attempt little more than a
verification of the observations of these trustworthy writers by
dissections of the material I have at hand, and thus fill in the
scheme of my memoir.

First, then, in the two specimens before me, with a scalpel I
carefully remove the integument entirely from the head and
down as far as the root of the neck. This done, the first
thing that strikes us is that we can easily discern the form of the
superior aspect of the brain even through the skull-walls, which
have here been rendered more or less transparent by soaking

344 DR. R. W. SHUFELDT’S MORPHOLOGICAL

in alcohol. It will be observed that the brain of Chordeiles
is considerably larger than the brain of the Whip-poor-will,
notwithstanding the fact that in the latter bird the skull is
markedly wider, longer, and flatter; while in the Nighthawk the
parietal region of the skull is more dome-like and rounded.
The eyes in the Nighthawk are rather larger than they are
in the Whip-poor-will; while in the latter the recurved limbs of
the hyoidean cornua are longer, more median, and reach higher
up on the cranium than they do in Ohordeiles.

Marked differences of course characterize the skulls of these
two forms ; but of this we shall have something to say later: the
inter-ramal layer of muscles is thicker in Chordeiles than it
is in Antrostomus, completely shutting out of sight the hyoidean
apparatus in the former bird, while its form can be easily made
out in the last-named type through this muscular layer.

We need not enter here upon a comparison of the structure of
the neck in these two birds, but proceed at once to remove
the skin from the body and limbs.

On the Mode of Insertion of the Patagial Muscles of the
Pectoral Limb.

These I not only examined in the specimens before me of An-
trostomus and Chordeiles texensis, but in a number of other
species of the latter genus, with the following results. Our
American Whip-poor-will, 1 find, has the tendons of these
patagial muscles of the arm inserted in precisely the same
manner as Garrod found them in Caprimulgus europeus, see
either in his “Collected Memoirs,” or in my copy of his figure
in my “ Contributions to the Anatomy of Geococcyx” (P.Z.S.
1886, p. 471). But it will be as well to mention here that
these tendons are far more slender than one would be led to suppose
from this anatomist’s drawing alone. They are exceedingly
delicate in structure. This remark, however, does not so well
apply to these tendons of the patagial muscles as we find them
in the genus Chordeiles; here they are decidedly broader and
stronger than they are in the Whip-poor-will, and also present
certain well-marked differences. Now, although the plan of
arrangement is essentially the same in the Nighthawks, we find
that the tendon (the main tendon) of the tensor patagii brevis is

STUDIES OF THE MACROCHIRES. 345

evidently composed of two longitudinal slips coming off from the
distal apex of the muscle together, they being but lightly held
together, as they descend towards the muscles of the forearm,
by a delicate connective tissue. The anterior or distal division
of this double tendon is the one which becomes inserted in the
extensor metacarpi radialis longior muscle of the forearm at the
juncture of its tendon and fleshy part. The inner slip of the
main tendon of the tensor patagii brevis, or the slip next to
the humerus, is directed asin the Caprimulgi generally. In other
words, the arrangement here is the same, only the tendon of
the muscle makes it appear somewhat different from the
arrangement in the Whip-poor-will, upon dissection, from the
fact that in the main tendon the two slips are so evidently
distinct. Both of these birds possess the “ bicipital slip,’ shown
by Garrod to be present in the Caprimulgi.

There is yet another point, however, present in the Night-
hawks which I have failed to find in the specimen of Antro-
stomus before me. It is this: when the tendon of the tensor
patagit longus muscle comes to be about opposite the points
where the slips of the tendon of the tensor patagii brevis are
inserted into the structures of the forearm, it sends off a delicate
little tendinous slip which is inserted upon the extensor metacarpi
radialis longior muscle, at the same point where the distal slip
of the tendon of the Jast-named muscle is also inserted, z. e.
at the point of union of its tendon and corneous portion.

On reading over this short description as detailed in the last
few paragraphs, it seems hardly necessary to give any figures to
make my remarks the clearer; it will be well to note, however,
that among the North-American Caprimulei at least—and it
will undoubtedly hold good for the entire group,—the method
of insertion of the slips of the tendons of these patagial muscles
of the arm may differ for the several genera very appreciably, and
on proceeding with my dissections of Antrostomus and Chor-
deiles 1 am the more convinced that, as genera, they are very
well-marked ones.

Of the Pectoral Muscles.

Both in Antrostomus and Chordeiles all three of the pectoral
muscles are present. Pectoralis major and pectoralis secundus

346 DR. R. W. SHUFELDT’S MORPHOLOGICAL

are in each case very well developed, while pectoralis tertius is
quite small and insignificant in comparison even with the second
pectoral. It arises mainly from the shaft of the corresponding
coracoid, and only the extremities of its most posterior-reaching
fibres arise from the sternum, and not as in many other birds,
where a proportionately good share of its bulk may spring from
this last-named bone.

All these pectoral muscles are inserted into the humerus in
a manner common to the great majority of the class Aves, and
require here no special remarks upon that point.

Notes on the Anatomy of the Pelvic Limb.

When examining that group of muscles of the thigh used so
successfully by him in classification, Garrod dissected specimens
of Caprimulgus europeus and Chordeiles texensis, almost identi-
cally the same forms as those before us. In them he found that
they possessed the “femoro-caudal, the semitendinosus, the
accessory semitendinosus, and the postacetabular portion of the
tensor fascie ;? but “the ambiens and the accessory femoro-
caudal are absent.” (Coll. Mem. p. 192.)

My observations tend to confirm these results for the genus
Chordeiles, and enable me to say that the same statement holds
good for Antrostomus; both limbs of the birds before me were
carefully examined, and all the muscles of the thigh dissected
out. I also saw that the main artery of the lmb was the
sciatic, as it is in the majority of birds.

Passing next to the foot, I dissected out the plantar tendons
of both feet in the Whip-poor-will, and the same parts in both feet
of Chordeiles texensis and C. texensis, var Henryt.

Here again I can confirm the observations of Professor
Garrod, who found that in Caprimulgus europeus “the two deep
flexors descend beyond the ankle-joint independently, as usual ;
after passing which, generally about one third down the tarso-
metatarse, they blend completely before any slip has been given
off. From the conjoined tendon thus formed, the tendons of
distribution spring, four in number, one to the hallux and others
to each of the three anteriorly directed toes (see Collected
Memoirs, fig. 4, p. 292), that to the hallux being generally sepa-
rated off before any of the others.” (Op. cit. p. 294.)

It struck me, however, that in Chordeiles the tendon of the

. 2." @ ™

STUDIES OF THE MACROCHIRES. 34:7

flexor longus hallucis enjoyed a greater degree of freedom than
had been found by the anatomist just quoted to exist in Buceros
rhinoceros, which I believe to be as he found it in Caprimulqus ; as
in Ohordeiles, it is easily separable from the deep flexor along its
side, thus approaching somewhat more nearly the condition as
found in Momotus lessoni.

Other Notes.

Cuvier, Nitzsch, and Beddard (P.Z.S. 1886, p. 147) have
all made careful examinations of the syringes of the Capri-
mulgi; and the tracheo-bronchial syringes of Caprimulqus and
Chordeiles are well known. I have investigated this part of the
anatomy of the forms before me, and find they agree in all parti-
culars with the descriptions given by the above authorities ; all of
which will obviate the necessity of my entering upon further
details here. Beddard’s paper, just alluded to, is a real contri-
bution to the anatomy of these parts for the Caprimulgi, and will
well repay reading in the present connection.

Antrostomus has two carotids present, taking the usual course
up the neck in the mid-vertebral canal. This agrees with what is
already known for Caprimulgus and Chordeiles ; and I verified the
fact in the latter bird in the specimen at hand.

Intestinal czeca are present in both Antrostomus and Chordeiles,
being in each case a long slender pair (fully 4 centimetres in
length), and each about one half the calibre of the intestine to
which they are attached.

Upon investigation I find that Antrostomus possesses a small
gall-bladder, while the several species of Chordeiles lack this
orgau: this confirms the observations of Mr. Beddard, who found
that in the latter case Garrod had algo left a MS. note to that
effect (P. Z. S. 1886, p. 151).

The form of the esophagus and stomach is pretty much the
same in both the Whip-poor-will and Nighthawk, although as
regards size it is comparatively larger in the latter bird. I find
it to be a flask-shaped pouch, somewhat compressed from side
to side, with the walls of a fairly uniform thickness, and com-
posed of strongruge. These, commencing at the lateral tendinous
centres at either side, curl round and round in double loop until
they come to the csophageal tube, which they ascend for a short

LINN. JOURN.—ZOOLOGY, VOL. XX. 27

348 DR. R. W. SHUFELDT’S MORPHOLOGICAL

distance and upon which they are gradually lost. These ruge
are so well marked that they may be easily discerned from an
examination of the external surface of the stomach; and upon
the dorsal aspect of the organ they seem to rise into a sort of
transverse ripple, a character present also in the Whip-poor-will.
The cesophagus is of large calibre in these birds, and, as stated,
thick and firm as it approaches the gastric pouch.

The small intestine is delicately constructed, and not especially
large as it passes from the wall of the stomach at a point situ-
ated at the upper right side of the organ, not far from where the
cesophageal tube enters.

Inside the stomach the gastric ruge are covered by a moderate
layer of corneous tissue, composing about one third the thickness
of the stomach-wall, which may best be seen upon a section of
the organ.

Of the Osteology.

For the purposes of classification 1 gave in my first memoir
on the present subject (P. Z.S. 1885) sufficiently full descrip-
tions of the skeleton in specimens of Chordeiles and Phaleno-
ptilus Nuttalli for all that is required in the present connection ;
so it will be only necessary here to make some additional remarks
upon the skeleton as found in my specimen of Antrostomus.
Judging from the figure of the base of the skull of the common
European Nightjar, which I copied from Huxley and repro-
duced in my first memoir, I should say that, osteologically,
the American form of this bird was very much like it in that
particular system of its anatomy; indeed, | expect that struc-
turally the two forms are very similar. Then, as one would
naturally have expected, I have found, upon a mere superficial
comparison, that osteologically the common Whip-poor-will and
Nuttall’s Poor-will (P. Nuttall) are very much more alike than
either of them resemble Chordeiles. In fact, it takes but a
glance at a skeleton of a true Whip-poor-will and a Nighthawk
to convince us at once of the marked differences that exist
between them. As I have elsewhere said, these two genera of
Caprimulgine. birds are separated structurally by very excellent
characters of a nature at once recognizable.

Figures in the plates of my first memoir, above alluded to, also
illustrated the skull of a Chordecles and the principal bones of

STUDIES OF THE MACROCHIRES. » 349

its skeleton ; and for further description these will answer all the
purposes required. Inthe present paper, however, I have thought
it best, for the sake of completeness, to add three figures of the
skull of Nuttall’s Poor-will (P. Nuttalli) in order to show how
well it agrees with Caprimulgus and Antrostomus, and differs from
the skull in Chordetles given in my former memoir (Pl. LIX.
figs. 1, 2, and 4).

Upon more careful and extended examination, I find that,
except in point of size, Phalenoptilus being about one-third less
than <Antrostomus, the skulls, mandibles, and hyoidean appa-

-ratuses of these two forms are very much alike indeed, in all

essential particulars. And as the characters of the skull of the
Whip-poor-will are well known, and, further, as I present here-
with figures in the Plate of the skull of the Poor-will, I believe
that any further comments upon this part of the subject would be
superfluous.

One point, however, in respect to the hyoid. In my former
paper I made the statement that in it the basibranchials in Nut-
tall’s Nightjar were in two pieces. This was true for the speci-
men examined, although in the skeleton of Antrostomus before
me these parts are anchylosed together, which may be the case in
all old birds of both these genera. Chordeiles has them in one
piece; and I am led to believe from this that it will be found
to be generally the case in our N.-American Nightjars.

Passing next to the remainder of the axial skeleton in An-
trostomus, I find my account of the corresponding parts for
Chordeiles and Phalenoptilus Nuttalli (P. Z. 8. 1885, p. 903) to

‘be so complete that it leaves but little here to be added.

Upon carefully re-comparing the axial skeletons of the three
genera Chordeiles, Phalenoptilus, and Antrostomus, now in my
hands, it confirms my previous notions as to their agreements
and disagreements ; and, as one would naturally expect, the
skeletons in the two Whip-poor-wills, or rather the Whip-poor-
will and Nuttall’s Poor-will, are most alike.

The skeleton in a specimen of a Nightjar has already been
described in the place just alluded to; and now I find that Antro-
stomus agrees with Phalenoptilus in having eleven vertebre in
the cervical division of its spine before we come to that which is
the first in the column to have free ribs attached to it. These

ribs in the Poor-will, however, are described as being rather long ;
rag

350 DR. R. W. SHUFELDT’S MORPHOLOGICAL

whereas in the specimen of the Whip-poor-will before me they
are quite rudimentary and small, although they have both head
and tubercle.

For the rest of the vertebral column in these two birds, they
practically agree, both in number and arrangement of the ribs and
vertebre. Their pelves are also very much alike, and wear the
same pattern for general outline, even to the pointed and in-turned
anterior tips of the ilia, which latter feature constitutes a very
excellent character for this bone, at once distinguishing it from
the pelvis of a Chordeiles.

Antrostomus also agrees with the Poor-will in having but five
free vertebre anda pygostyle in the skeleton of its tail; whereas
it will be remembered that the several species of Chordeiles, as a
general rule, have siz and a pygostyle. I have yet to find an
exception to this statement. All three genera seem to possess fen
vertebre in the series that anchylose together in the pelvis.

In Antrostomus in the dorsal series of vertebre, as in all the
Whip-poor-wills and Nightjars which I have examined, the hemal
spines are comparatively long and conspicuous, the anterior ones
being trifurcate at their extremities.

Essentially the form of the sternum in Antrostomus agrees with
the same bone in Phalenoptilus, and the general form it assumes
for the true Caprimulgine birds is very well shown in the figure I
gave of the sternum of Chordeiles texensis in plate 1xi. of my
first memoir, which can be referred to in the present connection.
With three specimens of this bone before us, one being
chosen from each of the three genera in question, they may
be in general distinguished by the following characters :—The
sternum of Chordeiles is the largest of the three, and that of Pha-
lenoptilus the smallest. The “costal processes” in the Whip-
poor-wills are simple erect spines (best marked in Antrostomus) -
whereas in the Nightjar they are more like laterally-compressed
plates, and as we find them in many other birds. All three have
the pair of deep rounded notches in the posterior end of the body
of the bone. They are all without manubrial processes.

The shoulder-girdle in Antrostomus is very like those parts as
I have already described them for Nuttall’s Poor-will, being only
proportionately larger.

Turning, now, to the pectoral and pelvic limbs in this American
Whip-poor-will, we find that they also essentially agree, except

STUDIES OF THE MACROCHIRES. Sot

in point of size, with the corresponding bones and parts in Pha-
lenoptilus Nuttalli, those in Antrostomus of course being the
larger.

With a skeleton of the latter bird now before me, and
carefully reading over my descriptions of the limbs of Chordeiles
and Phalenoptilus, as 1 gave them in my first memoir on this
eroup, I find that there is nothing special to add to that account ;
all the essential characters of these parts being duly presented
for the American forms of Whip-poor-wills and Nightjars.

There is one more statement I made there that seems, how-
ever, to demand correction; for in describing the proximal
phalanx of the index digit of the manus I said of its expanded
portion in Phalenoptilus, that of the two perforations which were
found in it in Chordeiles, they merged in the former bird “ into
one large one.” This is not so; for upon a more extended exa-
- mination I find that there are always two perforations in this part
of the bone in all the forms we have been considering.

This is all I have to state in regard to the descriptive part
of the structure of the Caprimulgine birds of the United States.
Should it become necessary further on to fall back upon this
descriptive part, for the sake of comparison with the remaining
groups yet to be described, it will be done; but, so far as I am
concerned, I am firmly convinced that, taken as a group, including
all other Whip-poor-wills and Nightjars, and such forms as Wyc-
tibius, Psalurus, Steatornis, and Podargus and others, they are
fully entitled to rank as an Order of birds, which I have elsewhere
designated as the CaPRIMULGI.

Not having personally examined such forms as Podargus,
Aigotheles, Nyctidromus*, Batrachostomus, and others, I am
not fully prepared to offer an opinion as to the families and
other divisions of such an Order, nor to state definitely to which
other groups the Caprimulgi are most nearly related; but I can
hardly agree with Prof. Huxley, who asserts that “the Capri-
mulgide come near Zogon, and more remotely approach Po-
dargus and the Owls” (P.Z.S. 1867); for believing, as I do,
that Podargus belongs to the Order, I am also inclined to the
opinion that we shall find that, through Steatornis and Podargus,

* T have since examined skeletons of Nyctidromus albicollis, var. Merrilli
sent me by my collectors in Texas,—R. W S.

352 DR. R. W. SHUFELDT’S MORPHOLOGICAL

the Caprimulgi are nearer the Owls, and only remotely approach
the Trogons.

Again, I can hardly agree with Mr. Beddard*, who would retain
such forms as Antrostomus and Chordeiles in the same “ sub-
family ;” for surely all the essential structural characters of these
two forms are of family and not subfamily rank: a comparison of
the skulls alone is almost sufficient to determine this point. And
the breach between Chordeiles and Steatornis must indeed be
wider than a mere subfamily line can indicate.

Anatomy oF THE NorrH-AMERICAN HIRUNDINIDA.

From my list of material at the beginning of this memoir it
will be seen that I have at hand specimens of every genus and
species of Swallow at present entitled to a place in the United-
States avifauna, and a sufficient series of each to enable me to
fully investigate their structure.

I will take them up, species by species, in the order in which
they occur in the ‘ Check-List’ of the American Ornithologists’
Union, but need not present a synoptical table of their ex-
ternal characters, for these are well known to ornithologists
aud ornithotomists the world over.

To commence with them, then, we will take a look at the ptery-
losis of a specimen of Progne subis, compare it with the figures
given in my Plate of Ampelis cedrorum, and with Nitzsch’s
drawing of the pterylosis of Hirundo urbica in his ‘ Pterylo-
graphy,’ and next with other American Hirundinide.

Now it will be remembered that we found the pterylosis of
Ampelis to agree essentially with most true Passeres, wherein,
upon the dorsal aspect of the body, the chief feature is that the
“spinal tract’ terminates in a lozenge-shaped pteryla situate
mesially between the thighs; and on the ventral aspect we have
another well-known distribution of the pteryle characteristic of
most Passerine birds. Progne differs from all this, and agrees
in the main with Hirundo urbica as figured by Nitzsch.

This author, however, does not present in his work a ventral
view of the pterylosis of a Swallow, but says in his text that
“the single genus Hirundo, which constitutes this group [ Hirun-
dines|, differs more than any other in its habitus from the
general type of the Singing-birds, and in this respect approaches

* P. Z.8. 1886, p. 153.

.
|
|
4
:

STUDIES OF THE MACROCHIRES. 353

very closely to some Cuculine, namely the Cypseli. For this
reason I usually place it at the end of the Passerine, in the
vicinity of the anomalous cuculine form just mentioned, which
stands in the same relation to the true Cuckoos as the Swallows
to the ordinary Song-birds. However, pterylographically, Hz-
rundo does not differ from the rest, but rather harmonizes com-
pletely with Diceum, in that the rows of single contour-feathers
uniting the saddle with the rump-band are either entirely deficient
(AZ. rustica, H. urbica) or indicated only by tworows of very sparse
contour-feathers (H. rupestris). The dilatation of the pectoral
part of the inferior tract is somewhat divergent at the end. The
number of remiges is eighteen, of which nine are on the hand,
and of these the first is the longest ; the first six secondaries are
remarkable on account-of their broad, emarginate extremities ”
(pp. 84, 85, ‘ Pterylography ’).

Now, in Progne I note that the “saddle” at the end of the
spinal tract is very broad, although forked as in Hirundo urbica,
but the posterior extremities of the limbs of this bifurcation are
joined, on either side, to the anterior end of the rump-tract by
distinct and well-marked rows of contour-feathers. Further, the
bifurcation of the “saddle” takes place at about the middle of
the back, and not nearly so low down, namely between the thighs,
as in Hirwndo. Another point to note upon this dorsal aspect
in Progne is that the “alar tracts” are very extensively joined
with the anterior endings of the “ humeral tracts.” In Hirundo
Nitzsch even seems to leave an unfeathered space, on either side,
in these localities. The “capital area” is the same, but in
Progne there are no naked areas around the eye and auricular
orifice, as in Cypselus, and as Nitzsch has also drawn them for
H, urbica. .

Under the throat in Progne and in most Swallows we find a
longitudinal naked strip running down close to and just within
the ramus of the mandible, on either side, which terminates at
about the angle of the jaw. It will be remembered that in the
Whip-poor-wills and others this feature is also present, except
in them it assumes a somewhat different type, the feathers of the
throat being arranged in regular rows. I am inclined to believe
that there isa reason for this, which ig, that in these birds, accus-
tomed as they are sometimes to swallow very large insects, an
operation which must distend the throat, or even momentarily

354 DR. R. W. SHUFELDT’S MORPHOLOGICAL

place the integument there on the stretch, these unfeathered
strips would spread to meet the action, but as the parts came to
rest again after swallowing, the feathered areas or strips would
again become juxtaposed and the throat apparently full-feathered.
In some Swallows (e. g. Chelidon) these naked strips are only
brought fully into view by stretching the integument of the
throat.

No special note is necessary to be taken of the ventral ptery-
losis of Progne, as it has all the essential characters of the pattern
seen in a Passerine bird, and departs but slightly therefrom.
It is more like Cypselus, however, than it is like such a form as
Ampelis, for instance, in that the ventral tract, on either side,
overlying the pectoral region, does not show that heavy feather-
ing to its external margin as seen in the latter type. In Swal-
lows, as in all Passerine birds, the oil-gland is nude.

Now I have plucked, with the greatest possible care, an adult
male specimen of every Swallow in our avifauna, and the birds
are now before me.

In Petrochelidon lunifrons the “rump band” on the back is
very wide, and is joined anteriorly on either side by a very
distinct double line of feathers from the corresponding fork of
the “saddle.” The ventral bands of the pectoral region are
broad but evenly feathered, while on this dorsal aspect the alar
tracts meet and blend with the anterior ends of the “humeral
tracts.” This last feature is invariably the case with all our
Swallows, and is best marked in Clivicola and Stelgidopteryx.

In other particulars Petrochelidon essentially agrees with
Progne in its pterylosis, and with the Hirundinide generally.

Ohelidon likewise has the posterior ends of the saddle-pteryla
of the dorsum joined by feather-rows, one on either side, with
the rump-band, which latter here is narrow again and strictly
defined. Neither this Swallow nor Petrochelidon have naked
annular areas around their eyes, nor the orifices to theirears. In
fact, none of these Swallows possess this last feature. Otherwise,
the pterylosis of Chelidon is characteristically hirundine.

Neither Zachycineta bicolor nor T. thalassina have the bifurea-
tions of the ‘‘saddle-pteryla” of the dorsum joined with the
“rump-band,” as in the foregoing forms, but the ventral tracts
are here again broad and evenly feathered.

In view, then, of the fact that the pterylosis of the Hirwndinide

STUDIES OF THE MACROCHIRES. 355

is pretty well known, it will not be necessary for me to enlarge
further upon my account of it.

But the principal thing to be borne in mind in the present
connection is, that Swallows, Swifts, and Humming-birds all
depart from the more typical pattern of pterylosis found in
true Passeres. And in the case of the Swallows and Swifts, so
far as Nitzsch’s figures and descriptions go, for I have not yet
examined the Cypseli myself for this character, the pterylosis of
the latter is of such a pattern that it requires but very little
modification to make it agree with the pterylosis of a Swallow.
Indeed, in those Swallows where the “saddle-pteryla” of the
dorsum joins its bifurcations with the anterier end of the “ rump-
band,” the pattern is nearly the same, differing principally in
relation, width of the tracts, and position of the bifurcation of
the saddle, which, in Cypselus apus, is between the shoulders.

On the Mode of Insertion of the Patagial Muscles
in the Swallows.

Scarcely any difference is apparent among the various species
of Swallows at hand in regard to the mode of insertion of
this group of patagial muscles, now known to be of so important
a character in the taxonomy of the class. I have carefully ex-
amined them in all the American species, and find that, so
far as the tensor patagii brevis is concerned, both its origin and
insertion seem to be almost typically Passerine. This observa-
tion applies with equal truth to the tensor patagii longus; and as
these muscles are now so well known to all working morpholo-
gists, I need not redescribe them here; moreover, in figure 2
of Plate XVII., I have drawn them for <Ampelis, which will
recall their appearance for the Passeres.

During the course of my dissections upon this region in the
Hirundinide, however, I came across, as I did in Ampelis, what
I am inclined to believe is a hitherto undescribed muscle, at
least so far as Garrod’s descriptions go. It first came to my
notice in a specimen of Progne subis, whereupon I at once dis-
sected a number of other individuals of the same species, and
found it equally well developed in all of them.

This muscle, in part, is a dermal muscle, and arises from the
integuments on the anterior aspect of the neck at about its lower

356 DR. R. W. SHUFELDT’S MORPHOLOGICAL

third; at its origin its fibres spread out fan-fashion, their terminal
ends meeting those of the muscle of the opposite side in the
median line. Here it is quite adherent to the skin, but its fibres
rapidly converge as they pass in the direction of the shoulder-joint,
opposite which region they gradually free themselves from the skin
to form a small fusiform muscle, which, ending in a delicate tendon,
runs along within the free marginal fold of the patagium of the
wing, in common with the tendon of the tensor patagii longus, to
blend with it just before reaching the carpal joint.

I propose to call this muscle the dermo-tensor patagii, it being
partially connected with the integumentary system of muscles in
the birds in which | have thus far found it.

Searching for it in all the other American Swallows, I find it
to be about equally well developed in every species, and absent in
none of them.

This muscle surely does not correspond with the “ bicipital shp
of the patagium,” as described by Garrod, and dwelt upon as the
tensor patagw accessorius by Professor T. Jeffery Parker in his
‘Zootomy’ (1884, p. 251) as occurring in the Common Pigeon,
for it makes no connection whatever with the biceps muscle.

Being desirous at this point of determining its presence or ab-
sence iu a few other groups of birds, I stepped aside for the
moment, and first examined a number of Passerine types, including
very diverse forms,—it was present in all of them. Next, with
the Caprimulgi, Trochili, and Cypseli, I found it completely ab-
sent, as it was also in a specimen of Tyrannus tyrannus, kindly
sent me by Mr. H. K. Coale of Chicago, from which I am led to
infer that it does not occur in the mesomyodian Passeres.
Further than this I did not pursue the subject, but left it for
subsequent investigation and the researches of others interested
in such matters *.

Of the Pectoral Muscles.

Every species of American Swallow has been dissected by
me to ascertain the character and number of these important

* Further opportunities for examining the literature of this subject now
enable me to state that the muscle here described is the “pars propatagialis
musculi cucullaris” of Fiirbringer and Gadow ; and it has been carefully consi-
dered by me in an extensive work upon the muscles of birds now in the hands of
the Smithsonian Institution for publication.—R. W. 8.

STUDIES OF THE MACROCHIRES. 357

chest-muscles as they occur in the group. In every individual
instance I found the state of affairs essentially the same, and
the Swallows agree with all true Passerine birds which I have
thus far examined, in possessing all three of the pectoral mus-
cles. The pectoralis tertius is, comparatively speaking, very
large, and arises nearly or quite as far back on the anterior aspect
of the sternum as the pectoralis secundus does ; it also arises, as
is usual, from the outer side of the shaft of the coracoid bone of
the shoulder-girdle. Pectoralis major makes a very broad and
strong tendinous insertion at the ordinary site upon the shaft of
the humerus, while the tendon of the second pectoral passes
through the usual canal formed by the juxtaposition of the bones
of the shoulder-girdle. Intexture the fibres of the great pectoral
in Swallows seem to be always coarse and of considerable size.

To these characteristics with respect to the pectoral muscles
as I found them in the smaller representatives of the group,
Progne subis forms no exception.

Of the Muscles of the Thigh.

According to Garrod all Passerine birds exhibit, for the classi-
ficatory group of muscles of the thigh, the myolovical formula
A. X. Y (except Dicrurus, wherein it is A. X); 2. e., they possess
the femoro-caudal, the semitendinosus, and the accessory semi-
tendinosus—the accessory femoro-caudal and ambiens being
absent. Upon carefully examining the Swallows, I find that this
is also the rule with them; and these muscles seem to be about
equally well developed in the several genera, although it struck
me that the accessory semitendinosus was, comparatively speak-
ing, rather feebly developed in Progne. Beyond these special
muscles, I did not investigate the myology of the pelvic limb of
these birds.

Notes on the Arterial System.

Swallows, in common with other Passeres, also have but one
carotid artery, the left, which courses up the neck, as usual, in
the hypapophysial channel at the mid-anterior aspect of the cer-
vical vertebre. And in the pelvic limb the main artery I found
to be the sciatic, which is likewise the rule among the Passerine
birds, and Professor Garrod found but few exceptions to this.

358 DR. R. W. SHUFELDT’S MORPHOLOGICAL

On the Prachea, Visceral Anatomy, and other parts.

More for my own satisfaction than with the expectation of
revealing any structure that would prove to be different from
what we already know of the morphology of the trachea or other
parts in the Passerine birds, I examined the wind-pipe, its mus-
cular and associate parts, as I did the several organs in the
chest and abdomen of these American Swallows, but found
nothing that required to be specially noted here.

The trachea exists as we find it in most true Passeres, as do
the several pairs of muscles at its lower larynx. I found the
‘“sterno-tracheales”’ to be very delicately formed indeed, almost
of hair-like proportions in some of the genera, as in Progne.

The gall-bladder is of good size, and the right lobe of the liver
the larger division of that organ.

Czeea coli are present in Swallows, but are of almost rudi-
mentary proportions, and in some cases might be easily over-
looked.

It is my intention to refer to a few of these points again, when
we come to consider the visceral anatomy of the Swifts and
Humming-birds.

The Osteology of the Hirundinide.

Skeletons of representatives of all the Hirundinide of the
United States are before me, and in sufficient number, so that
a general definition for this part of the structure of these birds
becomes quite possible, and will be given here. It is my inten-
tion, however, to be brief in this matter, not only on account of
space, but in view of the information already given.

Of the Skull—When I came to compare and examine the
skulls of our seven species of Swallows, I was surprised to find
them presenting such striking differences in their general form.
Not but that they could each and every one of them be recog-
nized at once as skulls of Swallows, but rather that they possess
characters quite distinct and peculiar to the species, and there
would be no difficulty whatever in telling, for instance, the skull
of a Barn-Swallow from one of a Cliff-Swallow—so diverse is the
general outline of each.

In Progne subis (Pl. XXI. figs. 18, 19, and 20) we find a skull .

STUDIES OF THE MACROCHIRES., 359

that exemplifies all the characteristics which pertain to the
Hirundine skull generally.

Its superior osseous mandible is very broad at the base, but
promptly tapers to a sharp and somewhat depressed tip anteriorly,
while all this portion of the skull is much compressed in the ver-
tical direction. The form of the external narial apertures can
best be appreciated upon a superior aspect, and are seen to be
long, elliptical openings placed longitudinally. Through either
one of them we may discern the upper surface of the anterior
part of the palatine of the corresponding side. The lateral free
edges of this mandible are sharp and turned downwards, while
the maxillary on either side is a horizontal plate fully three times
as broad as the slender jugal bar that continues this infraorbital
rod to the quadrate. We find no projecting processes from the
lateral margins of any part of this osseous superior mandible as
have been erroneously figured for the skull of Progne by other
anatomists (‘ Science,’ N. Y., No. 228, fig. 3). Just anterior to
the frontals, and posterior to the external narial apertures, there
exists a subtriangular area of bone on the top of the mandible,
which is formed by the proximal portion of the premavyillary and
the nasal bone on either side. In the adult skull, of course, the
sutural boundaries of these bones have been absorbed, but by
holding the skull up to the light the proximal end of the pre-
maxillary, and what was the median margin of a nasal, and finally
the anterior limit of the corresponding frontal bone can all be
easily distinguished, while the small triangular space they cir-
cumscribe, is also of bone, but considerably thinner than the
other parts mentioned. In all Swifts that I have examined this
thinner portion on either side has become absorbed, and a little
triangular opening is found at the site instead. My explanation
will be made quite clear by turning to Plate XXI., and com-
paring figures 22 and 23; in figure 23 at «# is shown the
thinned portion, while in the Swift’s skull, figure 22, an opening
actually takes its place on either side. Of course, in a skull so
vastly different from the Cypseline skull as the Humming-bird’s
is, no such comparison as this is necessary.

For the rest of the superior aspect of the skull in Progne we
find the frontal region narrow between the orbital margins, the
posterior edges of which latter are sharp, thin, and somewhat
tilted upwards. The parietal region is smooth and rounded,

360 DR. R. W. SHUFELDT’S MORPHOLOGICAL

while a shallow, mid-longitudinal gutter traverses this part of the
skull (fig. 19). ,

Regarding this part of the skeleton of the Purple Martin upon
a lateral aspect (fig. 18), we are to note the form and compara-
tively large size of the pars plana (p.p.), the slender and rather
small pterygoids, as well as the fact that the osseous interorbital
septum is pierced by two large vacuities of a form in most speci-
mens shown in the drawing. This figure displays so well the
characteristics of the lateral part of the cranium proper in Progze,
that any further account becomes superfluous.

Turning to the base of this skull (fig. 20), we are to note the
form of the vomer and the maxillo-palatines ; the first has very
much the character of that bone as we usually find it in the Passeres.
The maxillo-palatines have their median free extremities dilated,
and they, as in all Swallows, are separated by several millimetres
in the middle line.

The palatines articulate with each other for the posterior two
thirds of their length beneath the sphenoidal rostrum, and are in
close contact at their pterygoidal heads, as in the pterygoids
themselves in this latter locality.

As in all Cypseline birds which I have examined, the posterior
external angles of the palatines in Progne are somewhat drawn
out, and then squarely truncated (compare figs. 19 and 22, pl).
Swallows have the occipital condyle very small, while the foramen
magnum is relatively large, and its plane makes an angle with
the basi-cranial plane of some eight or ten degrees.

Posteriorly, the skull in Progne exhibits a large supra-occipital
eminence, and an occipital area which is nearly circumscribed
by a sharply defined occipital ridge or line, which defines its
form as reniform, and placed transversely at this aspect of the
cranium.

Coming next to the mandible of this bird, we find it to be of a
V-shaped outline, with its ramal sides shallow in the vertical
direction, and with a symphysis of some depth anteriorly at its
apex. There is a swell, on either side, at the superior ramal
margins at points about where the horny theca ceases and the skin
commences, when these latter parts are im situ. A small slit-
like ramal vacuity exists, and the posterior angular processes are
well-developea, though they curve up but very slightly.

Essentially, the hyoidean apparatus is Passerine in character ;

STUDIES OF THE MACROCHIRES. 361

I find, however, that the basibranchials are anchylosed into one
piece, while the glosso-hyal and the cerato-hyals are apparently
not ossified even in the adult Martin.

Several skeletons of Petrochelidon lunifrons have been carefully
prepared by me from specimens of the bird which I collected
a year ago at Fort Wingate, New Mexico, and they are now
at hand. So far as the skull and hyoidean apparatus of this
Swallow are concerned, we might almost cover the ground of our
description by saying that in these parts the bird is the veriest
miniature of Progne ; and, indeed, so true is this, that any detailed
description is rendered quite unnecessary.

Two points it will be well to note, however, for I believe, com-
paratively speaking, the cranial capacity in Petrochelidon is
relatively larger than it is in Progne; and although the palatines
are very much of the same shape, the postero-external angles in
the former are more inclined to be rounded than truncated as they
are in Progne.

Ohelidon erythrogaster in this part of its skeleton probably
typifies the Hirundine skull (Pl. XXI. figs. 21, 25).

In it the superior osseous mandible is very broad at its base, and
the postero-external angles of the maxillaries have a tendency to
project a little. The frontal region is more than usually narrow
between the upper margins of the orbits. Laterally, we note that
the yacuities in the interorbital septum are usually larger than in
other Swallows, though yet but two in number, and of the same
general outline. One thing characteristic of the skull of Chelidon
is its uncommonly minute occipital condyle; I cannot recall at
this moment any bird of the size of this Swallow which possesses
this feature in anything like such diminutive proportions. Its
pterygoids and the quadrato-jugal bars are also wonderfully
slender osseous rods.

Agreeing almost exactly with the mandible in Progne, save in
size, this bone in our Barn-Swallow requires no special mention.
In the hyoidean arches, however, it would seem that ossification
is regularly extended to the glosso-hyal and the cerato-hyals,
which was not the case, as we will remember, in the Martin.

Passing to the genus Tachycineta, we meet with a skull, in
either species representing it (7. bicolor, T. thalassina), which,
although essentially Hirundine in all particulars, yet bears a closer
resemblance to some of our other Oscines, not Swallows, than any

3862 DR. R. W. SHUFELDT’S MORPHOLOGICAL

of the other skulls of the Hirundinide. This is principally du
to the fact that in the skull of Lachycineta the base of the osseou
superior mandible is not nearly so broad in comparison ag it i!
among the other Swallows, and consequently more nearly ap:
proaches in appearance the skull of some of those Passeres which
possess mandibles with rather broad bases.

The structural details seen at the base of the skull in Zach
cineta thalassina I have already shown in a previous memoir,
wherein I have figured those parts in a specimen of that Swallow
(P. Z. 8. 1885, p. 899, fig. F); and as that figure is readily acces-
sible to the reader, a comparison of it with the figures in the
present paper may be made without difficulty.

Nothing worthy of special record is to be found in the man-
dible, nor in the hyoid arches of the skulls or the latter apparatus
in the genus Zachycineta ; they present all the true characteris.
tics of those parts as alread y described above with sufficient fulness
for the Hirundinide generally, and our present purpose.

What I have just said of the skulls and associated parts as found
in the two species of the genus Lachycineta applies with equal
truth to the corresponding structures ag found in Olivicola riparia
and in Stelyidopteryx serripennis, of which I have several examples
of each before me.

In their general form they, too, remind us more of the skulls
of certain other types of Oxcines than do the skulls of the other
Swallows which were described above, previous to our taking up
the skulls of the genus Tachycineta.

Of the remainder of the Axial Skeleton in the Hirundinide.—
My labour is considerably lightened here from the fact that I
have already touched with some degree of fulness upon the axial
skeleton of Tachycineta in my first memoir in the < Proceedings
of the Zoological Society’ (1885, p. 906); and then, again, the
sternum and shoulder-girdle of the Swallows is very well known,
making any detailed account of it here unnecessary.

By those who have read it, it may be remembered that I found
35 vertebre and a pygostyle in the spinal column of a Swift (M-
cropus), and the same number of segments in the column of a
Swallow (Lachycineta). Upon careful examination I am now
enabled to state that this ig the hormal number for all our
Swallows, and I have yet to find an exception toit. Should such
an exception be found, I predict it will simply be a free, and

STUDIES OF THE MACROCHIRES. 363

-erhaps rudimentary, vertebra at the end of the series of the
audal segments.

Further, I find the arrangement of the free vertebral ribs and
“heir uncinate processes the same for all Hirundinide, as I found
them to exist in the Violet-green Swallow in my former memoir,
“his arrangement consists in their having 12 cervical vertebre
‘Mat do not possess free ribs ; the thirteenth has a rudimentary
pair; the fourteenth has them better developed, and even may
have uncinate processes upon them; the fifteenth are the first to
connect by costal ribs with the sternum, as do the ribs from the
sixteenth to the nineteenth vertebre inclusive. The twentieth
is the first vertebra appropriated by the pelvis, and this latter
compound bone monopolizes ten of these segments, so that the
first free caudal is the thirtieth vertebra of the spinal column.

Thus far at least one Swift (dlecropus) was found by me to
exhibit an arrangement similar to this, and later on we may look
into the matter for Ohetura.

The Humming-birds possess, as I have elsewhere stated, but
32 vertebre and a pygostyle in their spinal column.

Every species of our Swallows possesses a pelvis of a pattern
characteristically its own, so that had we before us a dozen pelves
of Progne, a dozen of Chelidon, and a dozen of each of the others
we should have no difficulty, after once becoming acquainted
with them, in picking out the several varieties correctly. Then,
again, these pelves all strictly fall within the general description
applied to what we please to call a Passerine pelvis, so far as our
present knowledge and ideas of such a bone can be formulated.
Now there is nothing that I can at this moment place my finger
upon in the pelvis of a Swift that debars it from being classed in
the same category ; and indeed, when we come to examine into
the matter closely, the differences between the pelves of AZicropus
and Progne are no greater than are the differences between the
pelves of Progne and Chelidon.

Ornithologists have long ago placed on record descriptions of

the shoulder-girdle and sternwn of Hirundine birds, and the

morphology of these parts in them is so well-known that to say,
that although each species of Swallow has a characteristic form of

sternum and shoulder-girdle of its own, these elements of the

skeleton in all of them are strictly Passerine,—will sufficiently
meet our aims in the present connection.
In my memoir in the P. Z. 8. already referred to I made com-

LINN. JOURN.—ZOOLOGY, VOL. xx. 28

364 DR. R. W. SHUFELDT’S MORPHOLOGICAL

parisons of these parts as they occur in Micropus and TLachy-
cineta, and further on, when we come to examine the skeleton of
Chetura, a few more words on the subject may be added.

Of the Skeleton of the Limbs in Swallows.—A]] of the Hirun-
dinide agree with the true Passeres in having the little ossicle
known as the os hwmero-scapuiare at the shoulder-joint, but I
have failed to find it in the Cypseline birds.

In the Proce. Zool. Soc. for April 1887 I figured the humerus of
Tachycineta thalassina, and further on in this article I shall have
to refer to that drawing. Now, so far as the humeri of the other
Swallows are concerned, they all more or less resemble the bone
as found in Zuchycineta: they are invariably non-pneumatic,
proportionately short in the shaft as compared with the size of
the bird, and quite so relatively when taken in comparison with
the Passeres generally. Especially in Chelidon is this brevity of
the humeral shaft noticeable; and it becomes of interest to know
that in a specimen of this Swallow I find a humerus 15 millim.
long to an ulna 24 millim. long, and in Progne a humerus
22 millim. long to an ulna of 33 millim., while in a Swift
(Micropus) we have a humerus 11 millim. long to an ulna of
but 16 millim. in length, showing a difference of 9, 11, and
5 millimetres respectively.

Swallows have at least one good-sized sesamoid at the elbow,
but I thus far have failed to detect any such small bone in a
Swift; in Micropus, however, I find in the same tendon a small
nodule of dense cartilage.

The shafts of both ulna and radius are noticeably straight for
nearly their entire lengths, and in their general conformation
depart but little from the usual form assumed by these bones in
the Passeres at large.

I have already pointed out elsewhere that n a Swift (Micro-
pus) these bones are also markedly straight, and are, com-
paratively speaking, almost as short for a bird of its size as is the
humerus,—Swifts, as a rule, deriving their length of wing from
the long bones of the pinion, and not from those of the brachium
and antibrachium.

Radial and ulnar ossicles are found in the carpus of all
Hirundine birds, as usual, and in their form and method of articu-
lation no departure whatever is made from the composition of
the wrist-joint, as seen in all others of the group.

There are no claws on the finger-end in the manus and

Poa Mig = ene tn WP SE kht ot

STUDIES OF THE MACROCHIRES. 365

phalanges, and the carpo-metacarpal bone is much of the same
shape as we find it in Passeres generally.

To one point I desire to direct special attention, and that is—
that in all Swallows in their carpo-metacarpal bone the meta-
carpal which belongs to the index digit is considerably shorter
than the one which belongs to the annularis digit of this com-
pound bone. This arrangement is strikingly apparent in such a
bird as Progne swbis, and it will be remembered that in Zrochilus
this is also the case, though not so marked; whereas in Swifts
the reverse condition obtains, and the metacarpal of the index
digit is rather the longer of the two.

Little need be said here in regard to the osteology of the
pelvic limb of the Swallows, for from femur to phalanges it is
characteristically Passerine, and in every species the relative
lengths of the several long bones composing it are harmoniously
proportioned. Be it noted, however, that Swallows always
possess a patella, and that in them the pro- and ectocnemial
processes of the tibia are always well developed, while the fibula,
although often of only hair-like proportions (Progne), descends
below the middle point of the shaft of the tibio-tarsus.

Further, in the hypotarsial process of the tarso-metatarsus
there are four perforations for the passage of tendons, these
openings being arranged as though they were at the angles of a
square, one pair being next to the head of the bone, and the
remaining pair immediately behind them.

When I come to review, further on, the characters of the
pectoral and pelvic limbs of certain Swifts and Humming-birds;
it will be necessary to revert again to some of these Hirundine
characters as found in their limbs; and so it will not be necessary
to enter more fully into details at this point, but rather reserve
them for the more effective work of actual comparison.

On THE MornPHOLOGY OF CERTAIN CYPSELI AND TROCHILI.

Of the External Form and Pterylography of certain Oypseline

and Trochiline birds.

Very good hints sometimes as to a bird’s affinities may be
gathered from a study of its general contour and form after
it has been carefully plucked for the purpose. With this in
‘view, and in this way, I prepared specimens of Mcropus
melanoleucus, Chetura pelagica, and Trochilus platycercus, and

28*

366 DR. R. W. SHUFELDT’S MORPHOLOGICAL

present drawings of the same here to illustrate my meaning.
A glance at the contour of Mecropus will be sufficient to con-
vince us that in general outline it is strikingly, indeed actually,
far more like any one of our Swallows, as Progne for instance.
And, apart from the resemblance which its short antibrachium
gives it to Trochilus (Pl. XXIV. fig. 39), it has no other character
upon this aspect of its body to support the view that any true
relationship exists between it aud the latter bird. For the
rest, to my mind, shortness of the antibrachium amounts to
nothing as an indication of affinity unless correlated with actual
similarity of form in its details. Chetura having a deeper
carina to its sternum than has the other Swift, Micropus, it bears
a somewhat more general resemblance to the body of a Humming-
bird (fig. 89) than it otherwise would do, or as does Micropus ;
but some of the smaller Petrels might hold an equal claim to
affinity with Trochilus were it based upon such data as this.

Coming to a few of the true characters, we find the bill, the
position of the commissure of the gape, the feet, and some other
points widely different in a Swift from what the corresponding
characters are in a Humming-bird; and when Wcropus is the
Swift chosen for the comparison, the entire contour of its body
differs from that of Zrochilus in all important particulars.

Let us next examine the pterylography of these three birds,
and see what it indicates in regard to their affinity.

Nitzsch has presented us with fairly good figures of the
pterylography of Cypselus apus and Trochilus moschitus (Pterylog.,
ed. Sclater, pl. ii. figs. 16-19); but there are several points
requiring elaboration in his account, while in other particulars
his comparisons are deficient.

Taking his figures and descriptions just as they stand, how-
ever, and bringing into the discussion his figure 14 on the same
plate, of Hirundo urbica, we find that the pteryloses of the Swift
and Swallow, so far as their heads are concerned, agree, with the
exception that the Swift possesses those peculiar crescent-shaped
apteria, one over each eye; theseare absent both in the Swallow
and Humming-bird.

But the Humming-birds have a median naked space of a
spindle-shaped outline on the crown, situated longitudinally, and
between the eyes and the base of the superior mandible. This
is well marked in all species which I have thus far examined,
and it was overlooked by Nitzsch; moreover, it is absent in
the Swifts and Swallows.

STUDIES OF THE MACROCHIRES. 367

On the throat of Swifts and Swallows the feathering covers
the entire area, while in Humming-birds the median naked space
of the neck is continued almost up to the base of the inferior
mandible.

Again, Nitzsch noticed the naked “ nape-space [see his figure |
beneath the long cornua of the hyoid bone,” but “ could not deter-
mine with precision” whether or no it was a constant character
for the pterylography in the Trochili. My investigations con-
vince me that it is a constant character in them, and, further,
that itis never present in the Swifts nor Swallows. If any one will
take the trouble to pluck a Humming-bird and note, in the
natural position of its head, that the back of the head comes
very close to the body, he will see at once how this naked space
has come to be present there,

The arrangement of the pteryle upon the ventral aspects of
all of these birds is more or less alike, being apparent modifica-
tions of some Passerine type; but not so with the spaces upon
the dorsal aspects, for here we find that the true differences
among them come in (compare Nitzsch’s figures). And we must
remember that Nitzsch, in speaking of the pterylography of the
Macrochires, was forced to admit that :—‘“‘ In this family I place
the two genera OCypselus and Trochilus, which, indeed, present
but little external similarity, but are very nearly allied in the
structure of their wings” (p. 86). To the near alliance on
account of the latter character we will revert later on.

In the first part of this memoir I have attempted to point out
such differences as exist between the pterylography of a Swift
and a Swallow, so it will not be necessary to enter so fully upon
the details again here. Be it borne in mind, however, that,
upon this dorsal aspect of the two, in both the humeral tract
crosses obliquely at a point opposite the middle of the humerus
of the arm; in Trochilus, on the other hand, 7 is over the
head of the humerus. Swifts and Swallows both possess a femoral
tract; whereas it is absent as a rule (and, for all that I know
to the contrary, always) in the Humming-birds—certainly so in
Trochilus.

In Swifts the “ spinal tract’ connects the capital area behind
with the oil-gland, but just opposite the shoulder-joints bifur-
cates ; the bifurcations are as wide as the original tract, and
after passing the middle of the back they converge again, and
unite at a point over the anterior end of the sacrum. Thus we

868 DR. R. W. SHUFELDT’S MORPHOLOGICAL

find a spindle-formed figure produced, which is characteristic of
the Cypseli.

In Swallows the bifurcation does not take place until the
spinal tract arrives nearly at the middle of the back, and then the
ends of the fork fail to join the rump-tract below.

Now in Humming-birds, and I have examined a great many
excellent specimens of them, the “spinal tract” is altogether
different from this, for it consists of a very broad, lozenge-shaped
figure, spreading out over nearly the entire dorsal region, being
prolonged in a wide nuchal strip which merges with the “ capital
area” anteriorly, while its lower angle rests upon the uropygial
gland, and laterally spreads over the femoral region. Mesially,
and in the middle of this lozenge-shaped area, we have a short
longitudinal naked strip, but not nearly so conspicuous as it
is in the Swifts.

Indeed, the pterylography of a true Cypselus and Trochilus is
as different in character as any two forms of birds can well be in
this particular; and if one, unprejudiced in mind,-will look at
plate ii. of Nitzsch’s work, there will be seen a greater similarity
between the dorsal tracts of Cypselus apus and Coracina cepha-
loptera than between Cypselus apus and Trochilus moschitus.

We are already aware that, notwithstanding Swifts and
Humming-birds possess the same number of primaries and rec-
trices, it rather conveys the impression that this is more a matter
of chance, when we find that they essentially differ in their ptery-
lography and in the number of secondaries in their wings.

For another external character in the Swifts, and a very
excellent one, which I have failed to find elsewhere described,
we must turn to the integuments covering the pinion. Here we
find the entire skin exclusive of the border surrounding this part
of the limb, and on both sides, of a deep black colour, being pro-
duced by a pigmentary deposit in the skin itself. This peculiar
character is present both in Micropus and Chetura, while it
is entirely absent in Zrochilus. Swallows also lack this pig-
mentary deposit in the skin on both surfaces of the pinion.

To conclude this chapter, then, I will make a few comparisons
between the external forms and characters of Mdicropus—a true
Swift—and Trochilus platycercus—a typical Humming-bird.

So far as the general form of these two birds is concerned, a
glance at Pl. XXIV. figs. 37 and 39, will be sufficient to convince
any one that they are as different as they can well be.

iden...

STUDIES OF THE MACROCHIRES. 369

In the character of their beaks they are as widely different as
any two types in the entire class Aves.

They differ essentially in their pteryloses, and in the number
of the secondaries.

Their fect are radically different, quite as different, for instance,
as are the feet of a Swift and a Sparrow-Hawk.

The majority of these differences in these two types are abso-
lutely of an ordinal rank (for Aves).

And now, before entering upon their internal structure,
let me add here the well-known fact that these birds also
differ essentially in their habits, their mode of nidification,
and the manner of securing their food; indeed, in all these
particulars in their life history they are widely, very widely
different.

A critical Comparison of the Pectoral Limbs of certain Cypseli
and Trochili.

From time immemorial in Ornithology the two main charac-
ters upon which systematists have relied for retaining the
Oypseli and Trochili in the same group of birds, as related forms,
are the supposed similarity of the structure of their wings, and
the fact that both possess an unnotched sternum. Finding that
these birds widely disagree in so many vital, fundamental par-
ticulars, it is my object to compare them very critically with
respect to their wing-structure, and the present section will be
devoted to the results of my investigations in that direction.
Swallows, as we know, possess a wing-structure very similar in
organization to the Passeres generally, so it will not be necessary
to make many comparisons with them in the same connection.
We have just seen how essentially different the wing of Trochilus
is from the wing of MWicropus, so far as its external characters
are concerned: to be sure they have a superficial resemblance, as
both have short humeri and long pinions, but this resemblance
gives way when we come to compare the parts in detail.

First, then, let us examine the method of attachment of the
patagial muscles, surely a character which has proved itself to
be a useful one, and one eminently connected with the wing-
structure in birds, be they Swifts or Humming-birds. Now Prof.
Garrod dissected a Humming-bird with the view of ascertaining

_ the point which concerns us here, and he had a specimen of

Patagona gigas for investigation. Moreover he made a drawing

370 DR. R. W. SHUFELDT’S MORPHOLOGICAL

of his dissection of the parts in question, and it may be seen in
figure 1, plate xxiv. of his ‘ Collected Scientific Memoirs.’

With the exception of leaving off the lower extensor of the
forearm, his drawing is correct, and from it we see that the
tensor patagit longus arises and is inserted pretty much as we
find it in most birds; but with respect to the tensor patagi
brevis a very marked departure is met with, for that muscle is
as prominent as any other in the arm, more so than the majority
of them. It may be said to be somewhat pear-shaped in form,
with its larger end at the origin at the shoulder, while the smaller
extremity becomes attached to a tendon which passes directly
over the upper surface of the extensor metacarpi radialis longior,
longitudinally.

This tendon arises at the outer condyle of the humerus, and
passes to the carpus for insertion, and is very well shown in
Garrod’s drawing of Patagona.

I find it present in all the Trochili, where, so far as I know,
it constitutes a unique method of insertion for the tensor patagi
brevis, and to make it clearer I present a drawing of it for
Trochilus platycercus (Pl. XXII. fig. 28).

Since Garrod saw so clearly this very unusual insertion of the
tensor patagit brevis in the Humming-birds, I am surprised
beyond measure that he did not at once make careful comparisons
with the Cypseli in this particular; had he done so, he would
have found, as I have, that the mode of insertion of this muscle
in those birds is entirely different. In the first place the body
of the muscle is comparatively much smaller; it is also of a very
different form, being oblong and not pear-shaped ; finally it zs not
inserted into any special tendon, but directly upon a tendinous
fascia on the surface of the extensor metacarpi radialis longior,
and its fibres, becoming slightly tendinous, run down with that
muscle for insertion at the external condyle of the humerus.
In Pl. XXII. fig. 29, I present a drawing made directly
from my dissection of these parts in a specimen of Chetura
pelagica.

As both the Humming-birds and Swifts have short humeri
(though “shortness”? is not a character, I believe) and have
developed a large tensor patagii brevis (though “size” is not a
character either, I believe) it might not unnaturally be expected
that they should have this particular muscle short and thick;
but when we come to examine the true morphology, how vastly

STUDIES OF THE MACROCHIRES. 371

different is it! Quite as different, we may say in truth, as are
the humeri of these birds.

The tensor patagii longus in Chetura pelagica has the usual
origin and insertion that it has in so many of the Class.

Cypseli and Trochili both possess all three pectoral muscles,
but in such a form as Micropus they are none of them unduly
developed ; better so in Chetwra; while in the Trochili they are,
comparatively speaking, enormously developed.

Owing to the entirely different shape of the humerus in Swifts
and Humming-birds, the tendons of the pectorals make
dissimilar insertions. For instance, the pectoralis major in
Micropus is inserted upon the entire palmar aspect of the large
hook-like radial crest of the humerus of that Swift; but Zrochilus
possessing no such process upon its humerus, the muscle is
obliged to insert itself more or less upon the body of the bone,
at a point which would be considered as the base, upon the
palmar side of a radial process did such a thing exist there.

Now the pectoralis secundus in Micropus is inserted at the
head of the humerus upon its anconal side, between the summit
and radial crest or hook; while in the Humming-bird this second
pectoral sends its tendon across the head of the bone, to be
inserted at the distal margin of the pneumatic fossa. The
insertion of the third pectoral in these two groups of birds is
more similar.

So here, again, we see that Swifts and Humming-birds are
markedly different with respect to another class of muscles which
make up, in part, the fundamental organization of their wing-
structures.

Among the essential characters of the wing we still have left
the skeleton, but I have already published my views and drawings
in regard to that part of their economy elsewhere (Proc. Zool.
Soc. 1885 and 1887). I have there shown conclusively that
the humeri of Swifts and Humming-birds are very differently
formed bones indeed, and the reader has but to refer to the
figures in the papers to which I allude to be convinced upon
this point.

As I have elsewhere stated, the humerus in MWicropus is a
non-pneumatic bone as in the Swallows ; while all Humming-birds,
so far as I have examined, have pneumatic humeri. Still
my statement Proc. (Zool. Soc. 1887, p. 503) requires some
modification, for since that was written I have found that the

372 DR. R. W. SHUFELDT’S MORPHOLOGICAL

humeri in Chetura pelagica are pneumatic, but the bone is
shaped upon the same plan as the humerus of Micropus, and the
pneumatic fossa is, as in Passeres, on the ulnar side. From
what has gone before, we now know that in general form, and
other particulars, /icropus is nearer the Swallows than is such
a Swift as Chetura, and this last fact, with respect to the arm-
bones, points still more strongly to the truth of such a state-
ment. Even at this moment I am not acquainted with any other
bird in the Class that has the pneumatic fossa of its humerus
situated on the radial side of the bone, as the Trochili have.
This fact alone, and surely when taken in connection with the
otherwise vastly different form of the bone itself, is sufficient to
show that in their wing-structure Swifts and Humming-birds
widely differ.

Further, in the papers above alluded to I have already
pointed out how in the bones of the antibrachium, in Zro-
chilus and Micropus, the radius is actually bent to a bow in the
former, while it is as absolutely straight as any bone can be in
the Swift. The ulna, too, in these birds differs in its general
form. Moreover, we find sesamoids present in the carpus of
Humming-birds which do not exist in Cypseli, although, since
writing my first memoir on this subject, I] have found a sesamoid
at the elbow in Chetura and Micropus, such as the Swallows have.

Coming next to the carpo-metacarpus we find one great
and principal difference, in addition to minor ones—in the Hum-
ming-birds the middle metacarpal in this compound bone is
longer than the index metacarpal, the reverse condition obtain-
ing among the Swifts. This is enough to show that the bones
are essentially unlike in their most important character. The
proximal phalanx of the index finger is altogether a differently
formed bone from the corresponding segment in the manus
of the Swift, as any one may see by a comparison either of the
bones themselves or my drawings (P. Z. 8. 1885, pl. Ixi. figs. 3
and 4,7).

To briefly recapitulate, then, the absolutely essential and
fundamental characters in the wing-structure of a Swift and a
Humming-bird, I find that:—1. The parts markedly differ in
their external characters, inasmuch ag they do not possess the
same number of secondary quill-feathers ; Swifts have a very
peculiar pigmented (deep black) area of the skin centrally located
on both sides of the hand, while Trochili have not; the character

STUDIES OF THE MACROCHIRES, 373

of the plumage is quite different; and the position of the “humeral
tract” in the pterylosis is different, being across the middle of
the humerus in Swifts, and overlying the head of the bone in
Humming-birds. 2. The mode of insertion of the patagial muscles,
as well as the form and character of these muscles themselves, is
altogether different in the two groups. 38. The method of inser-
tion of the pectoral muscles is essentially different. 4. Through-
out the entire skeleton of this limb, the individual bones in Swifts
and Humming-birds differ widely in characters of the very highest
import, both morphologically and in the position, absence, and
presence of parts.

All this being so, | am firmly convinced that were the minor
details in structure in these two wings carefully worked out
under the lens of a good microscope, they too, of necessity, would
also be found to be at variance. Indeed, in making my own
dissections of the Trochili under a 2-inch objective I saw quite
enough to fully confirm this suspicion.

Finally, I must say, as 1 have already remarked in a previous
paragraph, that heretofore too much stress has been laid upon
the fact that both Cypseli and Trochili possess short humert;
and, further, to my mind, shortness, per se, does not constitute a
valid character, for if it did, some very remarkable forms would
surely be grouped together! My painstaking labours upon the
wing-structure of Swifts and Humming-birds convince me fully
that, in so far as this part of their organization is concerned,
there is little or no affinity at all.

Notes on the Anatomy of the Pelvic Limb in certain
Cypseli and Trochili.

Having shown how innately different the wing-structure in
Swifts and Humming-birds really is, let us now take a look at
their pelvic limbs.

It will not be necessary to pass the external characters of these
parts in review, as they are already well known; it will be suf-
ficient to remark that the pelvic limb of such a bird as Wicropus
differs from the pelvic limb of a Zrochilus in all its more essential
external characters.

My investigations tend to confirm the statement of Pro-
fessor Garrod, that Humming-birds and the American Swifts
Chaetura pelagica and Micropus lack the accessory femoro-

374 DR. R. W. SHUFELDT’S MORPHOLOGICAL

caudal, the semitendinosus, and the accessory semitendinosus
muscles from the group at the thigh ; in other words, their formula
is A. .

This from a physiological point of view would naturally be
looked for, as no members of these groups use their limbs for loco-
motory purposes; and consequently these special muscles have
long since been missing, or perhaps in neither of them bave they
ever been present. But to this matter I shall refer further on.

Coming next to the plantar tendons, I find the arrangement in

the Swifts at hand the same as described by Garrod for Cypselus
alpinus (Coll. Scient. Mem. p. 294), and as that has already
been made clear to us, I need not quote it here; but after having
carefully prepared the foot of a specimen of Trochilus platycercus,
and bringing the limb under the lens of a powerful objective,
which increased the size of this Humming-bird’s foot to that of a
Crow, I was enabled at once to discover that the arrangement of
the plantar tendons in these birds is very different from what —
obtains in the Cypseli; in other words, in Trochilus these tendons
are disposed very much as we find them in the Passeres, where the
tendon of the flexor longus hallucis is distinct from that of the
flexor perforans digitorum. It is just possible that in Humming-
birds a shght vinculum may connect the two, and although I
could not quite satisfactorily demonstrate this minor point, yet
I am inclined to think that such a vinculum is present.

I found the sczatic artery the main artery of the leg in both
Cypseli and Trochili, but that is the usual arrangement for
nearly all birds, which weakens its importance as a distinctive
character.

As to the skeleton of this limb in these birds I have already
contributed some work (P. Z. 8S. 1885, pp. 909-918), and little
or nothing need be added here. Suffice it to say that morpho-
logically the constitution of the pelvic limb, so far as its skeleton
is concerned, is radically different in Cypseli and Trochili. A
few points will be sufficient to convince any one of this fact,
for in Trochilus, for instance, we have a large patella present, a-
bone entirely missing in Micropus; in Trochilus we have the
hypotarsial process of the tarso-metatarsus both pierced and
grooved for the passage of the tendons, whereas in Micropus it
simply exhibits one deep groove for that purpose; finally, the foot
in each case is widely different, for in Zrochilus the joints of pes

STUDIES OF THE MACROCHIRES. 375

stand 2, 3, 4, 5, while, as we know, in Micropus they stand 2, 3,
3, 3.

As existing birds are classified, and were two such forms as
Miecropus and Trochilus classified upon the characters presented
in their pelvic limbs alone, all I can say is, that to my mind
there should be no hesitation whatever in placing them in widely
separated groups, notwithstanding the fact that the myological
formula of the thigh-muscles is the same. For even when we
come to examine these very muscles closely we soon discover
that they are quite differently formed and disposed, which should
also be taken into consideration in face of the fact of the mere
presence or absence of parts.

For the rest, the limb in these two groups of birds to its
very toe-joints is about as essentially different as are the limbs
of an Ostrich and a Coot.

On the Anatomy of the Head.

Were I asked to pick out any two forms of existing birds from
any part of the world which present us with the greatest
number of fundamental differences so far as the anatomy of
the head is concerned, it would puzzle me, I think, to select
two more diverse types than a true Swift and a Humming-bird.
Indeed, from tip of beak to nape it is difficult to find comparable
characters that show any affinity of the forms in question at all.
I have already pointed out above the very evident differences that
are exhibited upon a comparison of the external characters of ©
such a Swift as Micropus and any of the Trochili; while the
principal differences in the skulls of these birds have been already
dwelt upon*. And has the day yet arrived when differences
of the most manifest character in the skulls of birds are to be
ignored in taxonomy, and set aside as of no value?

At the present time I have before me upwards of a hundred
anatomical specimens of Trochili and a great many Swifts ; but
for a brief résumé of some of the distinctive cranial characters let
us choose a specimen each of Chetura pelagica and Trochilus
rufus, and see how they compare in these two types. We find
these characters to be as follow :—

* Proc. Zool. Soc. 1885.

376 DR. R. W. SHUFELDT’S MORPHOLOGICAL

Chetura.

1. Superior mandible wide and not
produced.

2. Triangular openings between nasals
and frontals, divided by the pre-
maxillary,

3. Cranium
rounded.

above smooth and

Vomer truncated.

. Maxillo-palatines prominent and
produced well backwards, tending
to approach mesially.

6. Postero-external angles of palatines

produced as prominent processes.

7. Palatine heads of pterygoids nearly
meet mesially.

8. Pars plana small and formed as in
Swallows.

9. Interorbital septum shows several
vacuities, and these are distinct from
those on the posterior orbital wall.

10. Mandible a wide V, without ramal
vacuity.

Trochilus.

1. Superior mandible narrow and
usually twice as long as the head.
2. No such openings present.

3. Cranium above showing a deep,
longitudinal groove for ends of
hyoid.

4. Vomer long and spine-like.

5. Maxillo-palatines not prominent,
rounded, and wide apart.

6. External margin of each palatine
nearly straight, and no angle
present.

7. Palatine heads of pterygoids widely
separated mesially (and I have seen
specimens where they anchylosed to
the palatines).

8. Pars plana very large, and very
different from the Swallows.

9. Interorbital septum never shows
but one vacuity, which merges with
one that absorbs nearly all the
posterior orbital wall.

10. Mandible a long and extremely
narrow V, with ramal vacuity.

In short, these skulls evidently belong to very different Orders
of birds, and their differences upon a lateral view can be well
appreciated by examining and comparing figures 24 and 27 of
Plate XXII. ; the Swift there figured, however, is Micropus, but
will answer just as well.

Carefully comparing the brain in several specimens of Hum-
ming-birds of different species, with the brains of Swifts and
Swallows, I find that, although in all three groups the brain
and its parts are strictly fashioned upon the true avian plan, in
the Swifts and Swallows its general and special form is far
more alike than it is when we compare it with the brain in a
Trochilus. This we might naturally have looked for, since
the inner shape of the cranial casket in the Humming-bird is
very different from the corresponding cavity in the Cypseli and
Hirundines.

Another structure which need not detain us long is the tongue.

STUDIES OF THE MACROCHIRES. 377

This organ is essentially alike in Swallows and Swifts ; while, as
we all know, in the Trochili it is more as we find it in the Wood-
peckers, indeed very similar to those birds, for I find after
careful microscopical examination that there is no truth in a
statement still current that this long, slender tongue of Trochilus
is a double-barrelled tube to suck honey with, but these supposed
hollow tubes contain the prolongations of the cartilaginous parts
of the glosso-hyal elements of the hyoidean apparatus.

With these few brief comparisons, which, however, are the
expressions of long and painstaking dissections upon the heads
of these several forms, I may state that, so far as this part of the
economy is concerned, Cypseli and Trochili are widely different
in all particulars, whereas Swifts show themselves to be but

highly modified Hirundine birds.

Résumé of some of the Points in the remainder of the
Axial Skeleton.

These I will tabulate in order to bring them into as bold
relief as possible for direct comparison. In the Proc. Zool. Soe.
1885, I have already made some remarks upon the skeletons
of Micropus melanoleucus and Trochilus Alexandri. ere, for
variety’s sake, we will take the Swift Chetura pelagica and
Trochilus rufus; they are essentially and respectively much
alike, atany rate the two first mentioned species, but I do this in
order to show that my first comparisons still hold good for the
proposed separate groups.

Chetura pelagica.
1. 12 cervical vertebrx that are with-
out free ribs; 13th and 14th ver-

Trochilus rufus.
1. 13 cervical vertebre that are with-
out free ribs ; only the 14th vertebra

tebrae possess freely suspended ribs;
while from the 15th to the 19th
they are true dorsals, connecting
with the sternum by costal ribs.

possesses freely suspended ribs; while
the 15th, 16th, and 17th are the
only three free vertebrx in the dor-
sal region which connect with the
sternum by costal ribs. The 18th
and 19th likewise do; but I here
propose to consider these two latter
ones as leading sacrals, as they evi-
dently belong to that bone. This
gives Trochili but three true dorsal
vertebré, quite as few as any other
existing bird, and it is all they have.

2. The last sacral vertebra is the 29th. 2. The last sacral vertebra is the 27th.

378 DR. R. W. SHUFELDT’S MORPHOLOGICAL

Chetura pelagica. Trochilus rufus.
3. The last caudal vertebra is the 35th. 3. The last caudal vertebra is the 32nd.
4. Pelvis much as we find it insome 4, Pelvis peculiarly formed; and two
Swallows; leading sacral vertebra entire vertebra project beyond the
does not markedly project beyond ilia (the 18th and 19th).

ilia.

5, Sternum untouched posteriorly; 5. Sternum unnotched posteriorly ;
possesses comparatively large costal very small costal processes; no
processes ; small manubrium; deep manubrium ; comparatively a much
carina; which latterand the body are deeper carina; sternal body and
always viddled with large vacuities. keel never perforated by vacnities.

6. Os furcula a very broad U-shaped 6. Os furcula rather of a very broad
one, with lateral abutments at its V-shaped variety, with small lateral
heads, and with rudimentary hypo- abutments at its heads, and with
cleidium ; the bone harmoniously rudimentary hypocleidium, with
proportioned for the rest of the the bone of hair-like dimensions as
skeleton. compared with others of the skele-

ton.

=~]

7. Coracoids much of the same form . Coracoids very peculiar, as the
as we find them in the Swallows. tendinal canal is closed by bone,
and the shaft perforated by a large
foramen below it. Totally unlike
the bone in the Cypseli.
8. Blade of scapula nearly straight. 8. Blade of scapula bent at a marked
angle at its posterior extremity.
9. General aspect of the body skele- 9. General aspect of the body skeleton

ton, aside from the unnotched ster- has no exact counterpart among
num and rather deep keel to it, living birds, that the writer has as
like the Hirundinide. yet ever met with.

Now a few words as to what the above table shows: first, it is
evident that the spinal column of Swifts and Humming-birds is
fundamentally different, both in the number and arrangement
of the vertebre. It should, however, be stated that upon going
over a large number of specimens, I find that it is the 15th
vertebra that first connects with the sternum by costal ribs, and
not the 16th as stated in my first contribution of 1885. This
gives the Trochili 3 true dorsals, which is as small a number
as any existing bird possesses; I found the same number ina
Californian Condor. Cypseli possess 5 true dorsal vertebre.

Some excellent characters, no doubt, are to be obtained from
any bird’s sternum, but the more I look into it the more I am
convinced that the facility with which we can say sternum
2-notched, sternum unnotched, sternum 4-notched (as the case
may be) has almost proved a detriment to avian taxonomy, for,

¢- STUDIES OF THE MACROCHIRES. 379

being satisfied with that (taken in connection with a few other
salient characters), very often the rest of the bird’s economy
has not been examined nor even taken into consideration at all.
Why the pelvis has not proved an equally valuable character
in the list of classificatory characters, is simply because the
systematist cannot so readily say pelvis 2-notched, pelvis un-
notched, and so on. Yet the pelves of birds, when carefully
compared, offer fully as good distinctive characters for taxo-
. nomic purposes as the sternum. I have already pointed out the
fact that the pelvis of a Zrochilus is as different from the pelvis
_ of a Oypselus as any two birds’ pelves can well be. Further, their
sterna, when we really take all their characters into considera-
* tion, apart from the fact that both happen to be unnotched, are
“very differently fashioned bones. Both are unnotched, to be
sure,—but so are the sterna of some Petrels! Were the fact that
the sterna of both Cypseli and Trochili are unnotched of any signi-
ficance, so far as affinity is concerned, then surely the remainder of
the organization in these birds would be more or Jess in harmony,
and not at the widest variance, as is the case! What I mean by this
is easily shown in the shoulder-girdles of the two types in question:
thus, the coracoid of a Trochilus is a very uniquely-formed bone
(P. Z. 8. 1885, pl. lx. fig. 5), and very different from the great
majority of birds. In the Swifts the coracoid is like that of the
Swallows. Again, the scapula in Trochilus is unlike the corre-
sponding bone in a Swift: consequently, this being the case, I
attach little or no importance, so far as affinity is concerned,
to the fact that their furcule happen to possess some marked
resemblance. For we well know that this latter component of
the girdle is that which becomes modified in accordance with
the flight of its owner, while the coracoid can be far better
relied upon for any affinity it may show as a character amongst
forms more or less related. Swifts are birds of long-sustained
flight, Humming-birds are great fliers, and so are Albatrosses; and
were we to increase in size the os furcula of a Swift and a Hum-
ming-bird to the size of the bone in an Albatross, we should be
surprised to find how much they resemble each other.

Seeing now how very different the thoracic and pelvic, or
really the trunk-skeletons of Swifts and Humming-birds actually
are, let us next examine into some of the organs and viscera
which they enclose.

LINN. JOURN.—ZOOLOGY, VOL. Xx. 29

380 DR. R. W. SHUFELDT’S MORPHOLOGICAL

The Heart and Carotids, Trachea, Viscera, Sc.

Cypseli as a rule possess but a single carotid, the left one ;
Professor Garrod, however, discovered that Cypseloides proved
an exception to this. In Chetura I found but one, which was
disposed along the anterior aspect of the neck in the most usual
manner; while in Micropus melanoleucus the left carotid, here
also the only one present, takes on a peculiar course, for being
so far over to the left, it passes up to the front of the neck
obliquely, and completely outside the protection of the muscles
and the hypophysial canal of the vertebre.

Past the middle point of the neck, however, it enters between
the muscles to the aforesaid canal, and then follows the usual
course to the head.

Swifts do not possess a heart of any unusual dimensions; but
Humming-birds, on the other hand, have a heart quite as unpro-
portionately large for their size as are the feet of these, the other-
wise pygmies of the Class. They too have but one carotid, so
far as I have examined, the left one alone being represented.

MacGillivray, in Audubon’s ‘ Birds of North America,’ under
the latter’s account of Trochilus colubris, presents us with a very
good description of the tracheain a Humming-bird. He says of
it that “The trachea is 9 twelfths long, being thus remarkably
short on account of its bifureating very high on the neck, for if it
were to divide at the usual place, or just anteriorly to the base of
the heart, it would be 43 twelfths longer. In this respect it differs
from that of all other birds examined, with the exception of the
Roseate Spoonbill (Platalea ajaja), the trachea of which is in so
far similar. The bronchi are exactly 3 inch in length. Until
the bifurcation, the trachea passes along the right side, after-
wards directly in front. There are 50 rings to the fork; and
each bronchus has 84 rings. The breadth of the trachea at the
upper part is scarcely more than 3 twelfth, and at the lower part
considerably less. It is much flattened, and the rings are very
narrow, cartilaginous, and placed widely apart. The bronchial
rings are similar, and differ from those of most birds in being
complete. The two bronchi lie in contact for 2 twelfths at the
upper part, being connected by acommon membrane. The lateral
muscles are extremely slender. ‘The last ring of the trachea is
four times the breadth of the rest, and has on each side a large
but not very prominent mass of muscular fibres, inserted into the
first bronchial ring. This mass does not seem to be divisible

STUDIES OF THE MACROCHIRES. 381

into four distinct muscles, but rather to resemble that of the Fly-
catchers, although nothing certain can be stated on this point.”
My own investigations upon other species than 7. colubris go
towards establishing in the main this admirable description of a
very painstaking anatomist, for whom I have always entertained
the highest regard both for his character and his work. It is
needless to add that such a trachea, the counterpart of which is
seen only in the Spoonbill, is sufficiently far removed from the
form it assumes in the Cypseli to satisfy the most sceptical as
to any affinity on that point! In Swifts it does bifurcate “at the
usual place;” it possesses but ¢wo pairs of muscles (the lateral

- ones, and those that go tc the sternum), and in all other points

is widely and fundamentally at variance with the windpipe and
bronchi of the Trochili.

Careful as MacGillivray’s account is, however, he neglected
to mention one very important difference, so far as these parts
are concerned in the birds under consideration, and that is, the
Trochili constitute one of those rare groups which lack the pair
of sterno-tracheales muscles; I carefully searched for them in
several species of Humming-birds, but failed to find them, and
am quite convinced they do not exist.

If the reader will kindly turn to figure 33 of Plate XXIII.
illustrating this memoir, he will find my drawing of the trachea
of a Humming-bird, and in figure 35 the position it occupies in
the thorax and neck with respect to the other organs.

Indeed, in figures 35 and 386 I have drawn the bodies of a
Humming-bird and a Swift, after having carefully removed the
pectoral muscles and sternum, in order to show this very thing.
A glance at these two figures will be sufficient to satisfy any one
as to the remarkable difference they present. In the Humming-
bird, we are struck at once by the position of the trachea; the
direct course of the left carotid, the enormous heart, and the fact
that the low position of the liver conceals from our sight all the
other viscera harboured in the abdominal cavity. Here, as in
most birds, the right lobe of the liver is the larger of the two,
which in the Humming-bird, as we see, curls round the apex of the
heart (more so in Z, platycercus), modelling itself to that extre-
mity of it. Still more at variance, as compared with the Swift,
is the digestive tract of a Humming-bird, for, so far as I am fami-
liar with the morphology of the group, in none of them do I know
of a species which possesses, as compared with the size of its
intestines, so exceedingly small a stomach! This organ, together

29*

382 DR. R. W. SHUFELDT’S MORPHOLOGICAL

with the relatively large intestine, with, too, its bulbous cloaca, I
have represented in figure 34.

Swifts possess a stomach, both in position and general form,
very much like the Swallows, and, as we now know, nothing
at all like the Trochili. True, neither Cypseli nor Trochili pos-
sess intestinal ceca ; but does this mean anything when no other
two organs in the bodies of these birds have any resemblance to
each other whatever, so far as affinity is concerned? Look at
them in the figures; are there many birds in the Class more
widely separated in this respect than these Swifts and Humming-
birds P

Upon laying open the stomach of a specimen of Micropus
melanoleucus, I found it packed full of insects ; but, what is more
important, anatomically speaking, [ discovered it to be lined
with a tough, corneous, inner coat, which was lifted out entire,
by simply using very gentle traction, with a pair of dissecting-
forceps. The stomach of the Humming-bird was also full of the
tiniest Coleoptera imaginable, which were very interesting to
study under a two-inch objective attached to my Beck’s binocular
microscope, and I wondered as I did so whether all these tiny
New-Mexican beetles were known to science.

Apart from the fact, then, that Cypseli and Trochili agree in
certain numerical and negative characters (“a single carotid,
and no ceca,” dangerous facts sometimes !), these birds are by
no means related, so far as the organs we have just been inves-
tigating are concerned.

Having now passed in review the characters of a Passerine
bird (Ampelis cedrorum), and gone very carefully over the
osteology of certain Trogons, and even yet more thoroughly
over the structure of many Caprimulgi, Swallows, Swifts, and
Humming-birds, I believe, as my views have been slowly for-
mulating during my painstaking dissections, that I am now in
a position to reconsider what I have already published upon
the classification of the MacrocuiRrEs, as well as to present the
conclusions at which I have now arrived, aided as I have been
by all this recent research. Before doing this, however, I desire
to present in a few paragraphs the results of my investigations
upon two specimens of 7. Calliope, nestlings only a day or two
old, and for which I am indebted to the generosity of Mr. F.
Stephens, of San Bernardino, California, who sent them to
me to be used in the present connection. One of these little
fellows I drew, life-size, and it will be found figured on Pl. XXIII.

STUDIES OF THE MACROCHIRES. 383

fie. 32, which givesits external characters sufficiently well to obviate
the necessity of aspecial description. Among the most interest-
ing of these features is the wonderfully short beak in this
nestling, as compared with the long slender one of the adult.

Supplementary Notes on Cypseloides niger and Nyctidromus
albicollis, var. Merrilli.

As this paper is passing through the press I am able to add a
few words upon the structure of these two birds—the Black Swift
and Merrill’s Parauque. This affords me particular satisfaction,
for inasmuch as every species of American (7. e. United States)
Swallow (seven in all) is anatomically described in this memoir,
I can add that I have similarly examined and compared every
species of Caprimulgine (except A. carolinensis) and Cypseline
bird. I am indebted to my friend Professor Newton, F.R.S., of
Cambridge, for the specimens of Oypseloides, which were collected
for him on my behalf in Jamaica by Mr. G. A. Waddington.
The specimens of Nyctidromus are from Texas, where they were
procured on the lower Rio Grande by two of my collectors.
Externally Cypseloides niger has a more Swallow-like appear-
ance than either Wicropus or Chetura. This no doubt is due to
the structure of the tail and feet, which have a more passerine
appearance than is seen in JZ. melanoleucus, and still more so
than in OC. pelagica or C. Vaux. Nevertheless Cypseloides is a
Swift, with the pterylography of the order as given above. It
also exhibits the peculiar black pigmentation on the palmar
aspects of its pinions, although the skin there is not quite so
dark as in other North-American Cypseli. The tarsal and pedal
integuments are skinny, but plainly show a scutellate definition.
The hind toe is somewhat elevated, though distinctly posterior
in position. In general form the plucked body presents the
appearance of the nude body of a Ohetura rather than of Micro-
pus, which is more compressed in shape.

Myologically, this Swift agrees with others already described,
the patagial muscles, the muscles of the thigh, and thorax being
almost identical with those of C. pelagica.

Upon opening the abdominal cavity we find that in these
parts also Oypseloides agrees with all true Swifts. The stomach
is notably large, and only overlapped by the lobes of the liver
above, in all these respects differing widely from the corre-
sponding organs in any existing Humming-bird.

384 DR. R. W. SHUFELDT’S MORPHOLOGICAL

In the anatomy of its air-passages, its heart and vascular
system, this Black Swift is likewise typically Cypseline.

Coming to the skeleton, I find Cypseloides in its osteology
agrees in the main with the group of birds to which it naturally
belongs ; that is, it is essentially a Swift so far as this part of
its organization seems to indicate; nevertheless, in several
particulars it has a skeleton nearer the Swallows than has either
Micropus or Chetura. It has, for instance, the interorbital
septum much as we find it in the Hirundinide generally, and a
large sesamoid at the elbow, as in Swallows. But, what is still
more significant, it has the vacuities, one on each side of the
posterior mid-end of the premaxillary above, just beyond the
frontal region, filled in by a thin continuous layer of bone—
agreeing in this particular respect with the Barn-Swallow (C.
erythrogaster). Cypseloides, moreover, has its external narial
apertures more circumscribed, or, in other words, more as we find
theim in certain Hirundines (see figures 22 and 28, Plate XX1I.).

Having compared the skeleton of Myctidromus albicollis
var. Merrilli with the skeletons of the other Caprimulgine
birds of the U.S. avifauna which I have described on former
occasions, I find that it agrees more nearly with the American
Whip-poor-will (Antrostomus vociferus) than with any other.
Osteologically, however, it may be found to agree still more
closely with the ‘“‘ Chuck-will’s- widow ” (A. carolinensis), but as
yet I have not had the opportunity of comparing it with that
bird.

The entire order of the Caprimutar stands much in need of
thorough revision, and extensive researches into structure will
be required before we can know much of the true relation-
ships and proper classification. Iam convinced that, so far as
the United-States forms of this group of birds are concerned, there
are certainly two very well-defined subfamilies of the Capri-
mulgide. From what we know of their external characters, and
from what I have shown of their widely different internal
structures, these might readily be characterized as the sub-
families Antrostomine and Chordeiline—the former to contain
the genera Antrostomus, Phalenoptilus, and Nyctidromus; the
latter the genus Chordeiles.

We have but to compare the skull of Nuttall’s Poor-will (2.
Nuttalli, Plate XX.) with the skull of Chordeiles acutipennis
var. texensis (P. Z. 8. 1885, pl. lix.) to be convinced of the wide
differences which exist in this part of the skeleton in these two

STUDIES OF THE MACROCHIRES. 385

very distinct kinds of Goatsuckers, and this, as we now know, is
sustained by other parts of the structure of the birds in question.
In this connection, however, | may add that I have recently
examined a nearly adult specimen of Chordeiles virginianus,
kindly procured for me by Dr. W. S. Strode of Bernadotte,
Illinois. In this I find that the maxillo-palatines do not meet in
the median line, but are pressed close against the sides of the
vomer on each side. This latter bone is bifurcated behind, and
into the fork the antero-median point of the palatines is wedged.
The vomer comes well forward, anteriorly, where it is bluntly
pointed and thicker thanit is behind. It is only in the immature
bird that these true relations can be studied, for in all species of
this genus, as they attain to maturity, these several bones indis-
tinguishably fuse, and present the appearance shown in the basal
view of the skull of Chordecles acutipennis var. texensis (P. Z.S.
1885, pl. lix. fig. 4), where, however, the vomer is not quite
correctly indicated, for the lines designated by Vo go to the
mesial fused portion of the palatines, and not to the vomer,
which in that skull is co-ossified with the maxillo-palatines, and
only its median line and anterior apex are seen.

Anatomical Notes upon the Nestling Trochilus,
a day or two old.

First, 1 remove the delicate skin from the specimen’s head,
and note that the ends of the hyotdean apparatus have not
proceeded beyond the posterior area of the parietal region, and
that, although the tongue is short, still it shows well the embryonic
condition of the two glosso-hyoidean rods which become go long
in the adult Humming-bird.

The nasal bones lap rather high up on the frontal region, and
mesially meet the backward-extending limb of the premaxillary
for their entire borders, thus leaving no vacuity in this locality,
as is to be seen in the postero-culmenar space of the superior
aspect of the upper mandible in an adult Cypselus.

In size, the lacrymal bones are exceeding small, and I am in-
clined to think that were we able to define their sutural bounda-
ries in the skull of the adult, we should find that they contribute
but a meagre share to the wide expanse of bone in the pars plana
of the mature T’rochilus.

At the base of the skull we note that the tiny palatines, the
jugals, quadrato-jugals, and even pterygoids are now considerably
ossified; and that the latter elements are separated at their

386 DR. R. W. SHUFELDT’S MORPHOLOGICAL

palatine heads quite as much in proportion as we find them in ~
adult skulls.

The premaxillary and mandible are also largely formed in
bone, more especially their tips and backward-extending limbs.

Removing the skin from the back, I carefully count the ver-
tebre of the column two or three times, distinguishing 35 seg-
ments, from which we may judge that 3 vertebre are incor-
porated in the pygostyle of the adult.

Without any difficulty whatever, and by the aid of a 2-inch
objective, I clearly make out the arrangement of the muscles of
the fore limb, and distinctly perceive the tendon into which the
tensor patagii brevis is inserted. Even still better can be seen
the muscles of the thigh, where the biceps seems to arise by a
double head from the pelvis, but otherwise the myological for-
mula here is the same as I stated it above for the adult Trochilus.
The plantar tendons also confirm all that is recorded in a pre-
ceding paragraph.

Coming next to the sternum, I find that even at this tender
age the posterior margin of the body of the bone is rounded and
unnotched. Six costal ribs articulate, on either side, with a
“ costal border.”

At the side of the neck in this specimen the cesophagus was
much distended by a small spider and two small beetles; but I
believe that this represents food that the little bird had not
swallowed at the time of its death, and that naturally no enlarge-
ment takes place in the cesophagus at the point in question.

We note that the bifurcation of the trachea is situated fuliy
halfway up the neck towards the throat in this nestling, so that
if the upper moiety of the anterior cervical region happens to be
covered with the finger at the time of microscopical examina-
tion, one is momentarily impressed with the notion that the
bird has two tracheex, so unusual is this arrangement in the Class
Aves.

Upon opening the thorax and abdomen, it disclosed the fact
that the sterno-laterales muscles of the trachea are not present,
and I am inclined to believe that Zrochili do not possess them.
Further, we find the heart is in about the same position and
relative size as it is in the adult; but the lobes of the liver are
proportionately much smaller, so much so that we can easily
examine the intestines and stomach below their hinder borders
without disturbing them, which is not possible in the adult.

On the other hand, the stomach is proportionately much larger

|
|
|

STUDIES OF THE MACROCHIRES. 387

in this nestling than it is in the parent bird (to be of any use, it
could scarcely be of a relative size), and in the present case
was crammed full of insects.

CONCLUSIONS.

Before touching upon the real object of the present memoir
as stated in its title, in these my final conclusions, I will briefly
allude to what may be gathered from my investigations as set
forth in the earlier sections of this paper, touching the morpho-
logy of representatives of certain outlying groups to the Macro-
cHtrEs. At the outset, believing it would be an advantage to
pass in review the structure of a suitable and average Oscinine
bird, I chose Ampelis cedrorwm for reasons already fully stated ;
and, in addition to the advantage of having its structural charac-
ters before us in the present connection, my brief account of
its anatomy, it is to be hoped, will prove useful in other par-
ticulars, more especially in throwing some light upon its own
probable relations to the Clamatorial birds and the Hirun-
dines.

It is believed that the account tends to show that struc-
turally Ampelis presents no special aflinity with the Swallows,
while in some respects it links the Mesomyodian birds with
the Oscines, though nearly all its entire organization points to
its more intimate relations with the latter group.

Judging from osteological premisses alone, it is very evident
that such forms as Zvogon puella and 7. mexicanus can claim
no special relationship with the Zrochili, while, on the other
hand, I consider that their affinity with the Caprimulgi is also
very remote. Further than this their kinship at present con-
cerns us not, as it does not especially bear upon the work in
hand; nor, even were I so disposed, would I hazard an opinion
in any such direction, until I had fully investigated the struc-
ture of other birds specimens of which, up to the present time,
it has not been my good fortune to possess, nor, in many
instances, even to see. How much Cuckoo stock they possess
in their economy is another point which can only be settled,
if ever, by exhaustive researches into the anatomy of the more
aberrant Cuculine types; it is more probable that they, the
Trogons, came up through some such tribe as the latter, than
through any other with which I am acquainted.

Still, and to hold this end of the thread for a moment
longer, it is difficult to see any near relationship between such

388 DR. R. W. SHUFELDT’S MORPHOLOGICAL

a bird as Trogon puella and Geococcyx californianus for instance.
Surely there must be a gap of no mean width when we come
to push them in that direction. Not long ago I published
(Proc. Zool. Soc. for 1887) some contributions to the anatomy
of Geococcyx, wherein, in my conclusions, I pointed out what
appeared to be the correct classification of the United States
Cuckoos, supporting Garrod’s original suggestion of placing the
true Cuckoos and Ground-Cuckoos in separate subfamilies. Still
maintaining, as I do, this opinion, I am free to confess that
I consider the subfamilies to be thus represented markedly
distinct, to say the least of it. For instance, how close to such a
bird as Coccyzus minor may Geococcyx be? Notwithstanding
the zygodactyle foot in the latter (a character sometimes of
uncertain meaning), I have always entertained the notion that
some day we may see an affinity between Geococcyx and the Da-
celonine, asin Dacelo gigantea; or, carrying it a little further,
a certain kinship with the Galbulide, more particularly those
which possess the two carotid arteries and the myological
formula A.XY.~ But here, again, the proper material has never
yet been at my disposal.

Since the appearance of my first contribution to the present
subject (P. Z. 8. 1885), nothing has arisen in the course of
my more extended researches which has in any way modified
my original opinion in regard to the Caprimulgine birds, nor
have I anything to add to what I have already stated in the body
of the present memoir. They constitute the first group which I
propose to remove from the old Order Picarta, and for them I
create a separate Order, the Caprimutet, first alluded to in my
previous paper, to contain all the true Caprimulgine birds of the
world, including such types as Steatornis, Podargus, digotheles,
Nyctidromus, Nyctibius, Psalurus, and others.

These birds have their nearest kin in the Owls, and they have
no special affinity with the Cypseli, much less with the Trochili.
With our present knowledge of their structure, these Caprimul-
gine forms may easily be relegated within this Order to their
proper family and subfamily positions, and in a way, too, I think,
that would meet the approval of all, as it would be based entirely
upon the structural characteristics of the several and respective
types, the best and only guide in such matters.

Coming now to the Hirundinide, 1 see in these fissirostral
Oscinine Passeresa group of birds, which, although they still possess
in their organization a majority of the structural characters of

STUDIES OF THE MACROCHIRES. 389

the original Passerine stock, have long since deviated from the
latter. They are true Passeres considerably modified, which
modifications in several instances may be traced to the adoption of
new habits, and are really physiological adaptations of structure.

For instance, through ages of time they no doubt have gradually
_attained their increase of size in the gape, which enables them
to take insect prey upou the wing with greater certainty and
ease, and at the present time constitutes one of their best
distinctional characters.

Whatever may be the physical principle involved that seems
to demand a brevity of the brachium to suit their flight, we cer-
tainly can now perceive that a proportional shortening of the
humerus is going on. Their flight, however, is not of such a
‘vigorous nature as yet to demand much increase in the size of
their pectoral muscles, and the consequent deepening of the carina
of the sternum for their due attachment, nor the unnotched body
of that bone to afford a more stable surface for the origin of
those muscles. But with the present configuration of the coun-
tries they inhabit, and where their food is now to be had in
abundance, no doubt they will long retain their present habits,
and consequently their present structural organization.

Suppose, however, at some time in the world’s history, ages ago,
there were certain large areas inhabited by the original Hirundine
stock, sufficiently differentiated from the existing Passerine types,
in which from some cause there was a diminished supply of
natural food—the insects which they had been accustomed to take
on the wing. This would at once seem to demand in the organi-
zation of the Swallows an increased rapidity of flight, in order
to secure for themselves and their young sufficient food during
the course of the day. It would also lead, perhaps, to an increase
in the size of the mouth, that this food migit be captured with
greater certainty. J*urther, they would probably be obliged to
remain longer upon the wing. Continued for a sufficient length
of time, such causes would be sure to work structural changes
in the economy of these birds, and modifications would in con-
sequence follow in their wing-structure, in the size and strength
of ‘their pectoral muscles, with an increase of the bony surface
of the sternum, both in body and carina, from which these latter
muscles arise; and finally, among certain other minor changes,
we might find in consequence of the last-named requirement a
suppression of parts in the feet and certain muscles of the pelvic
limb, as the owners would now rarely perch or walk.

390 DR. R. W. SHUFELDT’S MORPHOLOGICAL

And this is the way, I suspect, that certain forms which we
now see in our modern Swifts were differentiated from the early
Hirundine stock. That this occurred early in the chapter of
avian life-history, for the world is old, may be conjectured
from the fact that Cypseli{are now quite cosmopolitan birds,
and, moreover, have many representatives among them which
present highly specialized organization. Even at the present
time, however, we yet have forms that structurally are nearer
the Swallows than others of the same group. To instance this,
we have but to glance at two such birds as Micropus mela-
noleucus and Chetura pelagica, in the first of which we still find
the general Swallow-like form of the body, the average depth of
the carina of the sternum, the non-pneumatic humerus, and other
points, all of which are far more Cypseline in character in the
latter bird. Chetura, too, agreeing with other spine-tail Swifts,
shows its greater fixedness of characters in the very structures
which gives it its name, for the spines which terminate its rec-
trices are useful to the bird, yet can only have been developed
through ages of time. When we come to examine the still more
Swallow-like Swifts, Hemiprogne for example, and its allies, I am
sure we shall meet with other points in their anatomy which will
lend support to this view of the origin of these types.

In the present memoir I have, by extensive and careful com-
parative investigations into structure, attempted to point out
how entirely different these Swifts are from the Humming-
birds, a group with which they have long been associated, to
my mind upon very meagre claims. During the course of my
present researches I have shown that Cypseli differ from
Trochili, (1) in their habits; (2) in their nidification; (8) in
the method of securing their food; (4) in all their external
characters, and markedly in their external form; (5) in their
pterylosis; (6) fundamentally in their skeletons ; (7) every struc-
ture in their heads is as widely at variance as any two forms
of birds in the Class; (8) in their wing-structures; (9) in their
pelvic limbs; (10) in their respiratory apparatus ; (11) in their
visceral anatomy; and (12) in their digestive system. These
two groups have been associated together upon an entirely false
system of classification, which assumed first, that they are alike
in their wing-structure—a resemblance which I have shown to
be purely superficial ; secondly, that they both have an unnotched
sternum, although physiological law demands it, and when asso-
ciated with an entire organization that widely differs from that

mene

STUDIES OF THE MACROCHIRES, 391

of another form which may happen to possess an unnotched
sternum, it means nothing so far as affinity is concerned. This
becomes the more evident when the sterna themselves are
fashioned upon essentially different plans, as is the case in the
Cypseli and Trochili.

Truly related organizations never exhibit such an array of in-
harmoniously associated sets of morphological characters. And
it is to the detriment of comparative anatomy, and all we may hope
to effect by it, to summon to our aid such characters as “ short-
ness” (in the case of the humeri), as ‘‘ presence ” or “absence ” of
parts (as intestinal cece), and other matters of purely physical or
arithmetical interest, unless there can be shown in connection
therewith actual similarity in form and arrangement of parts.

Now in my first memoir (Proc. Zool. Soc. 1885) upon this
subject, I proposed that in the Passeres the Cypseli should be
placed next to the Hirundinide ; for convinced, as I was, of their
relationship, [ for the moment did not take into account the
artificial boundary lines of orders, genera, and what not, demanded
on the part of systematists, simply having in my mind (after
working many weeks over their several structures) their affinities,
and not how they really ought to figure in print.

Evidently this will not do, and we must assign them some
position in the system which they can occupy with propriety in
ornithological works, even if it does a little violence to the delicate
and intricate kinships, which the morphologist can so often see
with his mind’s eye, but which sometimes look so startling in
type.

There is but one way at present open to us to effect this, and —
that is, all the true Swifts in the world must have a group or an
order created for them, as the order Cyrsreri, which I now pro-
pose for their reception. This Order, were it represented by a
circle, would be found just outside the enormous Passerine circle,
but tangent to a point in its periphery opposite the Swallows,
which latter are to be found just over the line of the are.

For the Trocuitt I have already proposed a separate order in
a former communication, and am to-day more convinced than
ever of the correctness of that proposal.

The time may arrive when we shall see more clearly the rela-
tionship to other groups of birds of these markedly modified and
highly interesting little forms, but in the meantime a very great
amount of painstaking dissections upon avian types will have
to be successfully undertaken. Agreeing with the Psittaci in this

392 DR. R. W. SHUFELDT’S MORPHOLOGICAL

particular, I am inclined to believe that the order Trocurm1 will
be found to be an unusually well-circumscribed one, containing
upwards of 500 species, to represent it.

Since completing the main part of this paper, and especially
since closing the lst of acknowledgments at its commencement,
I have received many kind letters relative to the work from
fellow labourers in the same fields, and in some cases valuable
material for comparison.

Chief among these it gives me great pleasure to thank Professor
W. K. Parker, F.R.S., for many timely hints upon avian rela-
tionships, and for his ready encouragement of my work during
the time it has been in progress. I am grateful, too, to Sir
Edward Newton, C.M.G., formerly of the Colonial Office, of
Kingston, Jamaica, for his efforts to secure me specimens of
Hemiprogne zonatus; to Lieut. Edgar A. Mearns, of the
Medical Corps of the United States Army, for specimens of
Humming-birds from Arizona; to Mr. Robert Ridgway fer
having directed that the entire collection of birds in alcohol at
the Smithsonian Institution should be gone over with the view
of filling up gaps in my desiderata, although at that time it was
found that no specimens in alcohol of the Macrochires were in
the collections of that Institution ; and finally, to Mr. F. Stephens,
of San Bernardino, California, for the loan of many valuable sterna
of American Trochili, from his private collections.

EXPLANATION OF THE PLATES.
(All the figures in the Plates were drawn by the Author from the specimens.)

Puate XVII.
The pterylosis of Azpelis cedrorum.

Fig. 1. a. Ventral aspect. 6. Dorsal aspect. Considerably reduced.

2. Muscles of the patagium of the right wing in Ampelis cedrorum, seen
upon the outer aspect, and x 2. tp. J, tensor patagii longus; Zp. 0,
tensor patagii brevis; dt. p, dermo-tensor patagii; d, deltoid; ¢,
triceps ; 0, biceps; ¢. m. 7. 7, extensor metacarpi radialis eee h,
humerus; w, ulna.

. Right isteral view of the skull of Tyrannus eee 6; life-size.
1, the free lacrymal bone.

4. Same view of the skull of Ampelis cedrorum, 3; life-size: letters the
same.

. Same view of the skull of Hesperocichia nevia, g; life-size.

(Sw)

STUDIES OF THE MACROCHIRES. 3938

Fig. 6. Under view of the skull of Ampelis cedrorum, $; X 2. pmx, pre-

maxillary ; mx. p, maxillo-palatine ; pp. pars plana; pt, pterygoid ;
ju, jugal; g, quadrate; 8, foramen for vagus nerve; 9, for the
hypoglossal nerve; ¢. c, for the internal carotid; g. 7, quadrato-juga. ,
ew, Eustachian tube ; p/, palatine ; mx, maxillary ; v, vomer.

Puate XVIII.

Fig. 7. Pelvis of Ampelis cedrorum, X 2; dorsal aspect.
8. Basal view of the skull of Trogon mexicanus, x 2; the mandible re-

Fig.

moved. Lettering as in the preceding Plates, with 7. s, nasal septum ;
x calls attention to the basipterygoid process of the right side.

9. Anterior aspect of the body of Antrostomus vociferus, to show the

10.

23.

pterylosis. Somewhat reduced.
The same, shown from behind.

Puate XIX.

. Superior aspect of the skull of Trogon mexicanus ; life-size, with

mandible removed.

. Ventral or anterior aspect of the sternum of the same species; life-

size.

. Right lateral view of the skeleton of the same; life-size, with the ribs

of the left side removed.

. Dorsal aspect of the pelvis of the same ; natural size.

Puate XX.

. Left lateral view of the skull of Phalenoptilus Nuttaili,x2. Collected

by the author at Fort Wingate, N. Mexico, Lettering of the parts as
on Plate XVII.

. The same skull seen from above, x 2; mandible removed.
. The same skull viewed upon its basal aspect ; mandible removed ; x 2,

Puatr XXI.

. Right lateral view of the skull of a specimen of Progne subis, 9; x 2.

Lettering of the parts as before.

. The same skull seen from above, X 2; mandible removed.
. The same skull viewed upon basal aspect, x 2; mandible removed.
. Basal aspect of the skull of Chelidon erythrogaster, 8, x 2; mandible

removed.

. Superior view of the skull of Micropus melanoleucus, ¢, x 2; man-

dible removed: and letters as before. This drawing is made from
the same skull as the one from which I drew the basal view in a
former memoir on the Macrochires (P.Z. S. 1885, p. 899, fig. D).

Superior view of the skull of Chelidon erythrogaster, 3, X 2; man-
dible removed : letters as before. This is the upper view of the skull
shown in figure 21 ; and x directs attention to the thinning of the bone
in the triangular area on either side, between the nasal, frontal, and

premaxillary ; in the Swift (fig. 22) this entire triangular area becomes
completely perforate.

394:

Fig. 30.
. Ventral aspect of the pelvis of Trochilus rufus, x 33.
. Right lateral view, life-size, of a day-or-two-old nestling of Trochilus

Fig. 37.
38.
39.

MORPHOLOGICAL STUDIES OF THE MACROCHIRES.

Puate XXIi. Fi

. Right lateral view of the skull of Micropus melanoleucus, 6; X ?

Lettering of parts as before. This is the same specimen from whic.
I drew the basal view in a former memoir on the Maerochires (P. Z. 8.
1885, p. 899, fig. D).

. Right lateral view (x 2) of the skull of Tachycineta thalassina, 3.

Lettering of parts as before.

26. Superior view (x 2) ofthe skull of Tachycineta thalassina, $; mandible

removed. ‘This figure and figure 25 refer to the same specimen from
which I drew the figure in my former memoir (P. Z. 8. 1885, p. 899,
fig. F).

. Right lateral view of the skull and mandible of Trochilus rufus, adult

3; x4. Same lettering as before.

. Outer aspect of the muscles of the right arm in Trochilus platycercus.

Very much enlarged. ¢, triceps; ¢p. 0, tensor patagii brevis; tp. J,
tensor patagii longus; ¢. m. 7. /, extensor metacarpi radialis longus ;
tz, a tendon to the tensor patagii brevis.

. Outer aspect of the muscles of the right arm in Chetura pelagica.

Enlarged rather more than twice. Lettering same as in fig. 28.
This figure and the last were drawn by the author directly from his
own dissections.

Puate XXIII.

Ventral aspect of the pelvis of Micropus melanoleucus, X 2.

Calliope.

. Anterior aspect of the trachea of Trochilus rufus, x 44.
. Digestive tract of Trochilus platycercus, x 2%.
. Anterior aspect of Trochilus Calliope, adult, with the chest-wall and

other parts removed to show the relative size and position of organs.
l. c, left carotid; H, heart; r. 7, right lobe of liver; J. Z, left lobe of
liver. x 22.

. Same view and similar dissection of Micropus “melanoleucus. S,

stomach, with other lettering as in figure 35. Somewhat enlarged.

Puatr XXIV.
Left lateral view of a plucked specimen of Micropus melanoleucus.
The same of Chetura pelagica.
The same of Trochilus platycercus.

These are all life-size figures of male birds obtained by careful contour traces
directly from the bodies of the specimens, and are not intended to show any
part of the pterylography.

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STUDIES OF THE MACROCHIRES.

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STUDIES OF THE MACROCHIRES.

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MR. G. B. SOWERBY ON NEW SPECIES OF SHELLS. 395

Descriptions of fourteen new Species of Shells.
By G. B. Sowrrsy, F.L.S., F.Z.S.
[Read 20th December, 1888. ]
(Puatn XXV.)

Prevrotroma (Man@enia) Brionaz, sp. nov. (Pl. XXV.
figs. 14, 15.)

Testa elongata, anguste turrita, pallide fulva, transversim obscurissime
fusco multi-lineata; anfractus 9, convexi, spiraliter minutissime densissime
striati, lonyitudinaliter costati, suturaimpressa sejuncti; anfractus ultimus
parviusculus, leviter convexus, costis 10 elevatis levigatis munitus, ad
basin acuminatus, vix productus; apertura oblonga, mediocriter lata,
utrinque angustior ; columella rectiuscula ; labrum arcuatum, postice paulo
sluuatum.

Long. 20, maj. diam. 6 millim.; apert. long. 75, maj. lat. 25 millim.

Hah. Hongkong.

This species somewhat resembles an elongated form of the
British P. nebula, but is considerably larger, and distinguishable
by a very long and turreted spire. The shell is apparently
smooth, and the very fine spiral striz can only be seen through a
lens. The longitudinal ribs are rounded and smooth. Only one
perfect specimen was brought by Dr. Hungerford from Hongkong.

Oxttva Bitowt, sp. nov. (Pl. XXV. fig. 3.)

Testa subcylindraceo-fusiformis, lutea, obscure griseo nebulata, antice
conspicue rufo-fusco flammata; spira acuminata; anfractus 7, planato
declives, griseo-fusco obscure fasciati, sutura canaliculata sejuncti; anfractus
ultimus supra medium obscure angulatus, infra medium flammis rufo-fuscis
obliquis peculiariter pictus ; columella rectiuscula, 7-plicata, basi contorta,
albo callosa, unisulcata; apertura modica, antice paulo latior, intus albida ;
labrum fere rectum, leviter complanatum.

Long. 32, maj. diam. 13 millim.

Hab. New Britain.

This shell resembles O. emicator (Meusch.) in form, but the
colouring is very peculiar and characteristic, with conspicuous
reddish-brown flames only on the lower part of the body-whorl.
The type specimen is in the collection of Mr. Carl Biilow in
Berlin, and is the largest I have seen; the smallest is about
20 millim., and the average length is about 26 millim.

MiTRA FORMOSENSIS, sp. nov. (Pl. XXYV. figs. 4, 5.)

Testa fusiformis, fusca, albo-fasciata ; spira elongata, acuta; anfractus 11,
leviter convexi, sutura impressa sejuncti, albo unifasciati, primi 8-9
longitudinaliter costellati, spiraliter multisulcati, deinde spiraliter striati,
haud costellati; anfractus ultimus spiram subequans, rotunde con-

LINN. JOURN.—ZOOLOGY, VOL. Xx. 30

396 MR. G. B. SOWERBY ON

vexus, fascia alba supra et infra medium ornatus, infra medium valde
contractus, in cauda brevi desinens, spiraliter liratus ; columella quinque-
plicata; apertura intus albida; labrum crassiusculuin, leeve, nigro-fusco
limbatum.

Long. 50, maj. diam. 16 millim. ; apert. long. 24, lat. 45 millim.

Hab. Island of Formosa.

A species belonging to the “ Turricula” section, somewhat
resembling JZ. caffra, but with the whorls more rounded and the
body-whorl much shorter in proportion to the spire, rounded in
the middle and contracted at the base. Three specimens from
the Island of Formosa, all similarly marked, and differing but
little in form.

Mitra RECURVA, sp. nov. (PI. XXV. fig. 7.)

Testa fusiformis, alba, fusco-maculata; spira acutissima, mucronata,
subgradata; anfractus 10, superne angulati, demde levissime convexi,
longitudinaliter costati; costis numerosis, levigatis, rotundatis, ad angulum
tuberculatis, interstitiis transversim sulcatis; anfractus ultimus spiram
leviter superans, infra medium multo et abrupte attenuatus, in cauda
recurvata desinens ; columella sinuosa, quinque-plicata, superne tuberculo
albo rotundato munita; apertura longa, antice angustata, intus fusco-
maculata ; labrum arcuatum, tenue, pellucidum.

Long. 16, maj. diam. 6 millim.

Hab. Mauritius (V. de Robillard).

A very pretty little species, with a prolonged recurved canal.
The longitudinal ribs are neatly tubercled at the suture, the
interstices transversely grooved, and the turreted spire produced
and attenuated towards the apex. The brown blotches are for
the most part arranged in two zones above and below the middle
of the body-whorl.

Mirra SMITHI, sp. nov. (Pl. XXV. fig. 6.)

Testa elongato-fusiformis, albida, fulvo fuscoque sparsim fasciata et
maculata; spira acutissima, mucronata, subgradata; anfractus 11, con-
vexiusculi, longitudinaliter costellati; costellis numerosissimis, angustis,
confertis, rotundatis, levigatis, juxta suturam tuberculatis, insterstitis haud
striatis ; anfractus ultimus spiram equans, supra et infra fascia pallide fulva
fusco maculata picta, zona media albida, infra medium attenuatus,
spiraliter sulcatus et granulatus, in cauda recurvata desinens; columella
leviter sinuosa, triplicata, plicis crassiusculis; apertura longa, angusta.

Long. 16, maj. diam. 5 millim.

Hab. Mauritius (V. de Robiilard).
Another elegant little species, in several respects similar to the
last, but much narrower and more cylindrical in form; the ribs

NEW SPECIES OF SHELLS. 397

much closer, and the interstices not grooved. The colouring and
form of the apex, as well as of the canal, are curiously similar, as
also the beading at the top of the ribs.

Ovutum (Brrostra) Haynes, sp.nov. (Pl. XXV. figs. 1, 2.)

Testa elongata, angustata, albida, polita, utrinque rostrata, postice
attenuata, antice subtruncato-acuminata, extremitatibus acutiusculis,
lateribus leviter convexis; apertura medio angustissima, postice paulo
latior, antice dilatata ; columella convexa, nitidissima, labrum incrassatum,
leviter sinuosum.

Long. 34, maj. diam. 7 millim.; apert. maj. lat. 23, min. vix 1 millim.

Hab. Exmouth Gulf, W. Australia (J. H. Haynes).

A graceful species allied to O. Philippinarum, but larger and
differing in form.

The specimens are nearly white, being found attached to a
very handsome white species of Melitodes, which is also probably
new to science, and of which Mr. Haynes collected several
specimens in the above-named locality, one of which he presented
to the Natural History Museum at South Kensington. The
colour of Ovulum is probably determined by that of the place of
attachment, as is the case with several well-known species which
occur on West-Indian Gorgoniz, so that should specimens be
found attached to Melitodes ochracea (of which Mr. Haynes found
severa] in the same locality) they would probably be of an orange
colour. Hitherto, however, they have only been found on the
white species.

Trocnus (INFUNDIBULUM) BACCATUS, sp. nov. (Pl. XXV.
figs. 8, 9.)

Testa breviter conica, late umbilicata, luteo-albida, nigro fuscoque
punctata; anfractus 6, convexiusculi, spiraliter striati et granulis gem-
muliformibus nigro-fuscis, fulvis et albidis quinqueseriatim dispositis
instructi; anfractus ultimus ad peripheriam angulatus, infra planato-
convexus, liris 6, angustis, parum elevatis, fusco articulatis ornatus ; apertura
obliqua; columella oblique rectiuscula.

Alt. 12, maj. diam. 14 millim.

Hab. Mauritius.

A pretty little species, regularly and exquisitely spotted and
beaded.

THatori1a WILKIA, sp. nov. (Pl. XXV. figs. 10, 11.)

Testa conica, elata, grisea, albo fuscoque maculata et strigata; apice
fusco; anfractus 7, planato-declives, ad suturam undulatim angulati,
spiraliter multi-lirati, liris rugosis, hic illic irregulariter nodulosis; an-
fractus ultimus inferne angulatus ad basin levissime convexus, angus-

30*

398 MR. G. B. SOWERBY ON

tissime umbilicatus, spiraliter liratus, liris 8 (lira minima interveniente)
rugosiusculis ; columella leviter sinuata, laevigata; apertura subquadrata,
intus argentea.

Alt. 19, maj. diam. 15 millim.

A single specimen, locality unknown.

AMATHINA IMBRICATA, sp. nov. (Pl. XXYV. figs. 12, 18.)

Testa elongato-subtrigona, ineequilatera, sordide alba, utrinque angulata,
postice incurvata; dorso leviter concavo, laminis rugosis transversis partim
tubulosis sculpto, margine sinistrali bicarinato, dextrali unicarinato,
lateribus corrugatis, dextra concava, sinistra planulata ; apice minutissimo ;
apertura oblonga, antice quadrata, postice rotundata.

Long 13, maj. diam, 7, maj. alt. 5 millim.

Hab. Mauritius.

This species differs from the type (Amathina tricarinata) in
several particulars. Of the three keels, two are close together
on the left dorsal margin, and the broad space between these and
the right keel is curiously laminated and corrugated. A single
specimen of this interesting shell has been sent me from Mauritius
by M. V. de Robillard.

CocHLosTYLaA (AXINA) GLOYNEI, sp. nov. (Pl. XXV. figs. 16,
17.)

Testa globoso-depressa, imperforata, solidiuscula, nigro-fusca, ad apicem
pallida, epidermide albida hydrophana, in strigis obliquis et fasciis trans-
versis dispositis pulcherrime ornata ; spira convexa, apice obtusa; an-
fractus 6, convexiusculi, lente accrescentes, oblique striati, sutura impressa
sejuncti; anfractus ultimus subinflatus ad peripheriam obscurissime
angulatus, superne et inferne convexus; columella brevissima, albida,
oblique recta; apertura parviuscula, transverse oblonga, leviter arcuata,
ubique subequaliter lata, intus purpurea; labrum arcuatum, tenuiter
reflexum, leviter complanatum, purpureum, fusco limbatum.

Maj. diam. 33, alt. 24 millim.; apert. long. 15, lat. 6 millim.

Hab. Philippine Islands.

A single specimen, in perfect condition, of this interesting

shell is in the collection of Mr. C. P. Gloyne. The whitish —

diaphanous epidermis in oblique streaks and transverse bands
relieves the plain dark-brown colour of the shell, which, though
not particularly remarkable in form, differs considerably from its
congeners.

Ortuaticus MacAnpREWI, sp. nov. (Pl. XXYV. fig. 18.)

Testa elongata, griseo-fulva, fusco zonata et strigata, hic illic nigro
radiata; apice obtusiuscula, griseo-lilacea; anfractus 83, regulariter
accrescentes, convexiusculi, longitudinaliter leviter striati, sutura impressa

mer meet se

NEW SPECIES OF SHELLS. 399

sejuncti, lineis 3 vix conspicuis fusco alboque articulatis (in anfr. post-
penult. evanidis) spiraliter notati; anfractus ultimus spiram subequans,
rotunde convexus, zona supra griseo-fulva, media et infra fusca ; columella
fere recta, tenuicula, intus griseo-lilacea, extus nigra; apertura semiovalis,
intus lilacea; peristoma simplex, haud reflexum, vix incrassatum, nigro
limbatum, margine columellari callo nigro late interne suffuso induto.

Long. 70, maj. diam. 30 millim. ; apert. iong. 28, maj. lat. 16 millim.

Hab. Santiago de Cou, Peru.

This species, of which I have only seen a single specimen, is
similar in form to O. Benson, but it has no spiral sculpture and
the colouring consists principally of light brown zones with a
few longitudinal streaks of brown and black; the articulated
painting is very slight, and is only to beseen on the upper
whorls.

PECTUNCULUS CREBRELIRATUS, sp. nov. (Pl. XXV. fig. 20.)

Testa transverse ovata, crassa, subzequilateralis, albida, fusco hic illic
sparsim maculata et fasciata, costis circ. 30, parum elevatis, et liris
numerosissimis angustissimis radiata, stris concentricis cancellata, minu-
tissime granulata; umbones prominentes; area ligamenti recta angustius-
cula, mediocriter longa; latere antico rotunde arcuato, postico angulato ;
margo dorsalis posticus declivis; cardine arcuato, dentibus numerosis
divergentibus instructo, pagina interne alba, postice fusco tincta.

Long. 35, umbonis marg. 3] millim.

Hab. Moreton Bay, Australia.

Resembling Pectunculus angulatus in form, but quite differently
sculptured. It is densely radiately ridged throughout and
sculptured with minute transverse striz, which give a rough
granular character to the surface. The epidermis remaining on
the margin of the shell is velvety.

CRASSATELLA JAPONICA, sp. nov. (Pl. XXV. fig. 19.)

Testa ovato-trigona, crassa, inzequilateralis, fulva, fusco radiata, con-
centrice irregulariter striata; umbones acuti, approximati; area dorsalis
utrinque mediocriter excavata; latere antico rotunde arcuato, postico bi-
angulato ; margo dorsalis posticus recto-declivis; pagina interna carnea,
impressiones musculares fulvo imbutz.

Long. 32, umbonis marg. 27 millim.

Hab. Japan.

A very distinct species, though presenting no very prominent
characters. The style of radiation is common to many of the
Orassatelle, but the rough irregular striation in place of the
ordinary concentric grooves is peculiar.

400 MR. A. D. MICHAEL ON

CLAVAGELLA MINIMA, sp. nov. (Pl. XXV. figs. 21, 22.)

Testa parva, alba; tuba longitudinaliter compressa, apertura leviter
reflexa, haud fimbriata ; valva dextra lata, planulata, extus concentrice
irregulariter rugata, intus argentea, margo dorsalis fere rectus, ventralis
valde arcuatus.

Valve, umbonis marg. 7, antero-post. 10 millim.

Hab. Mauritius (V. de Robillard).

This small Mauritian species is certainly distinct from the
Maltese C. aperta, but it is impossible to say whether the speci-
mens are full-grown or not. I have seen four specimens differing
but little in size, two of which are in the National Collection at
South Kensington.

DESCRIPTION OF PLATE XXV.

Fig. 1, 2. Ovulum Haynesi, p. 397. Fig. 16, 17. Cochlostyla Gloynezt,
3. Oliva Biilowt, p. 895. p. 398.
4,5. Mitra formosensis, p. 395. 18. Orthalicus MacAndrewit,
6. Smithi, p. 396. p. 398.
G recurva, p. 396. 19. Crassatella japonica,
8,9. Trochus baccatus, p. 397. p. 399.
10, 11. Thalotia Wilkie, p. 397. 20. Pectunculus crebrelira-
12, 18. Amathina imbricata, tus, p. 399.
. p. 398. 21, 22. Clavagella minima,
14, 15. Pleuwrotoma Brione, p. 395. p. 400.

On some unrecorded Parasitic Acari found in Great Britain.
By A. D. Micnast, F.LS., F.Z.8., F.R.MLS.

[Read 7th February, 1889. ]
(Piatt XXVI.)

I spent the summer of 1888 in Derbyshire, near Chatsworth :
the cold and wet season, however, rendered ordinary open-air
collecting very fruitless ; I therefore took the opportunity of ob-
taining all the wild mammals which I could obtain alive, or imme-
diately after death, with a view to see whether I could find any
unknown forms of parasitic dcar?. My search was rewarded
by finding the three species recorded in this paper, which,
I believe, were not previously known. The Myocoptes adds
another to this curious genus, which was founded by Claparéde,
and of which there was only one species known previously ; both
are parasites of the mouse and rat tribe; they are strictly ecto-

= agen

Linn.Soc. Journ. Zoon Vou. XX. Pl. 25.

Hanhart imp.

NEW SHELLS

SOME PARASITIC ACARI. 401

parasitic, living among the hairs, and they are furnished with the
most remarkable apparatus for holding these hairs, to which the
females of the present species cling so tenaciously that the grasp
is often not relaxed even in death. The species now described is
very much smaller than that previously known.

The second species is a Symbiotes, one of the Sarcoptide, and
is a parasite of the hedgehog. I regret: that I was not able to
find the male of this species ; but I only had one hedgehog, the
parasites were extremely few upon it, and these few were most
difficult to catch, running up and down the quills of the hedge-
hog and about between them with great rapidity.

The third species, which is very minute, does not appear to fit
satisfactorily into any known genus ; I have therefore been forced
to institute a genus, “ Goniomerus,” for it; the species will of
course serve as a type for the genus; it would be too soon to
attempt to define the latter accurately in any other manner,
particularly as the present species is so extremely minute as to
render detailed observations of it most difficult.

Myocortrs renax, n. sp. (Pl. XXVI. figs. 1-7.)
Male. Female.

mm, mm,

Memtmr abou Ge eek. 15 ‘20 to '27
0 i rr ‘Li ‘10
Length of 1st and 2nd legs, without the

eee OOUD Ae. ‘06 ‘06
Length of claw of 2nd leg .......... ‘03 ‘02
Length of 3rd leg, without claw...... ‘04: ‘O4
Length of claw of 3rd leg .......... ‘03 ‘02

The colour and texture in both sexes is very similar to that of
the only other known species of the genus, viz. I. musculinus,
except as mentioned below.

Male.—Diamond-shaped, the division between the cephalothorax
and abdomen well marked by a nearly straight transverse line,
the body being slightly constricted at this point. Outline of
cephalothorax slightly and irregularly undulated; that of the
abdomen on each side convex anteriorly, then concave, and again
convex posteriorly. The abdomen is not divided posteriorly into
two pointed projections as in MZ. musculinus, but comes to a single
central bluntish point. On each side of this point is a square
projection, from each of the two outer corners of which springs a
very long and powerful hair. Thus there are two pairs of these

402 MR. A. D. MICHAEL ON

hairs instead of one pair, as in WZ. musculinus. These hairs are
nearly as long as the whole length of the body; both pairs curve
upwards at first, and then run backward and outward; but the
pair near the centre are much the straighter. There are two
other much smaller hairs springing from each of the square pro-
jections. There are two large hairs on each side of the body, one
a little in front of, and one a little behind, the constriction between
cephalothorax and abdomen ; the hinder is considerably the larger.
There is a pair of large hairs on the dorsum of the cephalothorax,
and several smaller pairs on that of the abdomen. The claws of
the first two pairs of legs are extremely long, very slightly curved,
and rod-like. The legs of the fourth pair, although of the same
general form as in I. musculinus, are very much smaller in pro-
portion, they do not project behind the body, and are scarcely
thicker than those of the first and second pairs; they are much
shorter and narrower than those of the third pair, which extend
considerably further back than the body. Each tarsus of the
fourth pair bears a long flexible hair and a short stout spike still
nearer to the tip. On the inside of the flattened claw of each
third leg, at the proximal end of the claw, is a curved chitinous
process projecting inward, and a similar process is found on the
preceding joint. There are several pairs of long hairs on the
ventral surface, and a pair of stout spines behind the penis.

The Female—The female differs from J. musculinus more
than the male does. The abdomen of the present species is almost
cylindrical, although somewhat flattened dorso-ventrally ; it is
slightly smaller at the ends than in the middle, but is without
the highly flattened form and the lateral angular projections found
in the other species. The whole abdomen is covered by strong,
slightly irregular transverse wrinkles or ridges; these are slightly
rough, but are not set with the well-marked and regular points
found on those of the female of IZ. musculinus. The length of
the abdomen varies greatly in different specimens, or more pro-
bably in the same specimens at different times: there is usually
one egg matured at a time, and as it is very long, it probably
elongates the extensile abdomen as it grows, and other circum-
stances will probably produce temporary variations in the length
of the abdomen. The variation in the length of the female noticed
in the measurements arises almost entirely from the abdomen.
The creature usually holds the hair of the mouse with the third
and fourth claws, aid keeps the rostrum sharply inclined down-

SOME PARASITIC ACARI, 403

ward, while the abdomen is somewhat raised ; thus the creature
has the appearance of possessing caput, thorax, and abdomen.
The abdomen has two very long hairs at its hinder end, and there
are two smaller, but still large, hairs on the lateral edge between
the second and third legs ; and two on the cephalothorax, as in the
other species, besides two sparse longitudinal rows of smaller
hairs on both divisions of the body. The first and second legs
are like those of the male, but the claws are shorter. The third
and fourth legs resemble the third pair of the male, but the claws
again are shorter, but, on the other hand, they are broader.
These great holding-claws in this species are much wider in pro-
portion than those of I. musculinus ; indeed, their breadth is
very remarkable. On the ventral surface the epimera of the first
pair of legs join in the median line and are prolonged by a short
sternum (so-called) ; those of the second pair of legs are forked
at their distal ends. Posterior to these, and almost touching those
of the third pair, are two conspicuous chitinous pieces of rather
more than half-a-circle shape with their outer corners slightly
prolonged; the chitinous pieces protect the vulva, which lies
between the third and fourth pairs of legs.

The Nymph.—The nymph is very similar in form to the male,
but without the square posterior projections ; there is ouly one
pair of large hairs posteriorly. The fourth pair of legs are similar
to the third, as in the female. The constriction between cephalo-
thorax and abdomen is very marked. Neither pair of legs are
nearly so long as the third pair of the male.

The Larva.—Al|most diamond-shaped ; of course there are not
any fourth legs, but the third pair are placed near the posterior
end of the body, giving a very singular appearance. Otherwise
the larva resembles the nymph. Both differ considerably from
those of A. musculinus.

Habitat. Parasitic upon the field-vole (Arvicola agrestis).

SYMBIOTES TRIPILIS, n. sp. (Pl. XXVI. fig. 8.) Female.

SEMEN ers eas Py ck eee reas ae es ‘33
SE ee en ee "26
Length of legs: Ist and 2nd pairs about ........ 15
Braparabout: sf Pik. ve. ‘06
PMA AMOUL Wenner s'il. 6) ‘03
Length of longest hair on 8rd tarsus about ...... ‘70

on hind margin of body about °40

9 +P)

4.04: MR. A. D. MICHAEL ON

This species is closely allied to the Chorioptes setiferus, var.
hyene of Mégnin; there are, however, well-marked specific dif-
ferences.

I have adopted Gerlach’s name of Symbiotes for the genus in
preference to Gervais’s name of Chorioptes, which Méenin has
employed, because Gerlach’s is the earlier, and Mégnin appears
to have used Chorioptes under a misapprehension. Mégnin says
that Gerlach’s name has priority, but he says that it fails because
Redtenbacher had already called a genus of Coleoptera by that
title ; this, however, seems to be an error. Gerlach’s name was
published in his monograph in 1857, whereas Redtenbacher’s
genus was in his ‘Fauna Austrie’ in 1858; it is therefore the
latter genus which fails, not Gerlach’s.

I have, unfortunately, not been able to obtain the adult male.

Female.—Body irregularly pentagonal, broadly truncated pos-
teriorly ; considerably wider anteriorly than posteriorly ; anus
projecting from the hind margin. Body almost white, semitrans-
parent ; rostrum, epimera, and legs strongly chitinized and brown.
First and second pairs of legs thick and strong; the tarsus of
each of these legs has a strong hair or rod springing from its
upper surface in the median line; this hair is of almost equal
thickness throughout, and ends quite bluntly, it stands upward
and is considerably longer than the tarsus. These tarsi are ter-
minated, as usual, by suckers. The epimera of these two legs nearly
join at their posterior ends. The third pair of legs are much
smaller, not above half the length; they are terminated by three
very long hairs of unequal lengths, the longest is more than twice
the length of the body. The fourth legs resemble the third, but are
so small as to appear quite rudimentary ; they are not above half
the length of the third pair, they also are terminated by three
long hairs, but these are small and short compared with those on
the third pair. The body bears two pairs of very long hairs on
the hind margin, the outer hair of each pair is considerably longer
than the inner, the outer hair is longer than the body. The vulva
is a rounded median depression, towards which the striz of the -
skin run from all sides, as in S. setiferus.

The Nymph.—Resembles the adult female, but, of course, is
smaller and less chitinized.

Habitat. I found the species upon the hedgehog (4rimaceus
curopeus). It is very active, running up and down the spines of
the hedgehog with great rapidity.

SOME PARASITIC ACARI. 405

GoNIOMFRUS* MUSCULINUS, nov. gen. et sp. (Pl. XXVI. fig. 9.)

2 mm.
, hs ee ce te eee cies "155
Ee "105

eeetn Oo lees (all pairs) about .............. 035

Colouwr.—Semitransparent white.

Texture—Much that of the ordinary Sarcoptide, 7. e. smooth

= and leathery, not polished. The skin is marked more or less
with excessively fine wrinkles or striations, so fine that they are
very difficult to see.

Form &§c.—Body almost oblong, but somewhat narrower poste-
riorly than anteriorly ; there is not any clear demarcation between
cephalothorax and abdomen. The rostrum projects, but is bluntish;
a pair of mandibles may sometimes be seen projecting from the
mouth-opening. The rostrum bears a pair of fine hairs. The
body is much compressed dorso-ventrally, and is divided into
four lobes on each side, the hind lobe showing a slight tendency
to be bifid. On the posterior margin there is a slight indenta-
tion above the anus, which is almost terminal, but slightly on the
ventral surface. From the hind margin spring a pair of long

7 hairs directed backward, they are fully two thirds of the length
of the body; immediately below them are another pair of similar
hairs proceeding from lobes on the ventral surface ; as this under
pair is entirely concealed by the upper pair, when viewed from

~ above, it is not shown in the figure. The legs are short, and are
the great peculiarity of the creature; they are all alike and all
directed forward, the fourth pair usually more so than is shown
in the figure, where they are spread out. They consist of five
joints ; the femur is very large and singular in shape, it projects
beyond the body and bends suddenly at rather less than a right
angle, so as to form an advancing corner which completely alters
the direction of the leg. The more distal joints become gradually
smaller ; the tarsus is terminated by a small claw, smaller in pro-
portion than can be shown in a drawing the size of fig. 9, and
there is another small chitinous projection close to it which may
be regarded as a mere peg or a second claw; 1 am inclined to
look on it in the former light. There are hairs on the underside
of each joint of the leg and one or more on the upperside of

* ywvia, an angle; znpds, the thigh.

4.06 PROF. B. T. LOWNE ON THE

each tarsus, besides a few others. The penis is placed on the
ventral surface in the median line between the hinder part of the
coxe of the third legs ; the vulva between those of the second
legs.

I found the creature on the surface of, or very slightly buried
in, a depression of the skin lining the inner side of the external ear
of the short-tailed field-vole (Arvicola agrestis). I believe it to be
unrecorded, and propose to call it “ Goniomerus musculinus.”

DESCRIPTION OF PLATE XXVI.

Hig. 1. Myocoptes tenax, 9. Dorsal aspect. Xx 175. Drawn from a
specimen with long abdomen.

2. 5 or lies Ventral aspect. x 175. Drawn from a
specimen with short abdomen.
3. ” La From the side. x 175. Natural position,

holding the hairs of the mouse.

* Pies oes Dorsal aspect. x 175.

- Ventral aspect. »X 175.

55 » nymph. Dorsal aspect.

a ara: Og 3rd leg, seen from the inner side. x 350,

8. Symbiotes tripilis, 9. Ventral aspect. x 180.

9. Goniomerus musculinus, $. Dorsal aspect. % 175, (There is
another pair of long hairs on the hind margin, below and
hidden by the pair shown.)

On the Structure of the Retina of the Blowfly (Calliphora ery-
throcephala). By Brensamin THompson Lowng, F.R.CS.,
F.L.S., Hunterian Professor of Comparative Anatomy in
the Royal College of Surgeons.

[Read 21st February, 1889. ]
(Prats XXVII.)

In 1884 I had the honour of reading a paper before this Society
on the compound vision and morphology of the eye in insects,
which was published in the second volume of the new series of
‘ Transactions.’

That paper received at the time much adverse criticism, and
Dr. Hickson published a memoir in the ‘ Quarterly Journal of
Microscopical Science,’ in which he convinced himself that he had
completely refuted my observations.

Id See to

;
af

x) &(\\2

ay:

*
ey a] : :

eT as pF eee ee ae eee ey ae

NEW peer ACARI oa rsa. a Si

== + /

> coaie Te a ae

is ie

RETINA OF THE BLOWFLY. 407

From that day to this I have continued to work at the subject,
and I now venture to bring before this Society evidence which I
think can hardly fail to convince even the most sceptical of my
opponents. Although I never had any doubt of the correctness
of my figures or descriptions, I felt it incumbent upon me to
produce preparations which would admit of no double interpre-
tation, but which would appeal at once to the eyes of those who
are only partially acquainted with the histology of the vertebrate
retina.

At the time I published my former paper I felt so certain that
the views I held would receive a ready acceptance, that I did not,
perhaps, enter sufficiently into minute details, and left many points
to be investigated by other workers. I have since examined
every structure in the greatest detail, and have much to add with
regard to the developmental history of the compound eye.

The retinal rods, which I figured correctly in my former paper,
correspond with the periopticon of Dr. Hickson, except that his
figures show that every vestige of nerve-structure and nerve-
terminal organs had been completely destroyed in his prepara-
tions, leaving nothing but the skeletal framework with the
tracheal vessels, which he has delineated most carefully and
correctly.

He states that my paper and investigations were unnecessary,
owing to the unanimity of previous investigators: none, however,
agree in any detail with Dr. Hickson, nor, so far as I am able
to judge, to any considerable extent with each other.

Putting aside for the moment the earlier observers, the so-
called periopticon of Hickson has only been described in detail
by Berger, Carriére, Ciaccio, Viallanes, Hickson, and myself.

To show how far these observers agree with each other and
with the older writers, I will quote a few sentences from
Dr. Hickson’s paper. He says :—

“ Previous to the publication of Berger’s paper the optic tract
of insects had been briefly described and names given to the
various regions. Thus Weismann called the opticon and epiopti-
con the ‘ bulbus,’ the region where the optic fibrils decussate the
‘ Stiel,’ and the periopticon the ‘Augenscheibe’”’ (J. c. page 27).

Even the most cursory acquaintance with the work of the
German naturalist would have shown Dr. Hickson that this is
an egregious misstatement of Dr. Weismann’s nomenclature.

408 PROF. B. T. LOWNE ON THE

Weismann’s ‘ Stiel’ was the optic nerve, and his ‘Augenscheibe’
the structure from which the dioptron is developed. I shall have
later to give Dr. Weismann’s views more fully. Dr. Hickson
continues (page 27) :—“ Since Berger’s paper appeared Carriére
has described the periopticon as ‘a layer of long palisade-shaped
cells, the number of which corresponds with the eye units; every
one of these palisade cells possesses an oblong nucleus at its
foremost, somewhat broader, end.’ My researches show that
this description is quite inaccurate. The elements of the periop-
ticon are not cells, and the large oval nucleus situated in each
element does not exist; nerve-cells, when they exist in the region
of the periopticon in Musca, lie between the elements and not in
them, as my figures show.”

These statements and others show that Dr. Hickson and Carriére
do not agree. With regard to the nuclei described by Carricre,
they undoubtedly exist, but not, as Carricre thought, within the
palisades, but externally to them, immediately beneath their in-
vesting sheath. Dr. Hickson is right when he says these bodies
are not cells, they are developed from cells, and each consists of a
bundle of fusiform rods. With regard to the terminations of the
optic nerve, Carricre distinctly traced the nerve-fibres into the pali-
sades; Dr. Hickson says they goround them. I trace them directly
into the fusiform rods which form the palisades. The structures
seen and correctly figured by Dr. Hickson are tracheal vessels.

Carri¢re supposed the nerve-fibres to pass out at the superficial
end of the palisades and to perforate the basilar membrane; from
this I entirely dissent. In support of this view Carriére has
figured, quite diagrammatically, what I believe is a tracheal vessel
seen behind the fusiform body. Carriére also saw the highly re-
fractive outer ends of the rods, or, rather, that part which is con-
nected with their inner portion, and says, “in Musca vomitoria
one sees in every cell a cylindrical axis.”

Dr. Hickson entirely put himself in the wrong in describing
the nervous elements as between the palisades; his nervous
elements are undoubtedly fine tracheal tubes. Dr. Hickson’s
figures accurately represent the nerve-sheaths and trachew as
well as the supporting neuroglia, but no vestige of nerve or
nerve-end organs appears in them. A careful examination of his
own figures at once leads to a dissent from all his statements,
which are as inaccurate as his figures are accurate. J cannot
understand how so good an observer could have been so misled.

RETINA OF THE BLOWELY. 4,09

Berger and Viallanes trace the optic-nerve fibres through a
series of small round cells, very conspicuous in the outer half of
my retina, Hickson’s periopticon. Hickson regards these cells
as of quite secondary import. They clearly belong to the sup-
porting tissue and are external to the sheaths of the retinal
elements, which are continuous with the perineurium of the optic
nerve.

Dr. Hickson and Dr. Grenacher suppose the sheathing cells of
the great rods, retinule of Grenacher, to be the nerve-terminals ;
and more recently Platten pretends that the optic nerve ter-
minates in the crystalline cone. There is therefore no unanimity
amongst previous writers, especially in matters of detail; as it is
impossible that they can all be right, it is quite possible, as 1
assert, that they are all wrong.

Dr. Hickson’s neurospongium, or terminal anastomosis, which
is inadmissible on physiological grounds, is no nerve-plexus
at all, but the tracheal plexus and the sustentacular framework
of my retina.

It is exceedingly difficult to prepare sections which show the
true retinal end-organs. This difficulty arises from the fact that
the chloroform and alcohol used in the process of imbedding
dissolve the fatty matters from the nerves, and the external
extremities of my retinal rods are completely dissolved or dis-
integrated by the action of aqueous media.

I have, however, on many occasions succeeded in obtaining
sections in which both the inner and outer extremities of the
retinal rods, as well as the nerves, remain more or less unaltered.
Another difficulty arises from the extreme transparency of these
structures in very thin sections, and from the fact that they
cannot be stained by any of the stains used in such researches ;
the outer ends of the rods are not affected by ainone solutions e
aniline dyes, except vesuvin *.

In thicker sections the numerous round cells between the
retinal nerve-end organs, which are not connected with nerves,
but with the sustentacular framework, entirely conceal the outer
ends of the rods.

There are two methods which give good results; in both the
tissues must be fixed either with osmic acid and absolute alcohol

* The best demonstration of these organs is obtained by staining with a
solution of vesuvin in aniline water. The solution must be quite freshly made,
and unfortunately such preparations fade rapidly when mounted in balsam.

410 - PROF. B. T. LOWNE ON THE

or in absolute alcohol, and imbedded in paraffin without the use
of ether or turpentine. Very thin sections are then cut and
fixed on the slide with shellac and kreosote. The cement must
be thoroughly dried in the oven at the melting-point of the
paraffin used, and on no account at a higher temperature.

The paraffin is next removed by turpentine. The slide is then
wiped on its back and edges, and flooded with pure spirit, which
is drained off, and immediately afterwards flooded with 75 per
cent. alcohol and rapidly drained ; Erhlich’s logwood solution is
then poured on the slide and washed off after a few minutes or
longer by agitating the slide for a few moments in water, and it
is again flooded with 75 per cent. alcohol. The washing is the
most dangerous process, as if the specimens are kept too long in
water the outer ends of the retinal rods will be entirely dissolved.
Instead of Erhlich’s logwood a solution of vesuvin in water may
be used ; it stains the retinal-end organs better than any of the
aniline dyes. Saffranine in 50 per cent. alcohol, or a solution of
fuchsine or eosine, may be used for staining, and the washings done
with spirit, the results of which are often satisfactory. Spiller’s
purple gives excellent results, but the specimens are not permanent.
The specimen, after flooding with 75 per cent. alcohol, is treated
with pure alcohol, rapidly drained and cleared with clove-oil
and mounted in balsam.

Or, after the first washing in water, the specimen may be
mounted in glycerine, gradually adding stronger and stronger
glycerine and water, and draining after each addition. I have
found that with aniline dyes a very dilute solution of sodium
carbonate, °5 per cent., or aniline water is not inadmissible for
washing out the excess of the stain.

Glycerine mounts, when successful, show the outer ends of the
rods, either vacuolated or frequently partially dissolved, more
plainly than balsam mounts.

The balsam mounts need very careful illumination, otherwise
it is impossible to see the outer ends of the rods.

If we trace the optic nerve, we observe that its fibres run in
larger or smaller bundles, invested in a very transparent sheath,
or perineurium. They terminate in the palisade layer by entering
the fusiform elements. The sheath is continued over the fusiform
elements, and terminates on the inner surface of the basilar mem-
brane. The tracheal vessels accompany the bundles of optic
nerve-fibres, outside their sheath, and continue between the pali-

RETINA OF THE BLOWFLY. 411

sades, and ultimately pierce the basilar membrane and run
between the great rods.

The figure given (Plate XXVII. fig. 1) is from the eye of
a Hawk-moth, in which these details are larger and more
easily seen than in the Blowfly. The palisade bodies do not
reach the basilar membrane, but are prolonged as extremely
transparent rods, 3 to 5 w in diameter, in the fly and in most of
the insects I have examined, and from 20 to 304 in length
(Plate XX VII. figs. 2and 3,a). These with the palisade cells, d,
form my bacilli or retinal end-organs, the whole length of which
is from 60 to 70. The outer transparent portion is rarely
straight, but usually strongly curved in a crook. They exhibit
a fine longitudinal striation.

The outer ends of the rods evidently consist of some substance
resembling mucin; they have the same refractive index and
general characters as the mucin of the intestinal epithelial cells
of the insect.

The inner extremity of the outer part of the rod is imbedded
in the fasciculus of elongate cell-like palisade bodies, fig. 2,
which form the inner portion of the retinal end-organs ; each outer
segment appears to be made up of a number of finer rods, 2 » in
diameter, pressed together into a cylinder; these produce the
longitudinal striz. Hach small component rod lies on the inner
surface of one of the fusiform cell-like bodies which form to-
gether the inner part of the retinal end-organ.

The outer ends of the rods are surrounded and, except in
very thin sections, concealed by the small round chaplet-cells of
Viallanes (fig. 2,¢). These are connected with each other by fine
processes and form a true adenoid sustentacular tissue, well seen
in transverse sections of the pupa (fig. 4).

Comparison of the Bacillary Layer with the Bacillary Layer
of the Vertebrate Retina.

In size and structure the elements of the retina are almost
identical with those of the vertebrate ; the optic nerve terminates
in the protoplasmic inner segment, whilst the outer segment is
transparent, resists stains, exhibits longitudinal striz, and swells
up with water in both. In both it is easily destroyed, and fre-
quently exhibits vacuolation.

LINN. JOURN.—ZOOLOGY, VOL. Xx. 31

412 PROF. B. T. LOWNE ON THE

In most insects the outer, highly refractive ends of the retinal
end-organs are imbedded in abundant pigment. The flies are
the only exception, and in these the cells surrounding the bacilli
are free from pigment.

The Trachee (Plate XX VII. fig. 3) form adense network around
the inner segments of the retinal end-organs in insects, and
branches extend to and perforate the basilar membrane. These
fine tracheze are without any spiral markings, and are easily
mistaken for fine nerve-twigs. The figure given (fig. 3) shows
these tracheee in a moth, and it can be readily seen that they
lie between the nerve-end organs, and that they branch
dichotomously between the great rods. The aniline stains at
once colour the tracheex, whilst they have no effect upon the
nerves. These stains, however, attack the nerve-sheaths, but
not the outer ends of the retinal end-organs. By the use of
aniline stains, especially Spiller’s purple, I have been able to
trace the finer tracheal vessels, which have been constantly mis-
taken for nerves, to the larger tracheal trunks and in one of my
photographs this relation is sufficiently evident.

The illustrations on Plate XX VII. show the large size of
the bundles of optic nerve-fibres with their terminations in the
retinal end-organs ; they also show that nothing bearing any
proportion to the magnitude of these nerve-cords passes through
or even up to the basilar membrane. The basilar membrane is
chitinous and has a cellular layer on both its inner and outer
surface ; that on its inner surface consists of branching or stellate
cells, which are continuous with the sustentacular framework of
my retina; the outer layer consists of pigment-cells, continuous
with the pigment-sheaths of the great rods. The perforations in
the chitinous layer of the basilar membrane are between and not
opposite to the extremities of the great rods, and transmit the
tracheal vessels.

The structure of the great rods has with some been the diffi-
culty in accepting my views. The appearance of these structures
in many sections is certainly perplexing. The reason is that ~
which I have already insisted upon. In life they are hollow tubes
filled and distended with fluid. In bad preparations they appear
stellate in transverse sections and present no central cavity ; in
radial sections they are separated from each other by wide spaces,
often filled by distended tracheal vessels.

ere

RETINA OF THE BLOWFLY. 413

In transverse sections, when unaltered by the process of im-
bedding, they are circular or hexagonal rings, with a large
central cavity; they touch each other at their periphery, and the
tracheal vessels appear as thick-walled but very small tubes.
Each great rod is seen in such sections to be lined by a thin
cuticular layer, which dips down between the sheathing cells ;
it is the folds of this membrane which appear as bright highly
refracting points under unfavorable conditions of illumination.
With direct central light, thin sections, with oil or water
immersion-lenses, no longer present these appearances; there
is no bundle of axial rods in such preparations when properly
examined, only a thin cuticular lining.

Further evidence in favour of my views is, I believe, shortly
forthcoming from the pen of an independent observer. Prof.
Plateau informs me that last year, at Cologne, Dr. Exner showed
the single image formed by the compound eye—the image in
the plane of my basilar membrane formed by the uninjured eye,
z.e. by my dioptron. I wait anxiously for the spring, as with
fresh insects at command I have little doubt the demonstration
of an erect picture in this region is perfectly easy.

The Development of the Compound Eye.

The development of the compound eye was described by Weis-
mann in 1864*. I have gone througha most laborious research,
and in the main points my observations agree with those of the
great German investigator. Weismann says it has long been
known that the eye in insects is developed from two perfectly
distinct parts—one from the nerve-centres of the larva, the other
from the optic dise (‘‘ Augenscheibe,” 7. c. p. 194).

If we follow the development of the optic disc, we find it at first
as a thin cellular expansion enveloping the anterior part of the
hemisphere (or supra-cesophageal ganglion). It consists of cells
(the optogenic cells of Viallanes) which are larger than those
of the other discs; they measure 15 4 in diameter at an early
period of the pupa state and have large clear nuclei. During
the formation of the head, the eye-disc separates considerably

* “Die Entwicklung der Dipteren,” Leipzig, 1864. Reprinted from Koll.
Zeitsch. f. w. Zool.
ai?

414 PROF. B. T. LOWNE ON THE

from the hemisphere, the interspace being filled with the granular
yelk-like substance of the somatic cavity of the pupa. The
whole dioptron is developed by a division of the optogenic cells,
as Claparéde long ago showed. Each original cell corresponds
to a single corneal facet. These cells form almost hemispherical
projections on the outer surface of the dise and are soon covered
by an extremely thin cuticular layer.

The cuticular layer is seen in my sections to dip slightly
between the cells, whilst the corneal lens is secreted subsequently
between the cell and the primitive cuticular layer. The lenses
are, as I have already described them, perfectly distinct from the
chitinous layer, giving rise to the condition I have designated
the kistoid cornea. In adult pupz the distinction is perfectly
apparent, although Dr. Hickson has denied that my description
is correct; the most patient reinvestigation entirely confirms
my former statement.

So far my investigations entirely accord with Weismann’s de-
scription. Weismann, however, believes that the great rods
contain a nervous structure, which he describes, from optical
sections, as resembling a bundle of fine, highly refractive, con-
ducting threads ending at the crystalline cone. He has nothing
to say of their manner of development, and only expresses the
opinion that they appear more like definite threads than the angles
of a solid rod.

These so-called axial threads, as I have stated above, are well
seen in numerous transverse sections to be mere folds of a
chitinous membrane enclosing a considerable empty cavity.

Weismann’s description of the development of the nervous
structures is as follows:—‘‘The thin nerve-cord (Stel) which
unites the optic disc to the hemisphere still appears on the fifth
day as a nervous cord; but on the twelfth day the pedicle car
no longer be seen.” He concludes, however, that it has spread
out into an invisible layer over the whole surface of the gan-
glion. That he should have arrived at such a conclusion without
sufficient evidence is quite unlike him. If, as he says and as is
certainly the case, the nerve disappears entirely between the
fifth and twelfth day, the opinion that the radial strie (which, he
says, appear later between the disc and the hemisphere) are the
same nerve spread out, is not founded on fact.

We must remember that Weismann regarded the discs as

RETINA OF THE BLOWFLY. 415

expansions consisting of epiblast-cells. It was Ganin who, ten
years later, first made sections and discovered their real structure.
He found three distinct layers—Weismann’s epiblastic layer ;
his own provisional layer, which covers it externally as a fine cellu-
lar expansion, which resembles the amnion of a mammalian
embryo in being continuous with the periphery of the disc, in
covering its whole outer surface, and in enclosing a cavity
between it and the epiblast of the disc; and the mesoblastic
layer, which fills the hollow cup-like cavity on the inner surface
of the epiblastic layer, and which consists of a network of fine
branching cells.

Weismann’s own figure (52, plate xiii., 7. c.) shows clearly that
his supposed optic nerve is the mesoblast of the disc. My own
observations show that the nervous pedicle of the optic disc
becomes atrophied and disappears, whilst the nervous retina is
developed as a papilla in front of the original optic pedicle.

In my former paper I described and gave figures of the manner
in which a new retina is developed during the skin-shedding of
the Cockroach ; the original nervous pedicle of the disc corre-
sponds to the nerve of the first few facets of the eye. As the
number of facets is far greater after each ecdysis, so a new retina
is developed from the nerve-centres asa distinct papilla; the first
formed nerve and retina at the same time undergo atrophy.

I regard the original pedicle of the disc in the Blowfly (figs. 5,
6, &7,st.) as arudiment. It exhibits few, if any, nerve-fibres and
consists chiefly of connective neuroglia continuous with the in-
vesting layer of the rudimentary hemisphere. The spongy meso-
blastic tissue which Weismann mistook for an expansion of the
nervous pedicle of the dise consists of the elements from which
the tracheal vessels and pigmented fringes of the dioptron and
neuron originate. This tissue extends into the dioptron, but only
between the ingrowing optogenic cells, which become first colum-
nar and then elongated rods, dividing during the process to form
the cone and the investing cells of the great rods, and separating
from each other to enclose the central cavity of the cone and
the great rod. Claparéde long ago correctly described the
manner of the development of the cones and great rods.

Viallanes, like Weismann, but with less excuse, mistook the
mesoblast of the dise for the optic nerve and believed that its
fibres perforate the axes of the great rods. It is easy in thick

416 PROF. B. T. LOWNE ON THE

sections to mistake fibres running between for fibres entering the
optogenie cells.

The nerve-papilla, from which the optic ganglia, the optic nerve,
and the retina are developed, gradually grows outwards towards
the dioptron (Plate XX VII. figs. 5-8, 7). It is at first covered
by a layer of columnar cells, which represent the epiblast of the
nerve-centre ; from this layer the bacillary layer of the retina is
developed. These cells become converted into the retinal end-
organs. The mesoblastic spongy tissue is gradually absorbed or
converted into tracheal and connective elements, which ultimately
form a thin layer between the retina and the basilar membrane
of the dioptron.

The retina, even when the insect is nearly ready to escape from
the pupa, is still separated from the dioptron by a space filled
with branching cells(Plate XX VII. fig. 8, mc) and secondary yelk,
so that the supposed entrance of nerve-fibres into the dioptron
cannot be explained by any known process of development.

The continuity of the trachee of the dioptron and those of the
mesoblast is the result of the penetration of the latter between
the great rods during their inward growth ; but during this period
the nervous papilla is separated by a wide space filled with
secondary yelk and reticular mesoblast from the ingrowing epi-
thelial structures of the dioptron.

Thus, if my observations are correct, the retina, like that of a
vertebrate, is entirely formed as an outgrowth from the central
nervous system, while the dioptron, like the erystalline lens
and the refractive structures generally, is formed from the ex-
ternal epiblast, which is more or less invaded by mesoblastic
elements. With regard to the retina itself, it is undoubtedly,
like the nerve-centres, no less epiblastic in the insect than in the
vertebrate, as the hemispheres themselves, as well as the ventral
ganglia, are formed from the embryonic epiblast.

In conclusion, I would add that it is scarcely fair to expect me
to prove a negative, z.e. that no nerve-fibres pass to the dioptron.
The onus rather lies with my opponents to prove that the great
optic nerve does enter the dioptron, and to find its terminals.
Even the most cursory glance at the works of Dr. Hickson, M.
Berger, M. Viallanes, and others will show that they have given
totally dissimilar representations ; of these Dr. Hickson’s are
correct enough as representations of tracheal and mesoblastic
skeletal tissues. I would ask, Which of the various structures

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RETINA OF THE BLOWFLY. 417

represented are to be considered as nervesP No one has yet
figured one satisfactory representation of the optic-nerve fibres
entering the great rods. Dr. Hickson says, ‘‘ Morphology teaches
us that the great rods are nerve-terminals.” To appeal to mor-
phology to settle the question appears to me to show on how
slender a basis of observation the received view rests, and I should
myself regard an appeal to morphology as one which is fatal to
the received view ; for, if morphology teaches us anything on this
subject, it is that the retinal end-organs belong to that part of
the epiblast from which the great nerve-centres are developed,
and that the dioptric structures arise from the superficial or
cutaneous epiblast.

DESCRIPTION OF PLATE XXVII.

Fig. 1. A section of the retina of a Hawk-moth ; partly drawn from a photograph
and finished from the section. g¢, great rods; m, basilar membrane ;
b, bacillary layer ; 7, optic nerve; ¢r, tracheal vessel.
2. A section of the retina of a Blowfly. c, chaplet-cells of Viallanes.
zs-inch objective, water-immersion.

3. A portion of the retina of a Hawk-moth; drawn from a photograph,
with details added from the specimen. The tracheal vessels seen
passing through the basilar membrane are much wore distinct in the
photograph than in the specimen seen by the microscope; these are
represented in the drawing as they appear in the photograph.

4, A transverse section through the bacillary layer of the retina of a Blow-
fly which had just emerged from the pupa.

5. A section of the optic disc and cephalic ganglion of a 3-day-old pupa.
0, optic disc; st, stalk; rt, retina. 1-inch objective.

6. A portion of the same, showing the retina and inner extremity of the
stalk.

7. A portion of the optic disc and stalk of the same. 0c, optogenic cells ;
mc, mesoblastic cells. 4-inch objective.

8. A section of the retina of a ten-day-old pupa. Showing the mesoblast
elements between the retina and the basilar membrane, 44-inch
objective.

(The letters indicate the same parts in all the figures. )

418 PROF. B. T. LOWNE ON THE STRUCTURE AND

On the Structure and Development of the Ovaries and their
Appendages in the Blowfly (Calliphora erythrocephala). By
B. Toompson Lownge, F.R.C.S., F.L.S., Hunterian Professor
of Comparative Anatomy in the Royal College of Surgeons.

[Read 6th December, 1888. ]

(Pirate XXVIII.)
. Introductory.
. Development of the Ovaries and Ova.
. The Oviducts and their appendages.
. The structure of the Gum-glands.
. Theoretical Considerations and Conclusions.
. Bibliography.

Oor © DOH

1. Introductory.
THE ovary of an insect is known to consist of a number of
ovarian tubes connected with a single outlet, the oviduct.

The ova lie within a thin membranous tube, the tunica pro-
pria, and form an egg-string; this is enclosed within a follicle,
the egg-follicle, the so-called peritoneal coat. In each ege-string
three parts are usually recognized—the terminal thread, the ter-
minal chamber, and the egg-chamber or chambers.

In the Blowfly each ovary consists of about a hundred ovarian
follicles, springing from the dilated anterior end, or calyx, of the
ovarian duct. When the ovary is mature (Pl. XXVIII. fig. 8)
the terminal threads exist as mere rudiments (¢.f.), and, properly
speaking, there are no terminal chambers. In the young ovary,
however (fig. 6), the terminal chambers are well marked (Z.c.).

Much discussion is found in the writings of various authors as
to the nature and import of the terminal thread.

Brandt (6*) maintains that there is no fundamental difference
between ovaries with and ovaries without terminal threads (J. e.
p. 21) —“ these are mere prolongations of the ovarian tubes or of
their peritoneal investment. In the former case they have the
same morphological significance as the rest of the ovarian tube,
in the latter they are mere connective or suspensory bands.”
My own observations have led me to exactly the same conclusion.

The Terminal Chamber (fig. 6, t.c.) is filled with small cells.
Stein (24) was apparently the first to distinguish the terminal
chamber as an important constituent of the ovary; he calls it the

* The numbers in parentheses following authors’ names refer to corresponding
aumbers in the Bibliography given at the end of this paper,

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 419

germinal chamber (Keimfach). He was followed by Prof. Huxley
(10, 11), Sir John Lubbock (19), Claus (8), and others. Stein
enunciated the view that the function of the terminal chamber is
the formation of germ-yelks; but he does not regard all the cells
in the chamber as germ-yelks. Sir John Lubbock went a step
further and wrote as follows :—“ In their earliest stage, the egg-
cell and the vitelligenous cells cannot be distinguished from each
other, and no one, I think, who has carefully examined the upper
part of the ege-tube inany Hemipterous or Dipterous insect can
fail to be of the same opinion.”

I agree entirely with Sir John Lubbock in this, that all the
cells in the terminal chamber are alike; but when he concludes,
“The egg-tube contains, indeed, at this end, cells which are
neither vitelligenous nor egg-cells, but which are capable of
becoming, under certain circumstances, either one or the other,”
I cannot agree with him, and my reasons will appear in the sequel.

The Egg-chambers.—This term was first applied, I believe, by
Brandt (6) to that portion of the egg-tube which contains definite
ova. In some insects each egg is formed from a single cell; this is
so in the Orthoptera; such ovaare designated by Brandt panoistie.
In other insects several cells are concerned in the formation of
the ovum ; these ova he termed meroistic.

In the meroistic egg Brandt calls the lowest cell the egg-cell,
the others he terms nutrient or yelk-cells.

The part played by these nutrient cells is a matter upon which
there is great divergence of opinion. Brandt’s view, which bas
been generally adopted in text-books and widely accepted, is this:
The ege-cell in the meroistic egg is the only cell enclosed by the
chorion, and the nutrient cells remain outside and ultimately
disappear. These are supposed to be in some way concerned in
the nutrition of the egg-cell. The great increase in the size of
the ege-cell is due to the deposition of yelk-granules within it,
around its nucleus, which Brandt regards as the germinal vesicle.
Similar changes also occur in the panoistic egg, which only differs
from the meroistic in the absence of the nutrient cells.

Weismann (26) maintains, on the other hand, that all the cells,
the nutrient as well as the egg-cell, are enclosed in the chorion,
and that they all take part in the formation of the yelk, ultimately
fusing into a single mass; and this, as I shall show hereafter, agrees
with my own observations.

With regard to the import of Brandt’s egg-cell there is less

420 PROF. B. T. LOWNE ON THE STRUCTURE AND

divergence of opinion; the identification of its nucleus with the
germinal vesicle has been regarded as of the highest importance,
and numerous attempts have been made to show that the nucleus
of the egg-cell differs from those of the nutrient cells, from the
very earliest stages of the egg-formation. Thus Dr. Claus (8)
(p. 44) writes :—‘ The questions, the answers to which are of the
highest importance, are: from whence is the germinal vesicle
derived? and what is its relation to the great yelk-cells? The
identification of the germinal vesicle appears to be difficult, and
the earlier observers came to no certain result.” After quoting
from Sir John Lubbock’s paper (19) he continues, “ 1 believe my
own observations enable me to prove that the epithelial cells, the
yelk-forming cells, and the egg-cell are modifications of originally
identical elements.’ Yet Claus thinks he distinguishes the
germinal vesicle in very immature ova by “its smailer size and
clearer contents”? from the nuclei of the adjacent yelk-cells.
Meyer’s (20) statements are in direct opposition to this ; according
to these there are several germinal vesicles, each nucleus becomes
invested by a membrane, each making, as it were, an attempt at
forming an egg, the lowest nucleus persisting and the others
disappearing.

In my opinion by far the best and most accurate description of
the development of the ovarian eggs in the Fly is due to Weis-
mann (26); it is as follows:—“ The ovaries are developed very
slowly in the Muscide; when the fly emerges from the pupa
none of the eggs are formed, although the ovary may be recog-
nized in the embryo” (J. c. p. 206). Weismann then refers to
a description of the ovary in the adult larva of the closely-allied
Sarcophaga carnaria (at page 134), “ Concerning the female sexual
organs in the larva of Musca vomitoria I have no observations ;
I must therefore fall back on those on the closely-allied S. car-
naria. Inalarva one centimetre long, the ovary has a diameter of
0:29", is flask-shaped, and differs in its histological structure from
the male sexual gland; here we find no mother-cells, the struc-
tureless sheath encloses only small round cells °018™ in diameter
with nuclei :01™, exceedingly clear and exhibiting nucleoli. These
cells are isolated with difficulty, as they adhere closely to each
other. If we tear the ovarium of an adult larva no further
structure is visible; but if a gentle pressure be applied to the
uninjured ovary one may distinguish the first rudiments of the
ovarian tubes. In the upper half of the ovary they appear as

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 421

cylindrical follicles lying parallel to each other. They are all
united above without any point (terminal thread ?) being visible,
below they are lost in the cellular mass. The diameter of a
follicle averages ‘(04™. The ovarian follicles consist of a sheath
of fine structureless membrane and its contents, which differs in
no way from the surrounding cell-mass. The sheaths are a cuti-
cular excretion from the outer surface of the cells forming the
cylinders.” (He continues on page 206) ‘So that, as I have
shown above, the original soft mass of cells with which the
ovarian capsule is filled becomes differentiated, in part, into solid
strings, which shed a cuticle from their surface, and the ovary
comes to consist of a small-celled ground-substance which fills
the capsule, in which solid cellular strings are imbedded, each
enclosed in a fine structureless membrane; of an outer and inner
epithelium, a tender albuminous contents in which free nuclei are
imbedded, as Meyer describes in the youngest condition of the
ovary, there is as yet no trace. The term egg-tubes is hardly
admissible at this stage, it is only later by the differentiation into
a wall and contents that they become tubes.”

‘On the seventh day of the pupa stage the egg-tubes still only
occupy a small zone of the flask-shaped ovarium (Taf. xiv. fig. 70);
they lie close together parallel to the long axis of the ovary and
still exhibit their original simple structure, only the contained
cells are somewhat larger and therefore more distinctly seen.
These cells are spherical, and their nuclei are easily distinguished.
The cuticular sheaths end above in rounded domes.”

“ By the fourteenth day the investing sheaths of the egg-tubes
are considerably more developed, and their outer form is altered;
the blind end is now drawn out into a point, the middle part is
swollen and the posterior part contracted. Still the lumen is
filled with cells disposed without definite order ; no regular epi-
thelium is yet visible, but there is a great difference in the size
of the cells, the central ones being larger than those of the
periphery. A little later these changes are more conspicuous,
and the egg-tube exhibits a stem, a chamber, and a nipple-like
appendage ” [Stein’s end-chamber], “ the narrowed blind end of
the tube. In the chamber there is a distinct separation of the cells,
small cells line the follicle in a single layer, as an epithelium
enclosing the larger cells ; from the latter the egg is ultimately
formed.”

“The development of the ovary shows that the life of the fly

4.22 PROF. B. T. LOWNE ON THE STRUCTURE AND

must last several weeks. A ripe egg is first found in the lowest
part of the ovarian follicle after the insect has flown about for a
long time; then a second, third, or even a fourth chamber has been
developed in which there are eggs in different stages of forma-
tion.”

“The development of these eggs takes place as follows. The
large cells which lie within the epithelium of the egg-chamber
enlarge, by their rapid growth they lose their original spherical
form and appear flattened against each other as more or less
hexagonal sections of a sphere.”

‘“‘ These cells each enclose a very distinct transparent vesicular
nucleus, and consist of homogeneous, but highly refractive cell-
substanee. With increase of the cells by growth this cell-sub-
stance becomes finely granular and afterwards dark and yelk-like.
The cell-membranes then disappear, and the yelk formed in the
cells fuses intoa mass; so algo all the nuclei disappear except one,
which becomes the germinal vesicle. It appears that the nucleus
of the cell which hes lowest in the chamber always furnishes the
germinal vesicle. This seems to have orginated Meyer’s state-
ments.”

Weismann concludes with the words*, “So far asthe Diptera
are concerned, my view accords with Lubbock’s ; we agree that the
egg of the Diptera is not derived from a single cell, but is a com-
pound formation, like the egg of Cestodes or Trematodes, in which
a germogen and vitelligen combine their products, for the com-
position of an egg.”

Stuhlmann (25) holds the same views as Brandt with regard
to the fate of the nutrient cells, and renews the old controversy
with regard to the germinal vesicle. The principal results at
which he arrives concerning it are summed up by him in the fol-
lowing words :—‘‘I have been enabled by a series of observations
on insects’ eggs to establish the extrusion of large balls from the
germinal vesicle which are afterwards lost in the egg-plasm.
Later the germinal vesicle disappears until at last at the upper
egg-pole we again find it as the segmentation nucleus ”’ f.

* Dass das Hi der Dipteren nicht yon einer einzigen Zelle abstammt, sondern
ein ebenso zusammengesetztes Gebilde ist als die Hier der Cestoden und Trema-
toden, bei denen Dotterstock und Keimstock ihre Producte zur Bildung des Hies
zusammenfliessen lassen ” (/. c. p. 209).

+ “ Es ist mir nun gelungen, an einer Reihe von Insekteneiern sicher einen
Austritt yon grossen Ballen aus dem Keimblaschen zu constatiren, die sich

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 423

Such an outstreaming of nuclear particles (Ballen) is un-
doubtedly seen in the lowest nucleus of the egg, but it also
occurs in the nuclei of the so-called nutrient cells, and in all the
nuclei of the various organs of the larva during their degeneration
in the first stages of the pupa. It is one of the most charac-
teristic phenomena of yelk-formation, whether in the egg or the
pupa, whilst it is quite unlike anything which has been observed
in relation to the well-known germinal vesicles of other animals.

Lastly, Henking (9) has quite recently figured and described
the ripe ovarian egg of the Blowfly with the nutrient cells
outside the chorion, and his figure has fortunately enabled me
to discover the error into which Brandt and his followers have
fallen. The appearance represented by Henking is an excep-
tional phenomenon which I have frequently observed. When
the eggs approach maturity they enlarge so rapidly that the
anterior pole of an egg is often pushed into a chamber above
it containing a half-developed ovum, which then assumes the
form of a cap over the anterior pole of the ripe egg. I
have sections which exhibit this phenomenon in several stages.
Whenever the young ovum in the chamber above the ripe egg
is present in an un-deformed condition the cap on the ripe
egg is absent, and whenever a cap is present there is no second
chamber in the egg-follicle. So many of the egg-tubes exhibit
transitional conditions in which the ripe egg impinges upon or
slightly indents the half-formed egg in the chamber above it that,
with good sections, I cannot believe anyone would have the
slightest doubt as to the nature of the phenomenon.

2. The Development of the Ovaries and Ova.

The earliest stage of development in which I have as yet seen
the ovaries of the Fly is in the four-day old pupa (Pl. XXVIII.
fig.4). In this stage they are apparently slightly in advance of the
stage described by Weismann as that of the seven-day-old pupa.
The discrepancy is probably due to the fact that I worked in
summer, and Weismann’s observations were made in winter. The
earlier stages of the pupa are well known to be greatly influenced
by the external temperature.

nachher im Hiplasma auflosen. Spater verschwindet das Keimblaschen von un-
seren Blicken, bis wir endlich am oberen Hipol den Furchungskern wieder-
finden” (7. ¢. p. 12).

424, PROF. B. T. LOWNE ON THE STRUCTURE AND

At this stage the ovary is pear-shaped ‘25m. in diameter and
34 to ‘4m.in length. It is enclosed in a thin but perfectly dis-
tinct cellular capsule (c). It consists of a stroma of small cells
less than 5 in diameter, enclosing the bundle of egg-strings (e.s).
These are closely packed together and occupy the anterior rounded
half of the ovary.

The posterior narrow part of the ovary (cl) is filled with small
round cells precisely like those which form the egg-strings, except
that the latter are slightly larger, 5 p.

The egg-strings present, even at this period, a narrower con-
stricted posterior and a more dilated anterior portion ; they are
like long narrow flasks, the neck measures 5p and the dilated
portion 15, in diameter. The whole consists of small closely
packed cells enclosed in a fine cuticular membrane. The necks
of the egg-strings appear to be open behind, where the cells of
the posterior part of the ovary, destined to form the duct, are con-
tinuous with those within the flask-like egg-strings. There is as
yet no trace ofa lumen in the solid rudiment of the oviduct.

The cells between and around the egg-tubes are, however,
already distinctly elongated and form a stroma, in which the egg-
strings lie, so differing entirely from the cells which form the
egg-strings and fill the calyx of the ovary.

The next stage of development is seen in the half-formed pupa,
about the tenth day (Plate XX VIII. fig. 5); at this stage the ovary
is apparently cup-shaped, it appears crescentic in lateral sections,
with the concavity of the crescent in front ; it has a diameter of
-5 m., bnt is still about ‘3 m. in thickness, measured from before
backwards in its thickest part. That part of the ovary not
occupied by the egg-strings 1s excavated by numerous chan-
nels; these form the calyx of the oviduct and cover the whole
convex surface of the organ. The egg-strings are now so broad
in front that 1 shall term them egg-follicles. The egg-follicles (os.)
are ovoid masses of small cells, each with a very narrow stalk (s¢.)
The stalk is apparently filled by a single row of cells, and its
investing cuticle is frequently transversely wrinkled, which often
produces an appearance of striation. Possibly this may have
given rise to the very remarkable view held by Schneider (23)
that the egg-tubes are developed within the alar muscles of the
dorsal vessel.

Each rudimentary egg-follicle is now surrounded by a pouch,

DEVELOPMENT OF THE OVARIES IN THE BLOWELY. 425

the ovarian follicle (of.), formed of fusiform mesoblastic elements.
This is the so-called peritoneal coat of Stein. The ovarian follicles
at this stage do not appear to open directly into the channels in
the calyx of the ovary, but to be closed by a cellular mass (#),
with which the stalks of the egg-follicles are continuous. This
cellular mass appears to me to be formed by the cells which filled
the posterior part of the ovarian capsule at the earlier stage of
development above described.

Between the ovarian follicles, which no longer lie close together,
the elements of the pseudo-yelk of the pupa (psy) are found in
abundance. These consist of globules (Kornchenkugeln) and
leucocytes ; they are derived from the breaking up of the fat-
bodies and the tissues of the larva. The pseudo-yelk, at this
period, forms the greater part of the bulk of the pupa. Indeed,
if such a pupa be opened it appears to contain nothing but a
milky fluid, in which all the tender half-developed imaginal tissues
are concealed.

There is as yet no differentiation of the contents of the egg-
follicles into epithelium, yelk, or germ-cells.

Even at the time when the fly is ready to escape from the pupa
the ovary remains in a condition which differs but little from the
stage last described, except in the form of the egg-follicles, which
now exhibit a narrow stalk and an ovoid middle portion constric-
ted above so as to form a small, but distinct nipple-like terminal
chamber (fig. 7, ¢.c.).

The peritoneal coat (of), or ovarian follicle, is also more
developed, is much thicker, and exhibits numerous tracheal
vessels (é.). There is still, however, no trace of differentiation in
the cells contained within the egg-chamber, and these only differ
from those in the terminal chamber in being slightly larger.

In the observation of the further developmental changes we
must have recourse to flies which have been -on the wing for
some time, and therefore the age of these insects is unknown.
Development progresses very slowly in captive insects, and as
these never unite with the males, there is no certainty that
development progresses at the same rate or in the same manner
as in free insects.

The next stage which I shall describe is represented in Pl.
XXVIII. fig. 6. The ovary now measures ‘35 m. in thickness and
is discoid. The calyx is very thin, as the ducts which form it
appear to be flattened by the lateral growth of the organ.

426 PROF. B. T. LOWNE ON THE STRUCTURE AND

The whole ovary has also altered its position ; the surface from
which the oviduct originates is now turned towards the ventral
aspect of the insect, so that the long axes of the egg-follicles are
transverse to the axis of the body. I shall still, however, call the
end of the egg-follicle which is nearest to the oviduct, posterior,
to facilitate comparison and avoid confusion.

The ovarian follicles are now more fully developed, and loosely
invest the posterior part of the egg-strings. The anterior part,
the terminal chamber, is closely covered by the anterior part of
the follicle, which is so thin that it can scarcely be traced as a
distinct layer. The posterior part of the egg-follicle exhibits a
distinct epithelial layer, which is continuous with the epithelium
of the calicine duct, and the follicles open freely into these duets.

The three parts of the egg-string are very distinct, within a very
fine cuticular tunica propria. The stalk is filled by a single layer
of epithelium ; there is no lumen and no double epithelial layer ;
but where the stalk enlarges near the egg-chamber there is more
than a single layer of cells, but these are irregularly arranged.

The egg-chamber, when fully formed, contains a group of
yelk-cells flattened by mutual pressure and surrounded by an
epithelial capsule (ec), which is continuous with the cells of
the stalk.

In some of the tubes a second egg-chamber is seen in process
of formation (Pl. XXVIII. fig. 7), with a cup-like epithelial in-
vestment. In others the first egg-chamber is in the same con-
dition; the cells destined to form the egg are still round and
scarcely differentiated from those of the terminal chamber. It
appears to me that the epithelial investment of the ovum grows up
from the stalk, and that all the cells of the terminal chamber
develop into yelk-cells.

The gradual transition from the small round cells of the
terminal chamber to yelk-cells is very marked, so also is the
upper edge of the epithelial cup (fig. 7, y), which ends abruptly in —
a thin edge, whilst the continuity of the epithelium of the stalk
and of the egg is equally distinct.

The large yelk-cells stain very deeply with alkaline carmine,
and the colour is not washed out by 5 per cent. solution of acetic
acid. They average 12 p in diameter, and all the cells in an egg
are precisely similar, and have nuclei which are rich in
chromatin granules. There is no cell with a clear nucleus, and
nothing which I can recognize as a germinal vesicle.

a 2...

bY (VS ee

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 427

In mature insects ready to lay eggs the ovaries occupy the
greater part of the cavity of the abdomen, There are about one
hundred egg-tubes (80 to 100) in each ovary, each having four,
five, or even six egg-chambers in different stages of develop-
ment (fig. 8).

A mature egg occupies the most posterior chamber (fig. 8, a), a
partially formed egg is seen in the second chamber, whilst the
third, fourth, and terminal chambers contain very rudimentary
ova. The whole terminates in a small empty, hollow, end-
thread (¢f).

The cuticular tunica propria closely surrounds the egg and
egg-strings, whilst the thin greatly distended egg-follicle passes
from one egg to the other, leaving a considerable space between
the eggs; it does not follow the outline of the egg-string
between successive ova. The egg-string between the second and
third chambers (¢p') is exactly similar to the egg-stalk of the
first chamber in the early stages of its development.

The two or three anterior chambers are filled with small round
cells like those of the terminal chamber or the whole egg-follicle
at an earlier period. The unripe egg in the second ‘chamber (ec’)
consists of large yelk-cells enclosed in an epithelial chorion.
The lowest cell in such eggs is, however, always much larger,
usually twice as large as the others, but its nucleus is also nearly
twice as large, and stains just as deeply. It also contains the same
kind of granules as the others. In these statements I agree in
no way with Brandt and Stuhlmann. Brandt states that the
nucleus of the lowest cell is large and clear, Stuhlmann that it is
very much smaller than the other nuclei, and that it is clear and
flattened against the chorion. According to my observations it
is neither one nor the other, and only differs from the nuclei of
the other cells in being larger.

The yelk-cells ultimately attain a giant size ; the largest cell,
when full-grown, measures 200 p» in its longest diameter, and
has a nucleus of 80 » in diameter. When the egg is enlarged
to about two thirds of its maximum size the granules in the
largest nucleus appear to stream out, the nucleus itself shrivels
and is ultmately lost, whilst the whole protoplasm of the cell
assumes a granular yelk-like appearance, in which the nuclear
granules can no longer be distinguished. The remaining cells

LINN. JOURN.—ZOOLOGY, VOL. XX. 32

428 PROF. B. T. LOWNE ON THE STRUCTURE AND

undergo the some changes, and soon become fused with each
other and with the yelk formed from the lowest cell.

The nuclei during these changes present a very variable
appearance; but all the changes of the nucleus are similar to
those which characterize the nuclei of the degenerating cells of
the larva, during the formation of the pseudo-yelk of the pupa—
a phenomenon well seen in the nuclei of the cells of the salivary
glands and fat-bodies of the larva during their histolysis.

I conclude therefore that the several cells from which the yelk
of the Dipterous egg is formed are of equal morphological
significance, that these all undergo histolytic changes, and so
form the yelk of the mature ovarian ovum.

So far as my observations go, there is no reason for supposing
one nucleus rather than another is the germinal vesicle.

When I first began this investigation, more than two years
ago, I looked for days in vain for some character by which I
might recognize the germinal vesicle. Sometimes one nucleus,
sometimes another presents a clearer contents and smaller
diameter, and frequently several nuclei appear to possess equal
claims in this respect to be considered the nucleus of the germ-
cell.

As the young ova approach the condition of maturity, the cell-
substance becomes more and more distinctly granular, the nuclei
lose their sharp contour, and exhibit what Stuhlmann describes
as an extrusion or outstreaming of nuclear particles, whilst
these are lost to view in the granular surrounding protoplasm,
and the cells themselves become fused into a single yelk-mass.
These changes commence in the lowest and largest cell of the egg;
but precisely the same changes afterwards occur in the remaining
cells as each attains its full growth.

The mature ova consist of a yelk surrounded by two mem-
branes, the vitelline membrane and the chorion. Such ova are
closely embraced by the structureless cuticular membrana
propria, and lie loosely in the distended ovarian follicle, which is
now a very thin-walled tube surrounded by a dense network of
tracheal vessels.

The yelk consists of an outer clearer layer (Pl. XXVIII.
fig. 11, @) and an inner granular substance (fig. 11, 6), but neither
contain any nuclei or cellular elements of any kind.

The clear peripheral layer of the yelk exists in the unimpreg-

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 429

nated eges whilst they still lie in the ovary; this layer was
described by Weismann, and called by him the Blastoderm plasma
(Keimhautblastem). He supposed that it is this layer which
forms the blastoderm. In my sections it projects in places as
if it possessed the power of amceboid movement, more especially
at the anterior egg-pole * (fig. 10, a); these may, however, be the
result of post-mortem contraction. The central granular yelk-
substance consists of small granules, 2 to 3 w in diameter
(fig. 11), imbedded in an apparently structureless, possibly in the
living egg semifluid, matrix. These granules are spheroidal,
stain deeply, and exhibit either a dark or light centre with
alterations of the focus of the microscope. In the ripe
unimpregnated ovum I have entirely failed to find any nuclei or
cellular elements of any kind, and I feel sure that if any such
elements were present they would be readily distinguished in my
sections.

It is not necessary for my purpose to enter into any details in
regard to the structure of the chorion and the nature of the
vitelline membrane; there are, however, some controverted points
upon which I would say a few words.

It is generally held that the epithelium of the egg forms the
chorion as an exudation from its inner surface (E. Korschelt, 14 ;
Weismann, 26). Whether this is so or whether the chorion is
formed from the cells themselves (Leuckart, 16), the manner in
which the ova leave the oviducts is entirely in favour of the
latter view. This is effected by the rupture of the remaining
rudiment of the egg-string between the ripe egg and the imper-
fect ovum immediately in front of it. Thus the thin tunica
propria and the epithelium of the egg descend in the ovarian
follicle and enter the oviduct together. The remains of the egg-
string attached to the unripe ovum in the ovarian follicle have
been seen and described by several observers, notably by
Miller, Landois (15), and Leuckart (16). I hold therefore that
whether the cellular epithelium is shed with the thin cuticular
ege-sheath in the oviduct, or whether it remains as the chorion

* The polar globules of Robin, which he described as formed by budding
and fission, are possibly only mobile processes of this layer in a contracting
yelk.

32*

430 PROF. B. T. LOWNE ON THE STRUCTURE AND

itself, it belongs entirely to the ovum, and cannot be regarded as
the epithelium of the ovarian follicle, which is quite distinct and
remains in the follicle.

I am also inclined to regard the vitelline membrane as the
cuticular exudation from the inner surface of the epithelium of
the ovum and the chorion as the modified epithelium itself. The
cuticular sheath which leaves the ovarian follicle with the egg
is, I have little doubt, the epichorionic membrane described by
Leuckart, Robin, and Kolliker. The shedding of the outermost
covering of the egg, probably the epichorionic membrane, and
possibly also of the epithelial chorion, was observed by Brandt
in the Field-crickets in transit through the oviduct, forming
what he designates corpora lutea.

The micropyle-canal, which, in the Diptera, extends the whole
length of the dorsal surface of the egg, is an infolding of the
chorion (Pl. XXVIII. fig. 9). It is extended over the anterior
ege-pole (fig. 10), forming a considerable chamber in the floor
of which the micropyle is situated (fig. 10, m).

The micropyle (figs. 12 & 13) is a small, almost quadrilateral
opening (fig. 12), 2°55 pw in diameter; it is surrounded by a
number of radiating folds which project on the outer surface of
the chorion, and by a circular area composed of small hexagonal
cells. These correspond in size to the hexagonal fields with
which the rest of the chorion is sculptured.

The open micropyle-canal is brought into relation first with
the orifices of the gum-glands, and later with those of the
spermatophorous capsules during the descent of the egg through
the genital canal. Henking (9) found spermatic filaments in the
micropyle-canal.

3, The Oviducts and their Appendages.

The general form and arrangement of these parts is well
known, so that the following description will suffice to indicate
their arrangement for my present purpose.

The ovarian ducts are two in number (fig. 2), and these form
a common oviduct (od) by their union. The common oviduct

opens into the pouch-like anterior extremity of the vagina on its
dorsal aspect.

— eo a ae ys

—_———— a 1

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 431

Stein describes it as opening on the ventral surface in Beetles.
I formerly fell into the same error; and it is exceedingly difficult
in dissections to determine this point. Sections of the entire
insect show at once the true relations of the parts.

The pouch-like anterior part of the vagina is very distinct from
the posterior part ; that portion of it in front of the orifice of
the common oviduct (fig. 3, 5) in the young insect is the bursa
copwlatrix of authors. The bursa in the egg-laying insect is no
longer distinct, but forms the anterior part of the vaginal pouch.

I shall call the vaginal pouch the uterus, a term applied to it
by Palmén (21) to distinguish it from the posterior tubular part
of thevagina. If the term is not morphologically, it is at least
physiologically correct, as an egg is frequently retained in it
until the embryo is ready to escape from the shell.

There is at present some uncertainty as to the exact manner in
which the common oviduct is developed; although it is quite
certain that the ovarian ducts are developed from the posterior
portion of the ovary, as Palmén has distinctly shown (21). The
same observer also shows that the vagina, uterus, and their appen-
dages are formed by an invagination of the external integument,
or rather of the hypoderm. My own observations entirely con-
firm Palmén’s statements. Although I have not been able to
trace the development of the common oviduct, its structure and
the manner in which the common duct of the testicles is formed
in the Fly (Weismann, /. c. Taf. xiv, fig. 68) indicate that it is
formed from the prolonged posterior parts of the ovaries.

So far I have stated nothing concerning the anatomy of
these parts which has not been frequently observed and generally
admitted. I must now, however, enter into some details which
are not, so far as I know, to be found elsewhere.

The common oviduct in the Blowfly terminates in two distinct
enlargements (figs. 1 and 3). The more anterior is due to a
thickening of its muscular coat where a thick retractor muscle
(m) is inserted into it. This witndraws the parts with the ovi-
positor. The second or terminal enlargement (0s) is, however, a
pouch or bulb lined by a greatly plicated intima, and capable of
distention, so that it encloses the entire egg during its passage
through the oviduct. One egg is frequently found in this section
of the oviduct whilst another occupies the uterine cavity.

432 PROF, B.. T. LOWNE ON THE STRUCTURE AND

The great interest of this pouch is that the gum-glands (g/) or
colleterial glands, as they are sometimes called, open by two
slender ducts (d) into it, and not, as is usually believed, into the
uterus itself.

Although I have frequently satisfied myself of this, both by
section and by careful dissection, this point is of such importance,
that I shall enter into an examination of the views of previous
writers with regard to the termination of these ducts.

Itis quite possible that several distinct glands have been con-
founded under the term gum-glands ; indeed it is generally used
for any accessory gland connected with the sexual canal. These
glands are generally described as opening into the vagina or
uterus. Stein (24) gives a great number of figures representing
the oviducts, uterus, and appendages in the Coleoptera ; in many
it is difficult to identify the gum-glands. In Hydrophilus (1. ce.
Taf. iv. fig. 111) he represents the gum-glands as opening into the
upper part of the ovarian duct. They are branching tubules
which evidently form part of the ovary itself; and, judging by
his excellent figure, are identical with the so-called gum-glands in
the Blowfly.

Except in the Hydrophylide, Stein considers the gum-glands
as a portion of what he terms the “ apparatus of fertilization”
(Befruchtungs-Organe), and represents them as if they opened into
the spermatophorous capsules or their duct; although in many
cases it is almost evident from his figures that they open into the
oviduct. In some.of his figures the spermatophorous capsules
are represented opening into the oviduct (Taf. 1. fig. vi), whilst
in others they are correctly represented opening into the uterus,
whilst the gum-glands open into the oviduct (Taf. 11. figs. i, ii,
and iii).

Tracing the gum-gland in the Blowfly from its ovarian ex-
tremity, it lies first under and close to the ovarian duct; it then
leaves this duct and comes into relation with the spermatophorous
capsule, around which it forms a loop. The duct of the gum-
gland commences at the termination of this loop, and is easily
overlooked, as itis in close contact with the duct of the spermato-
phorous capsule, round which it turns and runs forward in
close contact with the dorsal wall of the uterus and oviduct to
terminate in the bulb of the oviduct. It is not difficult to under-

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 433

stand how this duct has been overlooked, or how it has been
supposed that the glands open into or with the seminiferous
capsules. The gum-glands have also probably been confounded
with true vaginal glands, which appear to exist in some insects.

I shall again refer to the gum-glands in a special section of
the present paper in relation to their structure and functions.

The uterus (figs. 1, 2, 3, wt.) is a thick-walled sac lined by a
strong cuticular membrane, very different to the thin cuticular
membrane lining the oviduct. It has a diverticulum or pouch
(p) on its dorsal wall immediately behind the orifice of the
common oviduct. This pouch (sacculus, figs. 1, 2, and 38, p)
is lined by a very thick laminated cuticle with a projecting
median ridge which appears to divide it into two lateral pockets.
Each of these pockets opeus behind into the uterus, and is
usually filled witha clear colloid mass, which stains very deeply
with alkaline carmine. It has all the appearance of being the
same material as that which cements the eggs together when
they have been deposited. The same contraction of the uterus
which expels the egg would certainly expel some of this material
from the uterine pouch.

4. The Structure of the Gum-glands.

Although I have used the term gum-glands to designate
these organs, it will be seen that there is nothing in their
structure to justify its use. And although they are usually
regarded as secreting-glands which form a glue or cement for
the attachment of the eggs, a function first apparently ascribed
to them by Burmeister (7) and afterwards by Loew (18), Stein,
who has examined these structures with more care perhaps than
any other writer, entirely discards the view. He regards the
so-called gum-glands as accessory organs of fertilization except
in the Hydrophilide, where they open into the calyx of the
ovary ; and, curiously enough, disregarding the extreme impro-
bability that gum-glands would open in such a situation, makes
an exception in these insects, and regards the glands as gum-
glands. Stein further identifies these glands in the Diptera
with his “ glandular portion of the organs of fertilization.”

434 PROF. B. T. LOWNE ON THE STRUCTURE AND

With regard to the histology of these glands, very little, if
anything, can be said to have been recorded of a satisfactory
character. Stein gives several very remarkable figures (Jl. ¢.
pl. ix. figs. i, v, and xii) of their histological structure, with the
following description :—

“The fine structure of the glands is nearly the same in all
Beetles; they belong to that class of glands which yield a
fluid secretion, and which are tubular, follicular, or exhibit
bladder-like cavities. In the gland-follicles the proper elements
of the gland form a manifold layer of nucleated cells which pre-
pare the secretion. Between these cells very fine wavy canals
spread into the follicles, formed as outgrowths of the epithelial
coat (of the central cavity), and terminate either in blind ends or
within the cells*.

“In general the contour of the gland-follicles is the same as
that of the epithelial coat of the central cavity...... The
secreting-cells lie between the epithelial and peritoneal coats [the
italics are mine] without order, near and over each other, and
not united together. In form they are round, oval, or egg-
shaped; in the latter case the blunt end is turned outwards, and
the outlines of the cells, when one examines the entire follicle
under a certain pressure, are not generally distinctly seen, so
many lie over each other, and the cells, owing to their granular
contents, are so opaque”’ (pp. 102, 108).

Leuckart (16a), in his memoir on the Pupipare, gives a
figure of the corresponding gland of Melophagus ovinus, which,
although on a much smaller scale, represents a similar appear-
ance, and gives a description which corresponds nearly with
Stein’s.

These figures and descriptions are very difficult to understand,
except on the supposition that both Stein and Leuckart examined
glands with a quantity of adherent fat-cells. The fat-cells of the
ovary form a large mass on its posterior aspect, and closely sur-
round and adhere to the gum-glands. These fat-cells, when half
empty, as they always are in the egg-laying female, exhibit

* “‘Zwischen diesen Zellen verbreiten sich am Follikel sehr feine geschlan-
gelte Kanale welche von Austulpungen der Epithelealhaut gebildet werden, und
die nach aussen entweder blind endigen, oder an einer Zelle endigen.”

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 435

appearances, in optical section, which could be interpreted without
difficulty, as Stein and Leuckart have interpreted them ; possibly
the fine tubes are the fine trachex of the fat-glands, whilst the
cells figured by Stein are undoubtedly those of the fat-body, of
which I give a figure (Pl. XXVIII. fig. 15) for comparison.
Sir John Lubbock (19), describing the corresponding glands
in Ooccus Persice, gives a totally different description. He
Says :—

“They are six in number, four large and two small, the latter
bemg apparently attached by a short stalk to the peduncle of the
large one which is furthest from the vulva. They lie three on
each side, and their ducts open into the egg-canal close together
and about halfway between the vulva and the division of the
egg-canal into two oviducts. The internal structure is very
distinct and interesting. It consists of many cells lying loose in
the internal cavity, and resembling very much in form, size, and
appearance the vitelligenous cells of the egg-follicle.”

I have been unable to find any other published details on the
structure of these glands, which I shall now give from my own
observations.

The gum-glands in the Blowfly are simple tubes, tortuous
rather than convoluted, 2 m. in length, with an average dia-
meter of ‘175 m. ‘They have a glistening white appearance,
and are beaded over the surface from the projection of the cells
lining them. In sections these glands are seen to consist of an
outer musculo-cellular coat, like the so-called peritoneal coat of
the oviduct. This is lined by a single layer of large epithelial
cells. The lumen is filled by a granular fluid or semifluid sub-
stance. ‘This is coagulated by alcohol, the granules suspended
in it are blackened by osmic acid, and the intergranular material
is scarcely stained by alkaline solutions of carmine. In this
respect it differs entirely from the substance contained in the
uterine pockets and from that with which the eggs are cemented
together.

The epithelial cells which surround the lumen of the gland
are irregular in form and measure, on an average, 80 pm in
diameter, and from 30 to 40 pw in thickness. Many of these
cells contain very remarkable spherical corpuscles, usually
one in each cell (figs. 15 to 20). Besides these, some of the

436 PROF. B. T. LOWNE ON THE STRUCTURE AND

cells exhibit an oblong nucleus surrounded by a clear area
(Pl. XXVIII. fig. 16).

In females in which the ovary is still without ripe ova (fig. 17)
there are no corpuscles in the cells, but nuclei in an active state
of division. Each nucleolus or each group of nucleoli, two or even
four (figs. 17 and 18,2), is surrounded by a clear area. In some
_ there is a small speck of deeply stained chromatin close to the
nucleolus (fig. 19). :

The epithelial cells (fig. 20, e) consist of distinctly reticular proto-
plasm and stain readily. The largest of the contained corpuscles
measures 25 to 30, in diameter. A fully formed corpuscle
exhibits a clear outer zone (fig. 20, d), 4 in breadth, with a
distinct radial striation. This clear outer zone closely surrounds
a finely granular contents (¢) which stains feebly, and lying in it,
usually near one side of the corpuscle, a clear vesicular spot (6)
5 pw in diameter, with a bright highly refringent spherule 2°5 » in
its centre (a).

These corpuscles have, in point of fact, the closest possible
resemblance to the germ-ova of other animals.

The relation of these corpuscles to the nuclei of the containing
cells must at present remain a matter of conjecture. It appears
to me probable that one of the nuclei of the cells in the young
gland, when there are two or more, develops the corpuscle whilst
the others remain quiescent. The nuclei both of the young and
mature cells stain deeply, whilst the vesicle and highly refractive
body in the corpuscle remain unstained.

In several instances I have seen an appearance which leads me
to believe that the corpuscles when mature are discharged from
the cells in which they are developed into the lumen of the gland
(fig. 20). Either empty spaces in the cells or a distinct fissure
surrounding the corpuscle which lies close to the lumen of the
gland are not uncommon. In some preparations the corpuscles,
or some of them, have evidently fallen out in mounting the spe-
cimen.

On other occasions I have seen what appears to be a rupture
of the clear external zone, and a protrusion of the contents of
the corpuscle into the lumen of the gland. There is certainly a
close similarity between the contents of these corpuscles and the
material which fills the lumen of the gland.

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 437

I have, however, been unable to find either the corpuscles
themselves or the vesicular body they contain imbedded in this
material; but bright refractive nuclear particles like the central
highly refringent body of the corpuscles undoubtedly exist
in it.

5. Theoretical Considerations and Conclusions.

I am led by my observations to the following unexpected
conclusions :—

The ovarian eggs in the Blowfly, and probably in other insects,
are yelks, and contain no germ.

The so-called gum-glands are in reality germ-glands in which
the germ-ova are developed.
_ These germ-ova pass into the yelks during their passage through
the oviducts either (a) as naked germinal vesicles, or (0) as female
| pronuclei.
I shall now examine these hypotheses in relation to the work
73
:
]

of previous investigators, and discuss their probability.

1. I have already shown that much difficulty exists in attempt-
ing to reconcile the observations of previous writers on the deve-
lopment of the ova in insects. So recently as 1881 Prof. Balfour
(1) regarded the whole question as unsettled, and contented
himself with stating that the relation of the ovum to the germogen
and the relation of the yelk-cells to the ovum are points which
have been especially controverted. I make this observation to
show that the great number of researches which have been re-
corded by no means settle the question, which therefore still
remains an open one.

2. The existence of true germ-ova, if such they are, in what
has always been regarded as an accessory gland, although unex-
pected, is not inconsistent with the probable genetic relations of
the Insecta. |

Recent embryological observations show that the Insecta
exhibit resemblances, sufficiently startling, to the Nemertid worms,
and to the Trematodes generally, rather than to the Nematoid
worms. ‘This is seen by a comparison of the early developmental
stages of Lineus (Barrois (2)) and Chetognatha (Kowalevski) with
those of Musca\(Kowalevski, Biitschli), my own unpublished obser-

438 PROF. B. T. LOWNE ON THE STRUCTURE AND

vations being in complete accord with those of the above-named
authors.

3. With regard to the structure of the so-called gum-glands of
Musca and probably of other insects, I would submit that a com-
parison of the description of the gum-glands, as I have given it,
with the ovary of some Arachnids, Crustacea, and Worms is not
without interest,

That such ova are developed within cells and present appear-
ances exactly similar to those I have described, is pretty evident
from the figures of the young ovary of Spiders given by Stuhl-
mann (25), plate ix. figs. 190 and 197, and plate x. figs. 214,
215, and 216; whilst similar appearances are represented by
Van Beneden (8, 4, and 5) in the germogen of the solid-bodied
Worms and some Crustaceans. It is true these authors puta
different interpretation on the relation of the ova to the epi-
thelium of the germogen; but the fact remains that their figures
are such as to show the close resemblance of the germogen in
these animals and the so-called gum-glands of the fly.

4. With regard to the morphology of the gum-gland, so long
as we remain ignorant of the precise manner in which this struc-
ture and the common oviduct are developed, its morphology will
remain more or less doubtful. I have already given my reasons
for the belief that the common oviduct is part of the primitive
ovary; and this opinion is generally held. The condition of the
parts in the Hydrophilide is an undoubted indication, I think,
that the gum-glands are merely modified ovarian tubules, and
have a similar origin from the primitive ovary ; the connexion
which persists between these glands and the calyx of the ovary in
the fly is not, I think, unimportant in this relation. In the
Hydrophilide the gland is comparable with the germ-gland of
the Crustacea. Compare the figures given by Van Beneden (5)
and by Stein (24).

It is true that Palmén (21) states that the appendicular glands
(the gum-glands and the spermatophorous capsules) have the
same origin from the ectodermal invagination as the vagina and
uterus; but his observations are general, and he believed the
gum-glands to open into the uterus.

5. Supposing my corpuscle to be a germ-ovum, its discharge
from the cell in which it is developed may_be considered an un-

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 439

paralleled phenomenon. I am by no means sure that it is so.
The changes in the germarium of the Trematodes described by
Van Beneden (5) appear to me to indicate a similar condition.
The nature of the germ-ova in these has been discussed with
ereat heat, some holding that the germ is a naked nucleus, and
others that it retains a thin, scarcely demonstrable, layer of
protoplasm around it.

6. Until the actual passage of such a germ into the yelk has
been repeatedly observed, I admit that a serious hiatus exists in my
hypothesis. I am far from sure that the phenomenon has not
been already observed ; at any rate a passage in Leydig’s mono-
graph (17) on the ovaries and seminal pouches of insects is
worthy of note in this relation. Speaking of the chorion of
Musca domestica, he says, ‘‘ this has at the upper pole of the egg
a prominent micropyle which appears blocked by a highly refrac-
tive corpuscle. The corpuscle is not present in all the eggs, and
may be perhaps an exuding yelk-drop”*. I would ask, may it
not equally have been an entering germ-yelk ?

6. Bibliography.

As the bibliography of the subject is given in extenso by Dr. F.
Stuhlmann, 1886 (25), and Dr. A. Brandt, 1878 (6), I shall give
only a list of the works quoted or directly made use of in this
paper.

(1) Baurour, F. M.—Comparative Embryology. 1881.

(2) Barrois.—* L’Embryologie des Némertes.”” Ann. Sci. Nat. sér. 6,
tom. vi. 1877.

(3) BENEDEN, E. van.—“ Recherches sur l’Embryogénie des Crustacés.”’
Bull. de l’Acad. R. de Belgique, 2° sér. tom. xxviii. 1869.

(4) BenEDEN, E. v.—Ibid. tom. xxix. 1870.

(5) Benepen, E. v.—“ Recherches sur la composition et la signification
de l’Ciuf.”” Mém. couronnés et Mém. étrang. Acad. de Belgique,
t. xxxiv. 1870.

* “Dieselbe hat am oberen Hipol eine vorstehende Mikropyle und wie
verstopft durch ein fettglanzendes Zapfchen (Taf. iii. fig. 130, 140). Dieses
Korperchen ist nicht bei allen Hiern vorhanden und entspricht vielleicht einem
herausgetretenen Dottertrdpfchen ” (7. ¢. p. 35),

440 PROF. B. T. LOWNE ON THE STRUCTURE AND

(6) Branpt, ALex.—Ueber das Ei und seine Bildungsstiitte. Leipzig,
1878.
(7) BurMEISTER, HERMAN.—A Manualof Entomology. Translated by
Shuckard. London, 1836.
(8) Cuaus, C.—‘‘ Beobachtungen wber die Bildung des Insecteneies.”’
Z. f. w. Z. Bd. xiv. 1864.
(9) Henxine, H.—“ Dieersten Entwicklungsvorginge im Fliegenei und
freie Kernbildung.” Z. f. w. Z. Bd. xlvi. 1888.
(10) Huxtey, T. H.—Anatomy of Invertebrate Animals. London,
1887.
(11) Huxuey, T. H.—‘ On the Agamic Reproduction and Morphology of
Aphis.’’ Trans. Linn. Soe. vol. xxii. 1859.
(12) Kouuiker, A.—Observationes de prima Insectorum Genesi. Turici,
1842.
(13) KorscHe.t, E.—‘ Ueber eimige interessante Vorginge bei der
Bildung der Insecten Fier.” Z. f. w. Z. Bd. xlv. 1887.
(14) KorscuEe.t, E.— Zur Bildung der Eihiillen der Micropylen und
Chorionanhange bei den Insecten.”” Nov. Acta Acad. Ces. Leop.-
Car. 1887.
(15) Lanpots, L.—* Anatomie des Hundflohes.”” Nov. Acta Acad. Ces.
Leop.-Car. 1866.
(16) Leucxart, R.—‘“ Ueber die Micropyle und den feinern Bau der
Schalenhaut bei den Insecten Ejiern.”” Archiv f. A. und P.
1855.
(16a) LeucKxart, R.— Fortpflantzung und Entwicklung der Pupiparen.”’
Halle Abth. Nat. Gesellsch. iv. 1858.
(17) Leypie, F.—‘‘ Der Eierstock und die Samentasche der Insecten.”’
Nov. Acta Acad. Ces. Leop.-Car. t. xxxii. 1866.
(18) Lozw.—Hore Anatomice. Beitrage zur genaueren anatomischen
Kenntniss der Evertebraten, Abth. 1. Posen, 1841.
(19) Lussock, Sir J—‘ The Ova and Pseudova of Insects.’’ Phil. Trans.
1859.
(20) Meyer, H.—“ Ueber die Entwicklung des Fettkérpers der Tracheen
und den keimbereitenden Geschlechstheile bei den Lepidopteren.”’
Z. f. w. Z. Bd. 1. 1849.
(21) Parmén.—Ueber paarige Ausfiihrungsginge der Geschlechtsorgane
bei Insecten. 1884.
(22) Rosin, C.—‘ Sur la Production du Blastoderm chez les Articulés.”?
Journal de Physiologie, tom. v. 1862.
(23) ScHNEIDER, A.—‘‘ Ueber die Geschlechtsorgane der Insecten.”
Zool. Beitraige, Bd. 1.
(24) Svein, F.—Vergleichende Anatomie und Physiologie der Insecten

S --------5

iyo ee

BLL del:adnat,

OVARIES OF BLOWF

West, Newman imp.

: |) =" oo

DEVELOPMENT OF THE OVARIES IN THE BLOWFLY. 4A

in Monographien bearbeitet. Erste Monographie. Die weiblichen
Geschlechtsorgane der Kifer. Berlin, 1847.
(25) SruHLMANN, F.—“ Die Reifung des Arthropodeneies nach Beo-
bachtungen an Insecten, Spinnen, Myriapoden und Peripatus.”’
F Berichte der naturforsch. Gesellschaft zu Freiburg, Bd. i. 1886.
(26) Weritsmann, A.—Die Entwicklung der Dipteren. Leipzig,
= 1864.

DESCRIPTION OF PLATE XXVIII.

Figs. 1, 2, and 3. The uterus and its appendages in the adult egg-laying
insect. The different parts are indicated by the same letters in all the
figures.

Bursa copulatrix.
Duct of the gum-gland.
gi. Gum-gland.
m. Retractor muscle of oviduct.
o. Ovary.
od. Oviduct.
os. Terminal enlargement of the oviduct.
p. Uterine pouch.
(i: s. Spermatophorous capsule.
ut. Uterus.
Vagina.
Attachment of the gum-gland to the ovary.

v
2
Fig. 1. Dorsal view of the uterus. x 20.
2. The same, with the ovaries. The parts rendered semitransparent
with glycerine, and seen by transmitted light. x 10.
3. A median section of the uterus in the vertical antero-posterior

plane.
4. The ovary of a three-day-old pupa. c, capsule; c/, cells of the
b calix ; ¢s, egg-strings. x 200.

5. Two of the egg-follicles from a ten-day-old pupa. /, calicine duct ;
os, ovisac; st, stalk of egg-follicle of the ovary; Psy, pseudo-
yelk-granules of the pupa; 2, cell-mass to which the stalks of
the ovisacare attached. x 300.

6. An ovisac from a young fly. ¢c, terminal chamber; ec, egg-
chamber; s¢, stalk of ovarian follicle; c¢/, calicine duct; ¢r,
trachea. X 200.

7, Another ovisac from a young fly, a little more advanced. y, edge of

| epithelium of egg ; ec’, ec’, young egg-chambers. The other letters

: _ asin figs. 5and 6. x 300.

MR. W. E. HOYLE ON THE DEEP-WATER

The egg-follicle from the ovary of a mature egg-laying insect.
x 200. .

. Transverse section of an egg. mc, micropyle canal; ch, chorion ;

v, vitelline membrane ; cl, clear yelk ; y, granular yelk. x 30.

. A longitudinal section of an egg. mec', chamber at anterior pole of

the egg; m, micropyle.

. A section of the yelk. a, clear margin; 0, granular yelk. x 400.

. The micropyle. x 400.

. A section through the micropyle. x 400.

. The testes of a larval blowfly, showing the union of the prolongations

from which the duct is developed: after Weismann.

. The gum-gland and some of the adjacent fat-body. J, lumen of the

gum-gland; ¢, epithelium of gum-gland; 0b, capsule of fat-cells ;
a, ¢, d, stellate and flask-shaped cells enclosed within the capsule.
x 200.

. Transverse section of the gum-gland of the mature insect.

Transverse section of the gum-gland of the immature insect.
19, 20. Epithelial cells from the gum-gland, with the contained
corpuscles and nuclei in different stages of development.

On the Deep-water Fauna of the Clyde Sea-area. By Witt1am

Hoyzir, M.A. (Oxon.), F.R.S.E., Keeper of the Man-

chester Museum. (Communicated by Jonn Murray, LL.D.,
Ph.D., V.P.RB.S.E., F..8.)

[Read 4th April, 1889.]

(With Mar: Pirate XXIX.)

Since the establishment of the Scottish Marine Station in the
year 1884, Dr. John Murray has conducted an extensive series of
dredgings in the greater number of the lochs of the west coast of
Scotland. During these operations he was struck, as Forbes had
been before him, with the restricted distribution of certain forms,
as well as with the fact that some species occurred nowhere off
the British shores except in these depressions.

In the summer of last year, Dr. Murray suggested that I

FAUNA OF THE CLYDE SEA-AREA. 443

should continue these investigations, and endeavour to render
them as complete as possible during the months of July and
August, offering at the same time to give me the use of the
steam-yacht ‘ Medusa’ for dredging and trawling, and to allow
me the use of the materials which he had already accumulated.

Circumstances fortunately allowed of my accepting this offer,
and during the two months just mentioned I made Millport, on
the Island of Cumbrae, my headquarters, and thence made ex-
cursions to all the different parts of what is now known as the
“‘ Clyde sea-area.”

The physical configuration of this region has been very ably
described by Dr. Hugh R. Mill*, whose communication is illus-
trated by an admirable orographical and bathymetrical chart.
He regards the “‘ Clyde sea-area ” “‘as bounded on the south by
a line drawn from the Mull of Cantyre to Corsewell Point in
Wigtownshire, almost coinciding with the contour of 50 fathoms ;”
and within it he defines seven deep-water basins, which have a
depth exceeding 20 fathoms, and are separated from each other
by ridges, considerably shallower than the extreme depths of the
basins themselves.

1. The Arran Basin extends on either side of the north of
Arran, and up into lower Loch Fyne, being in shape like the
letter X. In the sequel I have regarded it as subdivided into
four portions, which may be termed respectively the ‘“‘ Brodick,”
“ Cumbrae,” “ Kilbrennan,” and “‘ Inchmarnoch ”’ basins.

The last of these is the deepest, and, indeed, attains the
greatest depth found anywhere in the Firth, namely 107 fathoms
off Skate Island. The Kilbrennan and Inchmarnoch Basins are
not so distinctly marked off from each other as are the remaining
ones, the channel which unites them just reaching the 60-fathom
line, the extreme depth of the former being 85 fathoms. The
Brodick basin, which is off the bay of the same name, has a
depth of 92 fathoms, whilst the Cumbrae basin descends only to
62 fathoms.

2. Upper Loch Fyne is 25 miles in length, and has a depth of
about 80 fathoms off Strachur.

3. Loch Striven runs up into the mainland due north of
Rothesay, and attains a depth of a little over 40 fathoms.

* Scottish Geogr, Mag. iii. pp. 1-7 (1887).
LINN. JOURN.—ZOOLOGY, VOL. XX. 33

44.4. MR. W. E. HOYLE ON THE DEEP-WATER

4. The Dunoon Basin occupies the channel of the river from
the extremity of Great Cumbrae northwards, and extends up
into the lower stretch of Loch Long. Its greatest depression off
Dunoon is 56 fathoms.

5. Loch Goil is only about 4 square miles in area, and its ex-
treme depth is 47 fathoms.

6. Upper Loch Long is of about the same extent, but has a
depth of only 35 fathoms.

7. The Gareloch has an area of about 5 square miles, and is
23 fathoms in depth.

The object which I set before myself was to ascertain as fully
as possible the fauna of each of these depressions, limiting them
by the contour-line of 20 fathoms, and then witi all the mate-
rials available to draw up comparative lists, and to endeavour to
discover their relations to each other.

Unfortunately I have been unable to make much use of the
published works of my predecessors, owing to the form in which
their results are stated. ‘Fairly common in depths of 5-25
fathoms,” with a few localities appended, is the type of a phrase
which occurs continually, but is, for the purposes of the present
inquiry, quite useless. The cases in which I have drawn infor-
mation from sources other than the records of the Scottish
Marine Station are all indicated.

The specimens collected by Dr. Murray had been sent from
time to time to the British Museum, and he had received from
the authorities of that institution lists of these consignments,
along with a number of named duplicates, which were of great
help in the identification of my own subsequent acquisitivus. I
have to acknowledge, with my sincerest thanks, the assistance I
have received, not only from the staff of the British Museum, but
from several other friends. Mr. David Robertson and the Rev.
Canon Norman, whose extensive knowledge of the British marine
fauna is well known, were at Millport during the greater part
of my stay there, and I had thus the advantage of being able to
consult them constantly. Messrs. Isaac C. Thompson and W.S.
M‘Millan, of Liverpool, have been good enough to draw up lists
of the Copepoda for me, and Prof. Herdman has given me the
benefit of his acquaintance with the Tunicata. To Dr. John
Murray, as above stated, I owe the suggestion of the present in-
vestigation, as well as the means of carrying it out.

pe ng tS ee
U

FAUNA OF THE CLYDE SEA-AREA. 445

Before proceeding to enumerate the species obtained, it seems
advisable to say a few words regarding the mode of procedure
adopted in collecting, for this probably explains certain general
features in the results obtained. In the great majority of in-
stances the instrument employed was a shrimp-trawl, which was
found on the whole more convenient for working in these loca-
lities than the dredge, owing to its bringing up a less amount of
mud and a greater variety of forms. Certain groups, such as
the Mollusca, are not obtained in such large numbers as by the
dredge, and hence the list of these animals obtained is small as
compared with that known to inhabit the district.

A tow-net was generally attached a short distance above the
trawl, so as to capture any Crustaceans which might be swimming
just above the bottom. These were almost invariably found to
be of a different species from those taken in the nets which were
dragged at the surface.

STATEMENT OF RESULTS.

For convenience I have drawn up the results in the form of a
Table, with a column corresponding to each basin. The figures
show the range in fathoms in that particular region, whilst there
is appended in another column a sketch of the distribution of
each form outside the British area. Those facts which have been
taken from the published writings of others are indicated by
italics. A note of interrogation indicates that I have obtained
the species in that locality, but the record of the exact depth
has been lost. d means that dead shells, not living specimens,

were obtained.

33*

446

34.
30.

MR. W. E. HOYLE ON THE DEEP-WATER

PIscEs.

. Pristiurus melanostomus

(Bonap.)
Acanthias vulgaris, Risso ......

. Raja clavata, LD. ........eeeeeeeees

fullonies, Liss. c.ntepe oe ae

maculata, Montag. ......+.

. Lophius piscatorius, L. ..-.....-

. Cottus bubalis, Huphr...........+-
. — Lilljeborgii, Collett.........

scorpius, L. .....-..sseseeeee

. Trigla gurnardus, L. ......-..ee

. Triglops Murrayi, Gthr. .........
. Agonus cataphractus, L. .........

. Liparis liparis (Z.) ....-.++.+0+++

. Gobius Jeffreysii, Gthr. «....-++-

MOMUbUS, (G:770.. sin seins 2-2

. Callionymus lyra, LD. ...+..+.+6

maculatus, Raf. .........0+

_ Centronotus gunellus (Z.) .....-

_ Sticheeus lampetreformis ......
. Gadus eeglefinus, LZ. .......:e+see-

luseus, i.c..c0 sence saccm esse
Esmarkii, Nilss. .......+.+--
merlangus, DL. ......++-0e+0e

ya PINS, 210, saison sense =k

WHOTE MUA, Cie. eee dens sere ee

. Merluccius merluccius (L.) ...

. Molva molva (L.)....sseesesseeees
. Onus cimbrius (L.) ......seeeeeee

maculatus (Risso) ......+--

. Hippoglossoides limandoides

(Bi.)
. —— platessoides (Fabr.).........
Rhombus megastoma (Donov.)

norvegicus, GtKr.......+++++
punctatus ( Bl.) .....-ss00
Pleuronectes cynoglossus, L. ...

ARRAN BASIN.

| nS EES SS ee

Bee Kil Inch
‘ brennan | marnoch
Basin. : ‘
Basin. Basin.

Cumbrae
Basin.

ceteee | (wre wieterm © 1)". ) pieeeetaias ee eee

PTT Goce i Or 8h OSGi

80-90 | 26-46 | ceseee | seeeee

qenees 65 40 aiemRe
seieee 40 45-100 kale
30-60 | 26-46 | 80-100 | .....
biceen 40 ‘aes omnes
Peek AB} ghee ak
fetes ol here 60

ee 46-70 | 80-100 sedan

FAUNA OF THE CLYDE SEA-AREA. 44.7

a — si: eg General Distribution.
ng
wenes ‘ hema biti ee. | “Seas of Europe, being common in
the Mediterranean.”
Ee eee ee Temperate seas of Northern and
Southern hemispheres.
= 2 a Around the seas of Europe, Medit.
ES ee eee Western Europe, rare in Medit., Ma-
deira.
sic 30-40 ee asia ..-es. | European coasts to Medit.; Madeira.
= 30-40 apes ania ..-... | Rare north of 60° N., Medit. to Cape

of Good Hope; Newfoundland to
Cape Hatteras.
| PS ee ee Arctic regions in both hemispheres ;
Baltic, North Sea, France, Spain.
| RS | eee. eee Norway and Feeroes (new to Britain).
| gaalie S | ere Arctie and N. Atlantic in both hemi-
spheres ; Baltic, North Sea.
| Se eee Deere Baltic; West Europe from Norway
to Medit.
Wear f) ecete- | ...<0- New species; an arctic genus.
oo ee eee Iceland, Norway, Baltic, German
Ocean.
EES Se eee Polar Regions to Cape Cod, and to
Scandinavia, Denmark, Holland,
France. ? Mediterranean.
Se ices | cdkese Shetland, Medit.
20-43 a oe 20 Scandinavia, Shetland, Channel,
Medit.
oo |) ce (ee Norway, Denmark, German Ocean,
very rare in Medit.
EE a ee eee Scandinavia, Denmark, Shetland,
Medit. (new to Britain).
SO yan fl awases 20 Greenland, Iceland, Shetland to
France.
| ee 40 20 Iceland, Scandinavia.
30-40 at Eo Ee Northern and Arctic Europe, New-
foundland to Cape Hatteras. ;
ED Se ere eee Scandinavia to Medit.
ME Ce en Scandinavia, Fzroes (new to Britain).
iis... ee vere Scandinavia to Medit.
ES) Oe eee Seandinavia to Medit.
i See eee Northern seas of Europe and America.
30-40 | ...... 40 20 Scandinavia to Madeira and Medit.,
Greenland to Cape Hatteras.
nesses a | eee Spitzbergen to Medit.
ES eee eee Coasts of Northern Europe.
ES ee eee Coasts of Europe.
eee eee Coasts of Northern Europe.

20-40 40 40 20 North Atlantic.
ES | ee rere Northern seas to French coast.
ES Ge Sweden and Norway.
ee ieee Northern Europe to France.
30-40 | ...... oe North Sea to France, American coast,

448 MR. W. E. HOYLE ON THE DEEP-WATER

ARRAN BASIN.

Upper
Brodick |. Hie | | Tale Wigs. Jona
Sroclc® | brennan | marnoch ee sue
Basin. : . Basin.
Basin. Basin.
36. Pleuronectes flesus, DL. ...cccccoss:| sescoe | emeece gl) (UntEE eect)
Hye Cac: 8 06 bg me eRe ne Oh So 20 |) gees ol a
38. microcephalus, Donov. ...| 50-60 | ...... 40500 Fi) cas... ||.
39. platessay. Ts cmt. .cakeekeaneeh | ovens 26. | deem eecees~ || scam
40. Solea variegata (Donov.) .........] sees GD || | aeceee nee
41. Argentina sphyreena;. 27. c..2si5| ° .-05.- . | | acneec.s [rn nn
42. Conger vulgaris, Cuv. ............ senses, | | oedema 56 | |...
42 species.
TuNICcATA.
| 43. Cynthia echinata, Linn. ......... g i 80-104. 0) ancs. | aa
44. Styela grossularia, V. Ben. ...... 50-60 2 80-108 1 fetes 70
AD rustica, ann.” Vacecscaecoes| | eee ee || meeeeeee 80-100 | ...... 10
46. Polycarpa pomaria, S@v. ....c.ca:) | eesseer a) Eoeeeeee omer | ie ee
47. Corella parallelogramma, O. F. G ‘ Ps Bi ears 50
Mill.
48. Ciona intestinalis, Zinn. ........- ? ? eR 508
49. Ascidia mentula, O. F. Miill,...| ...... ? 45°49 TE. 10%
50. -—— scabra, O. F. Mitll.......... ? i 45-104 nest ne
51 virginea, O. Ff’. Miill. ...... 2 2 SO=1G0 V1 Ges
9 species.
Mo.u.vsca.
52, Hledone. cirrosa, Lame. cpatcades| | paaves 22... | 0 Vee ee
53. Sepiola Rondeleti, Leach ......) «2... 22-101) ees
54. Rossia Oweni, Ball ............... 28-90 | 22-40.) ) Seas 30-65
55. Dendronotusarborescens(Miil/.)|...... 40-49 |) ..thee eee Soe
06. Scaphander lignarius (Z.) ......] eee 22... | eelete: HE apameraeee | en
57. Pleurotoma ‘(Bela): ‘turricula |) ...... 1) Gee. 45-49 5b) || 2a
(Mont.) ;
58. Chrysodomus antiquus (Z.) ...| 50-90 | 22-70 | 387-104 | 60-62 50 |
59. Fusus (Sipho) gracilis (Da @.)...) 22 22 100 60-62 | 50:
60. Buccinum wadatum, 2. ifeect.| «0.0 99-70 1042... ete 50-70
61. Aporrhais pes-pelecani (Z.)......| ...... | 92 37-49 60-62 ail
62. Turritella terebra (Z.)............ 50-90 22 37-49 50 ome
63: iissoa GbyssiCOlG | Ores ies.) icccs. |) ceddee 100 ee _
64. Velutina laevigata (Penn.) ......| ...... one 104. TT eee 50
G5. Natica Alderi, Forbes ...cscsesee.] sess: ie &6 | joacsad / Cia é
66. —— Montagui, Forbes....... aut}! OO |) goes 104 60-62 ;
67. —— sordida, Phil. ...........065. 90 22 37-104 d. | 60-62 an
68. Trochus (Gibbula) cinerarius, Z.) ...... | ..... 45-49, 50 30-7
104 d.
G9) ee) i531 oie aa seb ecciwut || sangae, |] setees 45-49 fies
70. ——(Zizyphinus) zizyphinus, Z.| ...... 40-64 | esses 1 eee
71. —— (——) millegranus, Phil. .| ...... 40-49 10d. | 1, cemcates 3
V2. Waanginilla crassa WOW, occwicsel — <asse0, |] aeddee 104.2. | | kese@e-c ik Ga

Loch | Dunoon
iven. | Basin.

f..... 40
: .. 30-40
37 | 32
S. | 40
mm. | 40
e..... | 30-40
40 | 30-40
ee |
...... 30-42
30-44 | 30-40
30-40 | 30-42
30-40 | 30-40
.. 30-40
vee | 80-42
iia | ay
| |...
B... 30-40

rr ay

eeeree

see nee

eee eee

st eeeee

eeeree

eeeeee

eaeeee

een tee

eeeeee

weeeee

ter eee

eeeeee

eeeres

eee eee

eereue

tenons

tenes

eee eee

FAUNA OF THE CLYDE SEA-AREA. 449

General Distribution.

Iceland, Northern Europe to France.

Iceland, Northern Europe to Bay of
Biscay.

Iceland to France ; Kamtschatka.

Iceland to France; rare in Medit.

Britain to France and Medit.

Norway to Medit.

Europe, Medit., East Indies, Japan,
Tasmania.

fi; S3Ae
Greenland, Spitzbergen, to Britain ;
Arctic to Belgium ; U.S.A.(?)
Arctic to Britain; U.S.A.
Seandinavia to Medit.
Scandinavia to Britain.

Arctic to Medit. ; Australia.

Greenland, Iceland, Scandinavia to
Britain; U.S.A.?; Medit.

Scandinavia to Medit.

Scandinavia to Medit.
Greenland to Scandinavia and Medit. ;
W. Africa, Canaries.
Scandinavia to Britain.
Arctic.
All European seas. 1-50 fms.
Arctic and Boreal Europe and U.S.A.
[38-100 fms.
Arctic and Boreal to France.
[5-30 fms.
Boreal Europe to Bay of Biscay, and |,
U.S.A. 5-80 fms.
Celtic and Boreal Europe, Greenland
and U.S.A. Low water to 100 fms.
All European coasts, Medit.
[3-100 fms.
Boreal and Celtic. 7-100 fms.
Scandinavia to Medit. Deep-water.
Arctic and Boreal. Isle of Man.
[Shallow water to 30 fms.
Atlantic and Medit.

Celtic Region. 12-90 fins,
Medit. and Atlantic. 20-60 fms.
Norway to Spain. Shallow water to

[20 fms.
Britain to Medit. 3-25 fms.
Norway to Medit. Low water to 50 fms.
Norway to Medit. 15-1} “ms,

Norway. 20-25 fms.

450 MR. W. E. HOYLE ON THE DEEP-WATER

Brodick

Basin
73. Emarginula reticulata, Sow. ...| ....--
74. Puncturella Noachina (Z.)......| «+++
15. Tectura fulod (MGiIl.) 2... <0.00ses|) ase
76. Chiton marginatus, Penn. ......] see.
77. Dentalium entalis, Z. ............ 90
78. Anomia ephippium, Z. ......... 22-50
79. —— patelliformis, DZ. ..........0.| s-++-
80. Lima elliptica, Jeffr, ........... 50-90
61." Pecten maximus (7..)) <5. .ccpcenc ee
82. opercularis (Z.) ............ 22
83. —— pusio (Z.)........ eakeeaue ens 50-60 d.
84. —— septemradiatus, Mili, ...| 50-90
85. —— striatus, Miill................ 80-90
86. —— tigrinus, Miill...........c0000] ceeeee
87. Modiolamodiolus(Z..) ii ..023-5.| :<a-05
88 Darbata(@7.) stccde. ce sehdeee | aneks
89. Modiolaria marmorata (Forbes)*| ......
90. Nucula nitida, Sow.......:......5- 50-60
5) MUCOUS (HES. di nacpnaetecens 50-60
92. Bulcata, Bron ......c..s0000- 80-90
93 Penis (Monr,)ai.s.-<0acskee

94, Leda minuta (AiU1.) .....0..0505|. 400-0
, f. brevirostris, Jeffr.| ......

95. Astarte compressa (Mont.) ......| ......
96 elliptica, Brown .........46. 50--60
97 suleata (WaC) ics, pace decnaee 50-60 d.

98. Cardium aculeatum, J. .......5.| 20...
echinatmmy 7). ce Soc .ccparalths ve=sse
fasciatum, Mont. ............ een
; WPIMNIRNGN, UI oemancteedal tenses
102.sCyprina islandica((h.)-.: s..c0cc) ae-2 <0
103.,lsocardia Cor (1.) 22.2%. Jescsctecons 50-60
104, Dosinta 6xoleta (Ag) ian caeeccciecns-| ) cose

LOS Fe inet PVE) wees escen | ols

106. Venus fasciata (DaC.) .........06.] seeeee
OFA, PER. Pak. deon ds thon 50-60
108. Cryptodon ferruginosus (Yorbes)) 70

109. —— flexuosus (Mont?.)............| sess.
110. —— croulinensis (Jeff”.)......... 70

111. Solen pellucidus, Penn. .........] .....-
112. Mactra elliptica, Brown .........| ...00+

* This form is. of course, only found along with the Ascidian on which it is parasitic.

ARRAN BASIN.

Kil- Inch-
brennan] marnoch
Basin. Basin.
5 Sa 104 d.
a 104
es. 104
ere 37-104
a 37
40-49 |...
Seni 80
Palsen 45-49
22-64 37-49
GB -:-+4) -Vaaewe
22-70 37-104
eens 37
Seaake 45-49
40-49) Bess
40 45-104
22-70 37-104
eae 80-104
22 45-104
sok “37
70 104
99° eee
Pahies 37
Le 104
65 37
eee | apa
Fees 37, 45-49 d.
40-49 37
eS 37-49
ao | eee

Cumbrae| Fyne.

Basin.

eeeree

eeeece

@eeree

aeseee

eerees
eeocees

eeeeee
eeeeee
eesees
sevens
sancee
seeeee

Upper
Loch |

weeeee

eereee

eeccee

@ecees

eercee

eeeeee

eeonees

eeeeae

eoeeee

eeetee

| wweees

| wees
eeeeer
ereeee
eeeeee

eateee
faeces
eeeeer
estes
tecces

Basin.

eeeeee

eesees

eeeree

eeeeee

seceee

seeeee

eeeeee

aeeeee

eeeeee

eerees

eeeeee

eeetee

FAUNA OF THE CLYDE SEA-AREA.

Loch | Dunoon} Loch
Striven.

Goil.

eeoreee

eeeses

eeseee
eoreee
eercces

eeneee

eeeeee
eeenes
eereee
eorees
eaeeee
sarees
sonore
oe eee
seeeee

eereee

eereee
aeeeee

eereoe

eorees

eoeres

eeecee

eereee

eerene

eoreee

eaeeee

eereee

eestoe

eereee

eoosee

eeeeee

eeossee

oases

eocvee

eoeeee

eeeeee

teeeee

eeeeee

eeneee

All West Europe, Medit.

451

General Distribution.

12-90 fms.

Greenland, North Atlantic, U.S.A.,
and Japan. 20-100 fms.
Norway. 20-80 fms.
Scandinavia, U.S.A., Vigo Bay.
| Shallow.

40-70 fms.
Low water to
[30 fms.
Northern Europe to Medit. 45-50fms.
All European coasts. 15-20 fms.
Norway to Gibraltar and Medit.

Norway to Spain.
All Kuropean seas.

[3-40 fms.

All European seas. 5-100 fms.
All European seas. Low water to
[90 fms.

Seandinavia, Medit. 20-100 fms.

Boreal, very rare in Medit. 12-60 fms,
Atlantic and N. European seas.
[12-60 fms.
Low water to 70 fms.
Lusitanian Region, Medit. 3-10 fms.
All European seas, Parasitic on As-
cidia mentula. Low water to 40 fins.
Sweden, Lusitania, Medit. Shallow
[ water to 34 fms.
Common.
[7-90 fms.
Scandinavia to Medit. (deep).
[30-100 fms.
Northern Europe, and U.S.A.
[40-100 fms.
Arctic and Scandinavia. 10-100 fms.

N. Atlantic.

All European seas,

Norway. 7-70 fms.
Greenland, Norway. 10-45 fms.
All European seas. 8-80 fms.

Scandinavia to Medit.
Scandinavia to N. Atlantic. 7—80 fms.
Atlantic.
Arctic and Norway.
Northern Europe.
Scandinavia to Medit,
All European seas.

[Low water to 80 fms.
All European seas.

[ Low water to 60 fms.
Norway to Medit. 4—60 fms.
Norway to Medit. 3-100 fms.
Arctic to Medit. 20-100 fms.
Arctic to Medit. 3-80 fms.
Arctic, Atlantic, and Medit.
Throughout European seas. 6-100 fms,
Arctic and N. Atlantic.

[Low water to 50 fms,

30-70 fms.
5-80 fms.
15-40 fms.

452 MR. W. E. HOYLE ON THE DEEP-WATER

ARRAN Basin,
a lll vm
j Kil- Inch-
yo brennan | marnoch ee Fyn z
* | Basin. Basin. — ‘.
113. Mya ftroncata, Ly ...ciccck cece] scenes] Coen ot ae ee 50 da :
114. Corbula gibba, Olivi ............ 50-60 22) |) “Sirens 50 36
115, Saxieava rugosa (L.) ...:0..00.0. roo
116. "Béllina bordida.ssa..scheented te ce | a Saas al
117. Semele (Abra) alba (Wood)...... $0.41) "See 37-80 50 36-50.
118.'=—— nitida, Mal... «...00stecenesh Satan 2) eee i en
119, ——— tenuis (Mont) v..c.ccccctse| tee gc] ae sce) fee a 4
120. Cuspidaria abbreviata, Forbes...) 1.0... 0 | cesses 36° ask) | a
121 cuspidata (O02) 5. Aaa pene naar Ba Ss a
70 species.
Bracuiopopa.
122. Terebratulina caput-serpentis| ........] ..... 80-108 |) Wn. 50
(i,)).
1 species.
Pouyzoa.
123. Scrupocellaria reptans (L.)......] ...... 64....| desea. | eae 1)
124. Bugula turbinata, Alder.........) ...... 30-50") - 9.5.88" Sai ctr
120) Cellatia fistulosa CZ.) ...0c05.<-00| fences 25-64 1 Sedeos, Qi eae, 9 ee
126, Wlustratoliaeea (1) Gracey 2.. 28 25-49 | wate | | Ee
127 securifrons (Pallas) ......) ....0 63-4) ale ee)
128. Membranipora catenularia | ...... 50. +} caktene ORT Mpeemei oS eee
(Jameson).
129, ——- Flemingi, Busk ............| ...00s se ee
130: ——« pilosa: (2. Wes. GAR eel oa: 50 104 bee nee ap
131. Microporella ciliata (Pall.)......) 0 ...... 50 - +] lsewee 8] CeeeeeN he ea
132. impressa. (Ava.)*its Geeta) 5.2.3: 50): +) ansiied | eee a
133. Schizoporella unicornis(Johnst.)| ...... i re | ee
134. Hippothoa carinata, Norman...| ....+ BO} | scat eee on ee
135. Porella compressa (Sow.) ......| se... 25 - | seweder® 1) paeeeeee) ee
136. Smittia reticulata (MacGill.)...) 0... 25 | <Basseee eal peeeeeee a ee
137. Mucronella Peachii (Johnst.)...| ...... 25-64 | ugsnsce 7
138 ventricosa (f7@98,) ).f.02525| ...00 50 | see eee eee
139. Cellepora avicularis, Hincks ...| ...... 50-65 | ue 5 1 eee le
140); ———— ‘pamiicosay wD. alii ees] 4. 50. «1? Qokd a of
141 PAMUNGEAI. fe dehie SR pacowe 25-64 | | .cacsze® |) eee

Basin.

eeetee

eeteee

eeeece

eet eee

eeeeee

eee eoe

eeeeee

ee eeee

eee eee

FAUNA OF THE CLYDE SEA-AREA.

30-35 | 30
35-45 | 30°
2 2 ae

ease

teeeee

eeneee

weet

eeeree

eee eee

ween

453

General Distribution.

—_———

Arctic and Atlantic.
[Shallow water to 34 fms.
All European seas. 7 to 80 fms.
Boreal and Celtic regions, N. Spain.
[Shallow.
Arctic. 53 fins.
Scandinavia to Medit.
[Common 1-49 fms.
Norway to Medit.
Britain to Medit.
Norway, Medit. 40-200 fms.
Norway, Sweden, Medit. 12-185 fms.

All European seas, U.S.A. 10-90 fins.

Scandinavia to Medit., Red Sea (?).
[To 100 fms.
British only. Shallow.
Seandinavia to Medit., U.S.A., Ma-
deira, Indian Ocean, N. Zealand.
[40-140 fms.
Norway to Medit., S. Africa, China,
Pacific. To 70 fms.
Spitzbergen to Medit., Labrador.
[10-300 fms.
North Sea to Medit., U.S.A.
[40-300 fms.
Greenland to Medit.
[‘‘ Tide-marks to deep water.’
Greenland and Norway to Medit.,
U.S.A., Indian Ocean, N. Zealand.
[Low water to 100 fms.
Cosmopolitan. 0-145 fms.
Norway to Medit. 30-40 fms.
Greenland to Medit., U.S.A., 8S. Africa.
Antrim, Birterbuy Bay. [80-170 fms.
Arctic to N. France. 40-170 fms.
Norway to Medit., Falkland Is., N.
Zealand. 40-80 fms.
Greenland and Norway to Medit.,
U.8.A. Low water to 170 fms.
Arctic, Norway to Medit., N. Zealand.
[10-20 fms.
Spitzbergen, Scandinavia, Medit.,
U.S.A.

Cosmopolitan. 5-50 fms.
Norway to N. France, Madeira.
[8-170 fms.

454 MR. W. E. HOYLE ON THE DEEP-WATER

ARRAN BASIN.

vee
ae, och
Brodick | ), a ies Cumbrae| Fyne.
Basin. ern marnoc Basin.
asin. Basin.

—— ——, var. producta.........) seseee 29 || Kae eeckees =|
144. Stomatopora granulata (M.-| ...... I ee

Edw.).

145. Idmonea serpens (Z.) ............| seseee 65...4 \igeeeee gy Seaee-e, | Goa
146. Diastopora obelia, Johnst. ......) sseeee 50-64 | | Geyer: | ce
147. Lichenopora hispida (FVem.) ...| ....+. 25-64 || | cuteness eeweeees

—— ,var.mexandrina,Peach| ...... 65 i Mie tn
148. —— verrucaria (O. abr.) ......| see. 5O |} nice React
149. Mucronella ventricosa ............| sseee. 5O 2 cpscchomeneeeeceee © 1! > co
150. Vesicularia spinosa (L.) .........] s-.00. 50 9.) eRe eee 7! oe
151. Cylindreecium dilatatum, Hincks.| ...... 65 | gee ieee Ge

29 species.
CRUSTACEA.
152. Inachus dorsettensis, Penn. ...} ...20. | seeeee 37-104 50-60 50 |
153. dorynchus, Leach c.s2.000| (esas 25 |) | Seeeee, OM aReneee = een
154) Miyas aranens, L.,...0cccceccensess| 0sere, | Besneeanl see a 5074
155 coarctatus, Leach ......... 60 49-65 45-49 60 75-80!
156. Stenorhynchus longirostris, M.-| ...... 40-49 37 BO
Edw.

Li MOSLTAUUG, Hrante, cteh oedema pecsae 40-64 | | weteds > Bees 4
158; Hurynome aspera, Pens..ccc.5.-| e<<s || Sewe-nm | eee BO 4. 3c
159. Portunus depurator, L. ......+. 80-90 ? 104 50-60 |...
160 noleais, LA07 ccc: .socssoscel. essaetall Meee ‘ie in a re Me
161 marmoreus, Leagch..ccceus| veces. ;.| ‘xeesen, sill needa ern '
162. pusillus, each... asencsln econ, al), Peete i eo
163. Ebalia tuberosa; Penn siccschaseedl) Seeevas cdl . eeeeecreal) ee 50 1) ae
1G64-dithodes: malas.) eeicesctet ed] Greece ? 37-49 | ances. 708
165. Eupagurus bernhardus (Z.) ...] ...... 20-49, 80} 37-104 | 50-60 | 35-70!
166. Prideauxii (Leach) .........) ve... 50 dl fae 50-60 |...
167. —— pubescens (Kroyer).........] sees 20-65 4 1oeean 60 | Wa .
168 CxOavatus, MMecrseceess|! eces,..| weeses 45-49 |) eceess rill ance
169. Anapagurus leevis (Thompson) | ...... ? 37-104 GO} 5 ase d
170. Galathea dispersa, Sp. Bate...) 50-90 | .s0.0s | | sanatan) een
17] nexa, Himbleton .......0000 BO-60 | ssee6,,. 4) 7 eeqeeee anne
iVOz squamifera, Leach .........| ss... 49 | | -gctoae a en 80
173. Munida rugosa (Fabr.) ..... ... 80-90 70 ee (mae
174. Calocaris Macandrew, Bell ...... 50-90) Wl... teste 80-104 BOW! | cae
175. Nephrops norvegicus (Z.) ...... 80-90 40 Bl. e bo Bee eae
176, Crangon Allmani, Kinahan ...| 25-90 | 20-70 | 37-80, 105| 40-62 | 50-75
ree echimaulatus) 2Rars clive) cccsck.s:| (aces CC ee
178. BiIGHOB: TGC00Is i coeesee onl Vasedae. | \ PROGR 45-49 | cece | one
179, Mika edulis; 721980. ccvesestssinee DS. | Pea eee LOL. | ‘eveae “9 > Soe
180. Hippolyte Gaimardi, M.-Hdw. .| ....... | seve 45-104 | ...... 50-75
181. pusiola, Kroyer isiccsescened| ..00..,| s0esoe,. | emer na nr |
182. —— securifrons, Norman ...... 50-90 | 50-60 37-80 60-70 | 35-80
183.

spinus, Sowd, ....... Sreees sal shi sevice 65 | newaen eee |

FAUNA OF THE CLYDE SEA-AREA. 455

Upper
Loch | Dunoon! Loch PP Gare- eee a

CE eee eee eee Arctic to Medit., U.S.A., Madeira,
S. Africa. [ 10-100 fms.
EE OE eee ee ee All British shores. 4-96 fms.
EE ee ee ee eer Nova Zembla, Scandinavia,
ED Aa eee ee ... | Norway to N. France.
[Low water to 170 fms.
ES ED eee ee oe eee Norway to Medit.
|2 fms. to “‘ deep water.”
I ee Arctic and Norway to Medit., U.S.A,
[2-20 fms.
EES eee ee ree Greenland and Norway to S.W.
France. To 170 fins.
re eee ee Shetland. 80-100 fms.
es ee Arctic, Norway, U.S.A. 10-150 fms.
Ee en ree Arctic, Scandinavia, to Medit., New
Zealand.
cece eee ee Norway to N. France.
a A) ne ee Belgium to Medit. 8-? fms.

SR eeerreee Soo oe Norway.
ES eres wade. ie site ...... | Norway, Medit. 25-35 fms.
: =e 30-42 30 30 20 Norway, Medit.
2 ba a ree eth Norway, Medit.
ES Pee ee eee 20 N. Atlantic to Medit.
ess . ene ahs Me rr eee Norway, Medit., Canaries.
ES eee eee Norway, Medit.
SD eee 40-45 30 ...... | Norway, U.S.A. Shallow.
40 380-42 | 30-40 30 20 Norway, Medit., U.S.A.
-- 30-40 datas ETE Wdaane Norway, Medit., Cape Verde.
I PE sccais., | codsces | ceanee Norway, Labrador, Arctic, U.S.A.
| Ee eee eee neous Medit., Atlantic, Senegambia.
eee eee eee Norway, Medit.
SE dikes f° vaseos | seccen | ceeeee Norway.
a 4 20-40 hobs seats Sa Norway, Medit.
ES eee eccees ...ee. | Norway, Medit.
40 B40 | SO-40 | wn, | cece Norway, Medit.
ee oan 40 Bae FAT e aiteck Norway, Medit., U.S.A.
40 30-42 2 ba ee 20 Norway, Medit.
40 30-42 | 30-45 30 20 Norway.
SS eee er Norway.
i) coe SS ee Meee Norway, Medit.
| = eS ae cee Norway, Medit.
40 380-42 | 30-35 30 20 Norway, Labrador, Arctic, U.S.A.
i eee 20-35 | 20-35 | ...... Norway, U.S.A.
40 30-42 | 30-45 | ...... 20 Norway, U.S.A.
Ee ee eee Norway, Labrador, Arctic, U S.A.

. Pasiphea sivado (Risso)
. Nyctiphanes norvegica (JM. Sars)
9. Boreophausia inermis(Kréyer)?

3. Gammarus locusta (L.)
. Meera Loveni (Lruz.)

_ Amathilla homari (Fabr.)
. Ampelisca macrocephala, Lid/).

. Kucheta norvegica, Boeck
. Pseudocalanus elongatus, Boeck
.Temora longicornis, Miil/. ......
. Centropages hamatus, Zi//).

. Dias longiremis, Lillj. ............
. Oithona spinifrons, Boeck
. Eetinosoma atlanticum (Br. &

. Secalpellum vulgare, Leach
34, Balanus hameri (As¢.)............

MR. W. E. HOYLE ON THE DEEP-WATER

———

4. Caridion Gordoni (Sp. Bate)...
. Pandalus annulicornis, Leach ...

brevirostris, Rathke

eeeeee

sete ewes

Raschi (M. Sars)

. Mysidopsis didelphys (Norman)
2. Mysis neglecta, G. O. S
. Cirolana hirtipes, 1/.-Hdw.......
. Conilera cylindracea (Movnt.) ...
. Munna whiteana, Sp. B. & W..

. Janira maculosa, Leach

eee eet ees

. Arcturus longicornis (Sowd.) ...
. Idotea parallela, Sp. B. g& W,...
9. Hippomedon Holbolli (Kréyer)
. Tryphosa longipes (Sp. Bate)...
. Callisoma crenatum (Sp. Bate)
. Bathyporeia pilosa (Lindstr.) ...
. Lysianax tumida (Kroyer)
. Leucothoé spinicarpa (Adi/d.)...
5. Stenothoé monoculoides (Monz.)
. Harpinia plumosa (Kréyer) ...
. Westwoodilla ceecula, Sp. Bate

. Monoculodes

eevee

Stimpsoni, Sp.
Bate.

longimanus, Sp. B. & W..

. Epimeria cornigera (J. C. Fabr.)
. Cheirocratus Sundevalli( Path.)

assimilis (Zilijeb.) ......4..

eonteveee

longimana (Leach)

eeeeecees

tenuicornis, Lz//7.

eee weee

. Haploops tubicola, Lillj. .........

setosa, Boeck

Podoceropsis Sophia, Boeck .

undata, Sp.Bate .......06:0.

3. Cerapus abditus, Zemplet. ......
. Evadne Nordmanni, Lov.
. Calanus finmarchicus, Gunner .

Rob.).

83 species.

Brodick
Basin.

eeeeee

eee ees

eesees
seeeee
ereeee
eeetes
eeceee
eectes
seeeee
seceee
sev cee
sence
serves
eeseee

eeecee
ee seee
eevee
eectee
woeree

eeeeee

seeeee
eevee
eereoe
e@ecese
eeveos
sence
eet awe

eoeree

eereee

Hee eee

ARRAN BASIN,

Kil-
brennan
Basin.

eeeeee
@ercees
eeceee
eeesee
eeceee
eeeeee
eectos
eecees

eecvee
feetee
eevee
eereee
eoocee
ee eee
seewee
eee eee

seesee

eeree
eereee
eececee
eeenee
seeees

seccee
eevee
eereee
eereee
eeeeee
teceee
eeeeee

eee eee

seen

Inch-
marnoch
Basin.

eoesee
eectes

eeceee
eorare
eee tee

eeewce
eecnces

eaves

fereee

eeeeee

ee eene

ee eeee

Cumbrae
Basin.

eereee
eeeeee
eeeeoe

eer eee

eeceees
eoorees
ee eeee

@eeees

eercee

rr
ences
eereee
eoeeee

eoeeee

weeeeee

eeteee
eeseee
eeerse
eeceee
te teee
vecers

ee eee

eeeeee

eereee

eeeeee

eereoe

oeseee
@erses
eoreee
eooeee
eeosees
eesees
ey
eeceee
eeeeee
ooseee
ceceee
eeseee

ween

see eee

FAUNA OF THE CLYDE SEA-AREA. 457

Upper
Dunoon Loch Loch Gare- General Distribution.
Basin. Goil. loch.

Long.

—

eee a Norway, U.S.A.

30-40 | 380-45 30 20 Norway.

SS ee ree 20 Norway, Arctic, Medit.
eee ene Norway, Medit.

it OOD veces | cannes Norway, U.S.A.

| a ae Salt ae Norway, U.S.A.

40-45 | ...... SM ck. Norway.

eee | see, |) ccaltiens Norway.
ee ir Norway.

i a bon IE A Medit.
eee | wt wet. | ccenne Firth of Clyde to S. of Britain.
ee ey ae ee Firth of Clyde to 8. of Britain.
ER Norway.

Bes... i ee Norway.
eee er Medit.
ES | ee eee ee Norway, Medit.
ED) ree reer senses. | Norway.
EE ee ee eee ee Norway.

ES eee eee eee Norway.

aayass ee Fy Rae wees. | Norway to Medit.*

RE Pe eee eee Norway.
ete ec 0 ewccan. |. eevee Norway.

a | | ae ee Moray Firth, Plymouth.

Ee NS ee ae | Plymouth.

Bee Co 2 ea | S. Norway to S.W. France.
| 2s SG eel aacecs. | see. Norway, Arctic, Medit.
| ree ee Norway to France.
RU wpeper, | ceacen.. | coveee Norway to France.
ee ee Norway, Labrador, Arctic, Medit.
I eee ee Greenland, Spitzbergen to Denmark.
NS || SS ee ee Norway.
ee ee ee Norway, Arctic.
DS eee ae Norway, Labrador.

RE eee ses aug (ee Norway.

z AEN sigivay,) | | <rncae, | veseee Norway, Arctic.

eS A eee De Norway, Arctic, U.S.A.
Bi... 39 ste nerere onexs | DOTWay,
Ee ee Northumberland,
er cesenc, | dense | senece Medit., Atlantic.

ee SS ee re North Sea, Medit.
20-42 35 40 20 Arctic, Norway, N. Atlantic, Southern
Seas, Medit.

EE aye eee Scandinavia.
20-42 35 40 20 Scandinavia.
Bs. VE ae ere 40 20 Norway.
iis. 20-42 | ...... Cg Dee North Sea, Medit.

3 20-42 35 40 20 Norway, North Sea, Medit.
> ee 50 20 Norway.

| a RES sei S| «dete N. Atlantic.

SE Se ee | European seas, Medit.
ee ee Iceland, Norway, Feeroes, U.S.A.

Carus (Prodr. faun. Medit. p. 409) states that this is confined to the Mediterranean,

r
ahd

458 MR. W. E. HOYLE ON THE DEEP-WATER
ARRAN Basin.
Upper.
: Loe
ms Kil- Inch- |
Brodick brea Eee Cumbrae| Fyne.
Basin. : . Basin.
Basin. Basin.
VERMES,
235. Pontobdella muricata,'\ ZL. 2252:] 20.05 | .cesens 0a Oe . oo
236. Aphrodite aculeata, DF 220.500] Sscstec s+] Baneee STO 7 vanes 65-75
237. Hyalinecia. tubicola (O. F| 2.5... | secon | See |
Mill.)
238. Eumenia Jeffreysi, M‘Z. ......... 80-90 40 104s ya Agee. 65-75
239. Pectinaria belgica (Pad/.) ...... 80-90 | ...... (i. ee
240. Sabella pavonia (Sav.) ......... SO). cal! eeceee es
241. Filigrana implexa (Berf.) ...... 80-90 | 22-70 Os Ml” aekiens 50
242. Serpula vermicolaris (L..) 2.00: .c0see |: | (asaeomal|” |) ene es 50
243. Leptoplana tromellaris: ......00-| ss... | seaeee 80
9 species.
EcHINODERMATA,
244. Cucumaria Hyndmanni | ...... 78-80.) axecace” Re” |)
( Thomps.)
245. Psolus phantapus (Strussenf.)...) ss... | seeeee HO... W Seinen” | >
246. Thyone fusus (O. Ff. Miill.)......| ....0- 75-80 sien DAN Rares | - So
247. Holothuria intestinalis, Asc. g| ...... 50 |. Mvgsstee 0 qt) tenes ot) (een
Rath.
246 Pichinus esculentus: 24.52.84 082] © ss soen7, ml aeons S1-60 faeces 30-80,
249 miliaris, P. 0.08. Mies.) .c.000 5+) cameos |) een
250. Brissopsis lyrifera (Forbes) ...... 50-90 | { 22-40,| 37-104 50. | cam
75-80
251. Spatangus purpureus,O.F Mii.) ....0...| sesso | Pe emmaes 50 36
952. Echinocardium flavescens, O.F".| ...:.. | csosese | ovens een) een
Miill.
253. Porania pulvillus (O. F. Miill.)| ...... 22-45 Oi > Weekes 0) © ee
254. Stichaster roseus (O. F’. Miill. ). ee 22. 923) eae 20-80 |...
255. Crossaster papposus (Linck) ...| ...... 22 abe ree eaters - oe
256. Solaster endeca, (G77t.) ...cccsoive.| cescoas | aensen pt || | pues
257. Cribrellaioculata (77eie)) rasa.<| <<<... 0] secume re oe ee '
258. Asterias rubens/iuastececkeeeea| sess 22-40 40-80. 4. Sesnes (> . ae
259. wiolaces, OREGUMUN. Wass | <caves 22. . |. \deengee alee eee
260. Ophioglypha affinis (L7/.) ......| sees. 75-80 104. | wsweee
261. albida (Forbes). .ccc..ccscse:| o-csee | seveee| |) SupRemEn 50-75
262 lacertosa (lenge) aecskeee| lees .ke ||! Bec SO) Agee 50
263. Ophiopholisaculeata(O.F.Miill.)| ...... 0 | sees 37-104. | Seen ‘ 50
264, Amphiura Chiajei, Forbes ...... 50-90 | {32-4°>| 25, 37-104| 20-60 | 36-50
265 filiformis (O. F. Mill.) ...;. 50-60 | 75-80 100 35-60 36
266. Ophiocoma nigra (0. F. Miill.). wooses | leieweae | | wiser nai
267. Ophiothrix Pentsphyilum 4 aceon. .| Gdawes 37 50 50-75
(Penn.)
268. Antedon rosacea (Limck)......00| seeeee | cereee | neneee | caeene | ceeees

25 species.

Loch | Dunoon
Striven. | Basin.

eereee =| eer eee

Cs
aeseee | 8 eeeee

eeseee | terres

Ce
seeeee =| we eeees

sercee =| 8 eeu ee

eeceee =| tee eee

eee ee | ae eeee
saaeeo06md06 6 OO Cee

eersee | eeeeee

FAUNA OF THE CLYDE SEA-AREA.

eee eee

eoeeee

toeree

eeeeee

eereee

ereees

weeeee

eeeere

eeneee

eoeees

eeeeee

Peeves

eeeree

eeeeee

eeeres

eens

teens

eereee

see eee

eeeeee

LINN. JOURN.—ZOOLOGY, VOL. XX.

459

General Distribution.

North Sea, Medit.

Scandinavia to Medit., U.S.A.

Scandinavia to Medit., Madeira.
[80-80 fms.

British seas.

Scandinavia to Britain.

Scandinavia, Britain. 20-100 fms.

Scandinavia to Medit. 20-300 fms.

Scandinavia to France. 15-80 fms.

Scandinavia to Medit.

Arctic, Scandinavia, U.S.A.
Scandinavia to Medit.
Arctic, Scandinavia.

Norway to English Channel.

Greenland and Norway to Medit., W.
Indies, Florida, Cape of Good

Hope. 0—-2435 fms.
Norway to Medit., Azores, Bermuda,
W. Indies. 0-45 fms.

Norway to France, Cape of Good
Hope, Carolina to Florida.

[0-150 fms.
Seandinavia. 15-106 fms.
Scandinavia. 2-50 fms.

Arctic and Norway to France, U.S.A.

[0-640 fms.

Arctic and Norway to France, U.S.A.

[0-150 fms.

Arctic to Britain, U.S.A. O-1350 fms.

Only British. 0-53 fms,

Norway to Britain. 65 fms.
Scandinavia to Medit., U.S.A.

[To 192 fms.

Scandinavia and Feroes to Azores,

Medit. 5-458 fms.

Arctic North Atlantic, Medit.,

Madeira.
Arctic and Scandinavia. To 560 fms.

Scandinavia to Medit. To 555fms.

Scandinavia to Medit. To 555 fms.
Arctic and Scandinavia. 7-87 fms.
France. &3 fms.

Hebrides to Madeira and Medit.
[100 fms.

4.60

. Lafoéa dumosa (FVem.)

MR. W. E. HOYLE ON THE DEEP-WATER

Ca@LENTERATA,

Hydractinia echinata (Flem.) ...

. Perigonimus repens (Wright) ...
. Dicoryne conferta (Alder)

Tubularia indivisa, Z.............

Campanularia angulata, Hincks,
volubilis (Z.)

eeceeeteeeeeres

fruticosa (Sars)

. Calycella fastigiata (Alder)
. Halecium Beanii, Johnst.

halecinum (27)ccassesnaeacee

muricatum (Il. § Sol.)...
Sertularella fusiformis, Hincks .
Gayl (Tiirin.)) feces oc
rugosa (L.)

. Diphasia attenuata, Hincks
. -—— fallax (Johnst.)

tamarisca (L.) teuiiar
pinaster (Hl. & Sol.) ......
Sertularia abietina, D.............

argentea, Hil. g& Sol. ......
Hydrallmania falcata (Z.)

. Antennularia ramosa, Lamh. ...
. Aglaophenia tubulifera, Hincks.
. Plumularia Catharina, Johnst. .

pinnate (Ein a.
Bolocera tuedize (Johnst.) ......
Virgularia mirabilis (0.7. Mill.)
Pennatula phosphorea, Z. ......
29 species.

PoRIFERA.

Suberites ficus (Johnst.)
suberea (Mont.) ?
Tragosia infundibuliformis
(SOGRHY To... Hes isang deg eos
Gianlina, BP... sagpscen darter esse

. Myxilla incrustans (Johnst.) ...
. Iophon Pattersoni (Lwk.)

feetes

Grantia ciliata, lem. ............
7 species.

ARRAN Basin,

Kil-
brennan
Basin.

Inch-
marnoch
Basin.

Brodick
Basin.

eeeres
@eevcee

eeceee

eec50ace

eocees | Terr | eaeees
eoeres
eoccee | “Ee | eevee
eovoeee | OW £j|{ esse
eeceos | eovcce
eeoccoe | jE | eee
eoeres [| eeeeee
cesses 9] PRISTON | wens
eocsne fF | WY I | \sisteeew
eooeee | CUTIE | = eeccee
coceme ~— WE "i | ‘ateinisien
neceme °F ME i | “semis
eepeese Pf | Wx 1) wena we
eevces

eoceceee | = dmddmd ENF | jeeeccve

Bebcae

eovecce

eeecee | jseevee | ( ef@8e08

eooeee Ff OI UtF | —— aeeeee
eereee
serece

feces | e80008

eeoecee

eeceee

eeseee

eeores

eoeeee

eeeree

eeotes

eestes

weseee
eoeeee

@eosvece

eervee *

@eoecse

eoctee

eevee |

eeeeee

Loch | Dunoon
| Striven.

teeees
seeeee
eeeeee

Basin.

ferees
serene
eostes

serene
tereee
seeeee
eeeees

eereee

eeeeee

eee

eeeeee

Loch
Goil.

eeteee
eeeees
eeeeee

eet eee

weeecee
eres
s@eeee

eaeeee

eee eee
eeeeee

weosees

seetes

seeeee

teeeee

eeeeee

tereee

FAUNA OF THE CLYDE SEA-AREA.

eeetee

eerees

eeeeee

General Distribution.

eeFeaor_r_—XS— OS _

France, U.S.A., Medit. ?

Medit.

Shetland, Northumberland.

Greenland and Norway to Bay of
Biscay, Medit.

N. Ireland to Channel Is.

Iceland, Norway, U.S.A., Medit.

[20-100 fms.
Norway, U.S.A., Medit, Tide-marks
[to 145 fms.
Iceland, Norway. 15-100 fms.
Shetland to Cornwall.
Medit.
Greenland and Norway to Medit.,
U.S.A. 80-50 fms.
Iceland, U.S.A. 30-50 fms.
Medit.
Normandy. 60 fms.
Greenland, Norway, and Labrador.
[30 fms.

Port Adelaide, Medit.

Norway, U.S.A. 30 fms.
Bay of Biscay, U.S.A., Medit.

North Sea, Medit. 40-140 fms.
Greenland and Norway to Medit.,

U.S.A. 30 fms,
Greenland and Norway to Medit.,
U.S.A. 4-50 fms.

Belgium, U.S.A., 8. Africa. 35 fms.
8. Africa.

Algoa Bay. 15-30 fms.
All British coasts, 40-60 fms.
North Sea, Medit.

Scandinavia, U.S.A.

Norway, Scotland.

European seas.

Hebrides, Northumberland, Mayo.
British coasts.

Shetland to Channel Is.
British seas.

Shetland, Patagonia,Tristan da Cunha.
British coasts.

462 MR. W. E. HOYLE ON THE DEEP-WATER

DISCUSSION OF THE RESULTS.

The above Table may be summarized in the more condensed
one given below, in which the number of species of each group
of animals from each basin is shown.

ARRAN Basin. 3 ap
2 8
a Fy a = bs je
S| |
se/ee|se/22/ 8 |F4/2@/slal4)e
os i S g 3 a3 io) 5 aq ro) 3 i es}
AQ) SA | oA | FA] A a|/51e6 a, | 0
~ ig Ss) oO ac wa boe b
5 =) A =
Piseer stig he 8 cc 10 29 12 7 36 61 6120 beat. 7
Tumicata °%..:.2..:. if ; 6 : 6 5 2 | eS 1
Mollusea ......... 24 26 46 16 61 20°} 1872134 i f
Brachiopoda ...... de ae 1 Ack 1 1
Polyzoa,323.6. 0. cor 3l 1 ee 3l
Crustacea ......... 24 29 46 34 75 20.) 16 1-31) a) a6.
Wermes hose % 6 3 6 Ba 9 4 1 41D 1 1
Echinodermata... 3 12 12 6 18 8 5 |. 6 6 3
Coelenterata ...... ai 26 5 1 27 +] 1 2 2 1
(PORMBER «cae scee ss ea ff 4 bee 8 rea Na | SAA
Motels: vas cucees 68 163 139 64 | 272 69 | 42 | 93 | 54] 39 | 33

In dealing with these figures great caution must be observed,
and it must always be borne in mind that no locality can ever
be said to be really exhausted. The number of dredgings upon
which the present inquiry is based will only suffice as a basis
for very general conclusions. This has been abundantly evident
during the progress of the work, for tables like the above have
been drawn up several times, and it has been noticed that each
successive addition of new data has increased the likeness of the
faunas of the different basius to each other.

Certain sources of error must also be avoided ; for instance, it
was apparent, from an examination of the various lists, that the
Polyzoa and Hydrozoa had only been exhaustively examined in

FAUNA OF THE CLYDE SEA-AREA. 463

the case of the Kilbrennan Basin. These groups must there-
fore be left out of account in comparing the different basins
with each other. Furthermore, the Worms and Sponges have
been very incompletely studied. Many specimens I was only
able to refer to their generic position, and hence it seems advi-
sable to omit these groups also from consideration for the
present.

Deducting the figures corresponding to them, we have the
following modified list of the total numbers of species from each
basin :—

ieromice Hasm:. .... 0. ee 62
fatpecninam basin «2.0.0. eS. 96
Imchmarnoch Basin ............ 1238
Rimipede basin -.......c......8- 6:

Total from Arran Basin ...... 197
Peper lipeln WyHe. 5.6. ccces ww oe os 60
Meee ow oe ee ks 40
Or ABI. Sse ee ee ee 87
C2 LS 44,
mieper Woek Long......<....... 36
ee 32

This revised series of totals proves beyond all doubt that the
richest fauna is in those basins which are in closest proximity to
the sea, and that it diminishes as we proceed into the more land-
locked portions of the district. An exception, which is, however,
more apparent than real, will be noticed in the fact that a larger
number of species has been found in the Inchmarnoch Basin
than in either the Brodick or Kilbrennan Basins. This is, I
believe, to be explained partly by the fact that the Inchmarnoch
Basin is much larger than either of the others, and descends to
a greater depth, and partly by the circumstance that more
dredgings have been carried out in it. If we take the Arran
Basin as a whole the truth of the above proposition is obvious. It
is, of course, just what might have been anticipated beforehand in
view of the marine origin of the whole fauna, but it is satisfactory
to have the matter established by actual investigation.

1t will be of some interest to consider the relationships of this
fauna as a whole, and particularly to ascertain which of the
neighbouring faunas it most closely resembles.

4.64. MR. W. E. HOYLE ON THE DEEP-WATER

For this purpose use must be made of the distributional notes
appended to each species in the list. According to this infor-
mation the species fall into three categories. The first contains
those which range from Scandinavia to the Mediterranean, or
even more widely still; these may be termed, for the present
purpose, “ Wide-spread ” species. The second consists of those
forms which are common to the Arctic and Scandinavian waters,
and hence may be termed “ Northern ;’’ whilst the third is made
up of species which may be called “ Southern,” as they extend
to the Mediterranean or the African coast.

The following Table shows the numbers of species of the
various classes of animals which belong to each of these cate-
gories :—

Wide-spread | Northern | Southern

Species. Species. Species.
PISGAH eA. oh t ees aee 23 17 2
DGGCALR sc deacecssloss ot 4 5
Mollusca and
Brachiopoda ... } = = 2
POly7ZGan poe. case tae 20 5 1
Crustacea ....>....... 382 41 5
WOTMIOS acca eciiess 3 3
Echinodermata...... 10 13 2
Ceelenterata ......... 9 4 9
Motalery.0.5:ar 147 110 22

From these figures it appears that the major part of the fauna
is composed of species which are dispersed more or less widely
over the north temperate regions of the globe, whilst the smaller
half is very unequally divided between the northern and southern
species, the former being five times as numerous as the latter.
In only one division of animals (the Ceelenterata) do the southern
forms predominate over the northern, and this subkingdom has
been hitherto very inadequately investigated in the Clyde area.

The depth of 20 fathoms as limiting what might be considered
the deep-water fauna in the Clyde sea-area was selected not
from any preconceived idea as to its significance, but because it
was convenient for practical purposes, and because it was appli-
cable to all the lochs, the extreme depth of the Gareloch, which

FAUNA OF THE CLYDE SEA-AREA. 465

is the shallowest, being about 23 fathoms. In the case of some
of the other basins, however, if is so far from marking out their
limits that it does not touch the tops of the ridges which sepa-
rate them. To Dr. Murray I owe the suggestion that it might
be worth while to compare the faunas of the deepest parts of
the basins, taking some other contour-line as the upper limit.
I have therefore gone over the Table given above and selected
from it those species which are found in proximity to the bottom
of each basin. The depressions themselves vary so much in
depth that I have thought it advisable to record both the forms
which are found below the 50-fathom ‘line and (in a separate
column) those which are found w..hin, say, 5 to 20 fathoms
of the bottom of each basin. Furthermore I have excluded
those free-swimming forms whose distribution in regard to depth
cannot be regarded as absolutely fixed. On this ground I have
omitted the Fishes, Cephalopoda, Amphipoda, and part of the
Macrurous Crustacea (viz. the genera Hippolyte, Pandalus, and
their allies) ; the Polyzoa, Hydroida, and Sponges have also been
neglected because they have been insufficiently studied.

In the subjoined Table the followmg symbols have been
used :—

*—occurring at the depth mentioned in the head of the
column.

t=occurring in the locality, but at a depth less than those
under consideration in the table.

W = Widely-spread species.

N = Northern species.

S = Southern species.

MR. W. E. HOYLE ON THE DEEP-WATER

466

AA vee e)ain) ee
NT ar coe * ba
ar dos oe % %
eM * * % *
"NI x * * *
= a “ Pi eee
Nr are eee % *
"mM. eee i ome a
“AA ele is eee 28
NT vee % * we
“M eee aai6) te oe
“NI eee % 4 a
“M. eee % * he
NI eee eee oe ath
a % 5 oC

“Mg coe ece eee 238

‘moTynq
“EVIC) sguory

qooy
soddq,

Top | “Useg | “Weal
yooT | uwoound} yooTy

| | | |

hi ie 56
ae oe %
AAR eee *
f ra oon
ern x *
eee x *
cee SAO %
& x One
eee * 22
eee * oe.
% * :
% * :

le eke 6D
04 04 o

G9 0¢ Og

UISUT

Aeke evrquing

yoory todd

Pi % eee
% % ne eee
% x eee eee
eee nt eee il
eee It ene alt
% % eee x
% % ooo m
x % eee x
eee J. ope er

% % . ee
4% x ee eae
x % eee Pie

‘POT | POT | “G8 | “G8
OWL Comtey) onl py onl
08 | 0G | OL | 0G

eye ee

* *

aie.) eee

0 eee
Aon eee
wisie eco
eee ee
a0 eee
eee eee
eee Cyr)
p7ita| ae
eee bo)

SS | | |

‘C6 | G6
0} | 04
GL | 0G

‘ulseg | “Ulseg
qooureul ueu
-youy | -uolq [ly

‘NISVG NVUWV

“UISeg,
yorporg,

erate Pena en em SRO LO ‘TLOPLV BOTPEN
shacetelaye (‘wuag) VIVSIAL] BUIINTIA
reeeerees saga “epootsshqe BOSssty,
wee ccecevcce (‘T) BIQI10}4 VTOpAAn YT,
"1+** (rr) tuvoojed-sod sreytsody
pe senile ‘unjyepun wnuUloong
“(Co nq) stpoeas (oydig) susnyy
“+ (rr) snnbiyue snuoposkayQ

eee cone rere ee reeeecesereee (“quo )
wnoaing § (vpg) wvuloyotno,g

‘"VOSATIOP

pelea <P gap ‘BOUTS ITA

ee ocencecncnceccce AN “eq Bos
alelainioretelelelerets “0T “eynjuoUL BIPlOsy
mie) a eveis\eleielxiein "T ‘SITVUIJSOJUT BUuOory)

“rnp “euUeAsOTaTTVAed vT[eA109
pacrenent cacy ‘ervemod edavos]og
ee eee cereessrese aac dA ‘eorqysnd
“ts wag ‘a “elrepnssoas epotyg
wee eens ccesces 19; ‘ey BvUlyoe eryyudg

‘VIVOINO T,

FAN H SOROS

467

FAUNA OF THE CLYDE SEA-AREA.

Bae

PEEE

AAazeEaa

(Bae EE

‘AESEESEEE

EEEEE

Se ee i (7) esosni VAVOIXVG .

se eeeeeree eveee uno ‘eqqis B[nqioy .
unoug “eo1ydrye Vayoryy *

eae (‘ufap) stsuoutpno1a —-
eee weeeeees (‘2uopy) snsonxep
(saquog) snsoulsnatey uopoydArgn
weesee seeeeses “Uuad “@yeAO snue A
Se eed (7) 1090 BIPIVIOST

baie trees (rr) vorpuvyst vutadsg -

4_ “cnuryayon
“a “qabg aT. ‘uingelosey wnipaszey)
Scie. “Ear eters nT) eyeoTns
sree umoug “eoIydtt]e o41e]SV
SS AAI ee CW) eqynUlu VparT
306.0. cueu eas - sree (‘guopy) sinuey e
woe rece cceseres UU0L “eqyeoTns

Sule'w'e erwin a go ewES bees (-T) snepnud

seteeeeeceeroes smo “BDIqIt BONY
ee eecece eeee (7) Byeqgieq B[OLpojy
tee eee eee eeeee ‘7nyl ‘snyetays
"++ nr ‘snyerperueydes
eect ecesccce ecccccecs (7) oisnd

rrrseseeeeee(rr) srapmnodedo weyeg °

Selsele Wales Mewes “far ‘eotydry]o Bully
sete er eesoes Mf ‘srumtojiy[oyed

cece cecscces "T ‘aniddryde elulouy

8" wag ‘SnyeUulsieul WOJTGO
steers (Cr) BULOVON BTeInjoung
*7yq ‘snuURISeT[IUL ( )
“T ‘snuryddziz (snurydéz1z)

“T ‘sniaeiould (vpnqqry) snyoouy, *

ee ee "104d “epIp.ios
aces ‘TnseyUOPL ——

“7 ‘siyeque waniTeyueq *

LINN. JOURN.—ZOOLOGY, VOL. XxX.

MR. W. E. HOYLE ON THE DEEP-WATER

468

EEEEZ

a Ee
+ 32

‘bE
04
0€

“uorng

“SIC! -Suory

qoo'T
seddq

‘Tl0H
yoo]

* * * *
e eee * *
% eee eee eee
* ¥ * *
x eee eee %
% : sg x
ee ° eee x
os * eee .
* * oes *

— enn
i em | | | |

4 0} 04 04
OF c¢ ¢9 0g
‘uIseq | ‘U9ATLT}G ‘oud

uooung | yoo y | yoory zoddg

“UISeg
oviquing

04 | 04

“UISeg
yooureul
eeu

x | *
gt
ae
28 -Foe
“| 4
*
"8 | G8
07 | 03
OL | 0S
“uIseg
ueul
“Ted T

* | *
saw, tae
* | *
"C6 | °G6
0} | 04
GL | 0G
"UISe
yorporg,

‘NISVG NVULV

* *(7) snpaequieq sninsedny ‘89

eeeeee

ANBocnacs (7) wIBUL SepoyWT “19
“8 Uuad “egoaaqny Bryeqy “99
seeee . yooaT ‘snqjisnd "GQ
"+ «7 Soyeandep snunyJog ‘$9
** wuag ‘etodse ommoudking *e9
De oy ‘snqeqjsor "29
saaeeraneee iteseeeteees equ

“py ‘slajsoaisu0T snyoudyAoueyg *[9

evese .

eeecoe

es -yonaT ‘snyezoIVOD ‘09
cece cere eece CIE ‘gnourre sedy]T "eg
‘uuag ‘sIsueyjes1op snqeuy "gg

*VHOVISOUO

cece vce ee reese resescers ee (7)

styuedaes-jndeo ‘eurpnyeaqeiay, "LG

*V@Od0IHOVUG

*'SaQuouT ‘eyelAorqqe viaeprdsny “9G

eeeree

eceeee

Pee Scarce (: QUOT) SINUS} ‘GG
eevcee eee WLAN “epryiu ‘$C
(poo) vq[e (vaqy) ojeteg “ge

469

FAUNA OF THE CLYDE SEA-AREA.

I AE OC en ee ee ee ae ee

Aaeae

7 Zz 2

Zz, 2

PEE ZzZe

PAS

EZ,

*
|

°K OK OK Ok

ZO) oorgny

Cyr CE )staeqpourasy vuepdoydeT “6G
reese (Cry) stavpoolmtiaa Bndtag “$6
erent (‘ysag) @xo[duit BUBISITLT “SG

criteetens (ang) eruoard vaqeyg “76
| ceeces (‘190c) BOLSTOq vVIIVUuloIg “16
[tresses TA ‘tsholpor vImeUINy “06

de yisis vinieleielnlelsisflvisie se sicwisibiers (172
vroaourpeAy] “68

"T ‘eyeotnov ojtporydy “8g
‘STNUA A

‘M PE dy ‘ejaqyered voqopy ‘18
“** (‘gmog) STULODISUOT SNANPIV “98
5S apee yaT “eso[novul BALUBP “CR
“** (guopr) BoovapuryAd BIBTLMOKD “FS
“pgp ‘sedyary BUBpolty “Eg
(sung “pz) tposey visneydoaiog *Zg
se eeeeee eee sete eee ere ress eee (sung

‘W) vorseatou souegdnody “Tg
ossvy ‘St[Np? VAIN “08
"8" spy "AT ‘snyepnUTyoe "6L
“+ wnyoury ‘TaeUTy wosueID “gy,
rseree +r) snotseadou sdoryde yy *)),
are 1ag ‘watpuvseyy strBool[VQ “9/,
seeseesee «(yon 7) BSOBNA VPIUN *C),

seer eere eeeaeee

steeseeee onary “exoyturenbs ‘PL
woere rereees WORaIQUyy ‘exo "Sh
ts : +g dy ‘esaadstp voyyeley “ZL,

(wosdwoyy,) stawyt snansedeuy ‘TL
ssseeeees(sahOLY ) susosaqnd ‘OL
eeareeree (yov0T) Iixnvepltg oo ‘69

MR. W. E. HOYLE ON THE DEEP-WATER

470

wD
*

ZEEZ

aze B Baaaeee

. .

‘mOTynG
"Leer . Zuory

Yooy
sodd

“OF
04

TIO
yoo]

‘eg
0}

“UIseg,
uoound

* * x
* ae |
eee eee _*
is A a
nf ae %
* ae eb
ve oa |
* % x
“LP ‘LL | “LL
0} 04 04
cs g9 | OS
"UOATIYQ ‘oud
yoo, | yoory soddg

69
oF
OG

ULSUg
evaiquing

i ° eee eee eee
* * * * ae *
* * * * * *
* % a eee eee eee
% % . eee ove ees

*k
ok

‘POT | FOT | “E8 | °€8 |°6 | G6
0% | 09 | 209 1407 | 09 |) 04

08 | Of | OL | OF | GZ | Og

‘uisegq | ‘ulseg
qoouwu | ueu
“youl | “Gerda

“UIseg
shy SERS

‘NISVG NVUUW

srevsereseesaeeseneeesseens: (cya)
wnyAqdejued x1aqyorydg “g]T
CRW AO) etsra vuos01ydg "ZIT
“* (Comp Wf ‘QO ) Stus0j. —— “TIT
“* saquog ‘ale vantqgdmy ‘OTT
‘IT °O) ‘eByeoqnoe stjoydoiydy ‘¢o]
srreserreeee(quaq) B8041OORT ‘801
cislatele Wieieipisiere “(saquou) epiqye “JOT
reeres Cog) stume eyddjsorydg ‘901
ses aT aT “GQ “BOOB[OIA "COT
Naeem Me ‘gueqnt SVILIISV “FOL
eet" (YOUWT) BYL[NDO VTTOIQIID "SOT
‘QO) +suedseARp WNIplvooulypY ‘ZOT
Pca entiae aanete c eneeeetenes ear
‘7 ‘Q ‘sneindind snsurjzedg ‘TOT
“*9*(saquo7) BAOJLIAT stsdossig” “OOT
SO ‘snjue[nose snuUlyoy "66

Sew ears steer es eeeesserere (y2ner

/ ‘osp) sI[VUl4soyUL VIAMYAOTOFT “9G
CUMAT A °O) snsny ouodgy, “16
See er er (‘sdwoyz,)
Tuueurpus FT VIAVUININY “96

“VLVNUACONINOW

FAUNA OF THE CLYDE SEA-AREA. 471

If the preceding table be summed up in the same manner as the
first one the result is found to be as follows :—

Arran Basin. d|/ 2]. la
= 3 Sed i (S)
Upper | 5] A] SS Ae
Loch mia m a
Kilb Inch Siro tees
Brodick | “ibren- ne a ae Fyne. 3 S cae
Basin. nan mMarnoce. rae = a =)

Basin. Basin. Basin.

50 | 75 | 50 | 70 | 50 | 80 50 50 | 65 | 35 | 40 | 35 | 30
bon \eto.} to. | to | to | to to ton} to. | to- | to: } to..| to
92.| 92.) 85.| 85. | 104.) 104. 62. hie tee) Al: 55.1 40.) 34

TE Meee ies | OY OT ie Merete terse? ot |. ol
BPOHUSCA ...000-0000- oe) Oo; 2) 3 | 23.) 21 16 Pe Or) 1% 16: | 7
Brachiopoda .........] ... emiecr beds Wy te) Bil) cesses 1
BEPUSEACER ......05050. ey oP oye 2 is | 12 15 weer, clisy 6 | 6
BPGETICS «20.5 sc0ccce0. eC ae a eo ieee tl ab 4! OI
Echinodermata ...... Sezer yor) 9} 9 5 Ge zRi Gol 2 bh-3 | 6
Totalact: 3s:::. 40 | 24 | 26 | 12 | 54 | 51 36 39 | 22 | 26 | 42 | 32 | 21

It is cbvious at once that these numbers do not show so
clearly as those previously obtained the gradual diminution in
the number of species in the different basins. There is a slight
tendency in this direction, but the exceptions are rather nume-
rous, and if we take the bottom faunas of each basin instead of
that below 50 fathoms the series of numbers is :—

94, 12, 51, 36, 22, 26, 42, 382, 21.

This result is extremely interesting because it seems to show
that the bottoms of the remoter basins have a fauna which
approaches the more seaward basins in respect of variety more
nearly than do their faunas taken as a whole. It suggests the
possibility that we have in these basins, in addition to the fauna
derived from the present outer seas, which seems to be gradually
making its way into them, a fauna which has been in them for a
much longer period.

Regarding the range of distribution of the species which are
confined to these depressions we find :—

LINN. JOURN.— ZOOLOGY, VOL. XX. 36

472 DEEP-WATER FAUNA OF THE CLYDE SEA-AREA.

Wide-spread | Northern | Southern

Species. Species. Species.

THRIGAER “ancides sree 4 5
Mollasea> ..cas0c002 30 15 2
Brachiopoda ......... 1
Crustacea ......scce0- 15 12 2
WRPMICS, ...xesi desea 3 3
Echinodermata...... 10 6 i

Totals wcscccors 63 41 5

Here, curiously enough, the wide-spread forms preponderate
over the others more than was the case in the previous list; but
it is noteworthy that the percentage of southern forms has
diminished, which emphasizes still more strongly the Arctic and
Scandinavian affinities of the Clyde deep-water fauna. These
results illustrate in a very interesting manner several of the
generalizations of the late Edward Forbes.

In conclusion, I may be allowed to express the hope that the
facts recorded above may furnish the nucleus of more extended
series of observations. I hope to continue the work as oppor-
tunities arise,and I shall be extremely grateful to any naturalists
who will furnish me with records of the occurrence, with the

exact locality and depth, of any species in the Clyde sea-area or
neighbouring seas.

Upper
Loc hy, Av

| Inch- |

Raeeraaey

rene Basin.

Basin.

. = <a Cumbrae
ne) NS.

Mintern Bros btn

BATHYGRAPHICAL CHART OF THE CLYDE SEA AREA.

ON THE ZOOLOGY OF FERNANDO NORONIIA. 473

Novres oN rHE ZooLocy oF Fernanvdo NoRoNHA.
By H. N. Rivtey, M.A., F.LS.

[Read 7th June, 1888. |
(Pate XXX.)

INTRODUCTION.

On July 9th, 1887, the writer, with Mr. G. A. Ramage, of
Edinburgh, started for Brazil to thoroughly explore the island
of Fernando Noronha, lying in long. 32° 25’ 30" W. and lat.
3° 50' 10" S., at a distance of 194 miles N.E. from Cape San
Roque, coast of Brazil. On arriving at Pernambuco we were
joined by the Rev. T. S. Lea, who came as a volunteer at his own
expense. The cost of the expedition was defrayed by the Royal
Society. After some delay at Pernambuco we embarked in the
‘Nasmyth’ steamship, trading to Liverpool, which was permitted
to land us at the island, as the regular steamer trading between
Pernambuco and Fernando Noronha was detained for a long time
just as she was due to start. We arrived at our destination on
August 14th, and remained there till September 24th, when we
returned by the little Brazilian steamer to the mainland. We
occupied ourselves in exploring, and in collecting plants, animals,
and rock-specimens in all parts of the main islands, and visited
also most of the other islets which were accessible; but owing
to the absence of boats, which, on account of the convict-station,
are not permitted on the island, we were unable to obtain much
by dredging. The coral-reefs, however, at low tide afforded an
abundant harvest of marine animals and plants.

Having in the ‘ Introduction’ to my ‘“ Notes on the Botany
of Fernando Noronha,” printed in the ‘ Journal of the Linnean
Society ’ (Botany, vol. xxvii. p. 1), given a detailed account of the
eroup of islands of which this is the chief, as well as a history of
its discovery by Amerigo Vespucci in 1503, it will be unnecessary
to repeat what has there been stated. For the better under-
standing, however, of the special reports on Zoology which are
now furnished, the following extracts from the Introduction
referred to may be found useful.

Vespucci’s description of the trees and innumerable birds
is evidently correct, though most of the trees are destroyed,
and the birds far less abundant than they were then. The
lizards with two tails may have been a confusion of the

LINN. JOURN.—ZOOLOGY, VOL. Xx. 37

4.74. MR. H. N. RIDLEY ON THE

very abundant and conspicuous Gecko with the Amphisbena,
which is often called the snake with two heads, or may have been
suggested by finding an accidentally fork-tailed lizard, of which
anexample was obtained by our expedition. The ‘serpents”’
were doubtless the Amphisbena. The large rats are much less
easy toexplain ; at present the only rats occurring on the island
are Mus rattus, the common introduced black rat. It is impos-
sible that the animals seen by Vespucci could have been this
species, which could not at that time have been introduced.
1t is possible that there was formerly an indigenous rat-like
mammal, which became exterminated by the black rat. We
could find no tradition even of this big rat, and I fear it is quite
extinct. The only hope of recovering its remains lies in an
examination of the guano deposits of Rat Island, where its
bones might be preserved.

The number of insects belonging to the orders which are
well known as plant-fertilizers is surprisingly limited. A few
small species of moths haunted at night the bushes of Scoparia
dulcis, Cassias, &c. on the open spaces. A single species of
butterfly was very abundant on Rat Island and the main island,
but we never saw it visiting flowers.

The most important fertilizer was a small endemic hornet
belonging to the genus Polistes, which gathered honey from the
Leguminosx and Cucurbitacee ; and three small black species of
Halictus were caught in the flowers of the melons, Momordica
charantia, Oxalis Noronhe, and the mustard. The last plant
was also haunted by Yemnoceras vesiculosus, a pollen-eating
Syrphid. The only other insects which could also be considered
as possible fertilizers were Zuchytes inconspicuus, nu. sp., and
Monedula signata, two sand-wasps, Pompilus nesophila, n. sp.
(Hymenoptera), and Pstlopus metallifer (a Dipteron), but none
of these were seen at or near flowers. A small black beetle also
was found in the flowers of an Acacia in the Governor’s garden.

Though the number of species of insects was not large,
the individuals, especially of the Polistes and Halicti, were very
numerous, but at the same time they seemed out of all propor-
tion to the immense number of flowers to be fertilized. It is
very probable, however, that the majority of the Leguminose
and some of the other plants were self-fertilized.

The lake on the main island contained a species of Nitella and
an alga, an aquatic beetle and an Hemipteron, a new species of

ZOOLOGY OF FERNANDO NORONHA. 475

Planorbis, and an Ostracod, the latter also occurring in all the
streams of any size. The remaining streams and puddles pro-
duced dragonflies, a species of Gammarus, and a few algw. One
may compare this state of things with the freshwater fauna and
flora of the other Atlantic islands. The absence of freshwater
fish and amphibians is common to most small islands.

Just as with plants, a considerable number of animals have been
introduced by man into the islands intentionally and by accident :
such, for instance, as the Gecko (Hemidactylus mabouia), the
American Cockroach (Blatta americana), and its curious parasite
Hvania, a spider, centipede, scorpion, rats and mice, and Sztoplidus
oryz@. These, though usually plentiful on the main island around
the houses, are markedly absent from the smaller islets.

There are also many visitors which have arrived here by the aid
of their wings, probably assisted by a suitable wind. They
include a number of the peculiar terrestrial fauna, the land-
birds and the insects. On looking over the lists of species
taken here, we may note that the smaller birds are endemic, and
a large proportion of the smaller insects, The small butterfly
and almost all the moths are known from the mainland of South
America, and the dragonflies are also widely distributed forms.
All the winged fauna have a South-American facies, whether they
are endemic or of wider distribution.

There are other creatures unprovided with means of traversing
the ocean and not introduced by man. They include the
Amphisbena, Skink, the freshwater and terrestrial Mollusca,
and perhaps some of the feebler-winged and apterous insects,
the endemic ostracod, &c.

The Planorbis, Gammarus, and Ostracod, all supposed to be
endemic species, may possibly have been brought over on the
feet of Wading birds, which migrate here.

The presence of some others is more difficult to account for.
The Mollusca are almost all peculiar, and the two that are not
so are West-Indian. The Amphisbena and Skink are endemic,
and allied not to Brazilian but to West-Indian forms.

It is commonly said that reptiles and terrestrial mollusks find
their way across the ocean by secreting themselves, or their eggs,
on floating trees, which are drifted to islands; and though for
several reasons this does not seem a satisfactory explanation of
their distribution, yet the appearance of these animals here
suggests this as the means by which they may have arrived. As

37*

476 MR. H. N. RIDLEY ON THE

i have said, they are West-Indian in their affinities, and it is a
striking fact that the marine fauna and flora are mainly West-
Indian, while at least one of the plants (Ipomaea Tuba) whose
seeds are known to be constantly drifted about at sea, and

thus carried from place to place, is also only known from the:

West Indies. Another fact of interest in connection with
this sea-travelling fauna, if I may use the expression, is that
almost all the species noted occur on all the islands suitable for
their existence. Thus, on Rat Island the Bulimus Ridleyi, the
Amphisbena, and Skink are common on St. Michael’s Mount; the
Skink is a large species, but the island, being a mere rocky peak,
is unsuited for the Amphisbena.

On Platform Island the lizard and several terrestrial Mollusca
were found, while at the same time almost all the animals of
more recent introduction were absent from these localities, just
as is the case in the distribution of the plants. I believe, in
fact, that this part of the fauna and flora was established on the
island before it was broken up into the little archipelago of rocks
and islets of which Fernando Noronha now consists*. Perhaps
even this portion of the fauna and flora was introduced previously
to the deposition of the basalt over the masses of phonolite
which form as it were the skeleton outline of the island.

MAMMALIA.

No indigenous Mammals are to be found on these islands, and
notwithstanding their proximity to the mainland, where Bats
are abundant, no Bat of any species was observed by us, nor
had the convicts ever.seen any. Rats and Mice are exceed-
ingly common. The Rat (dus rattus) is here much paler
than usual, and generally of a grey colour, while albinos are
sometimes met with. It frequents the melon-fields and the
tops of the cocoanut-trees, and is very destructive. The com-
mon House-Mouse, JZ. musculus, is even more abundant, and
has suggested the name Rat Island (Ilha do Ratta), where it is

* On reference to A. Vespucci’s description of the place, it will be found
that he speaks of it as one island, so the breaking-up into an archipelago can
only have taken place within the last 400 years.

ZOOLOGY OF FERNANDO NORONHA. 477

as common as on the mainland. It swarms everywhere, and is
so tame that it is often caught by the hand. I have seen one in
the evening on the top of the inflorescence of a Crotalaria,
apparently devouring the young seed-pods. Albinos are often
seen. There being no birds or beasts of prey to keep these
animals in check, and food being particularly abundant, they
have increased enormously, and one of the employments of a
convict is to capture a certain number of rats and mice once a
month. At the monthly rat-hunt while we were on the island
over 3900 were taken; but we were assured that, in the dry
season, when the herbage which covered the greater part of the
island was dried up and burnt, the mice were compelled to leave
their holes, and many more were taken. The hunts are then
undertaken weekly, and 20,000 have been caught ina day. The
bodies are piled up in the square after evening service, and the
numbers counted.

The Cat is said to have become feral on the main island; and
on Rat Island and one or two of the other islands we saw a large
black Cat which had escaped from an Italian vessel wrecked
there, and which had run wild.

In Amerigo Vespucci’s account of the island above quoted, he
mentions “ Mures quam maximi.” What these were we cannot
now determine, but it is highly improbable that they were dus
rattus.

A species of Dolphin was constantly seen in San Antonio Bay
and also off Rat Island. One was captured during our visit; its
stomach contained many cuttlefish and prawns, the latter very
similar to the common edible prawn of Pernambuco. Whales
also passed within sight of the island on one occasion, but we
did not see them.

AVES.

By Rk. Bowpter Suarpre, F.L.S., &e.,
Assistant in the Zoological Department, British Museum.

The birds of the island are not very numerous as regards
species, and apparently there are only three indigenous Land-
birds. The species of Sea-birds found by Mr. Ridley are
precisely what one might have looked for, but it is a little
remarkable that no Petrel was observed.

478 MR. H. N. RIDLEY ON THE

Fam. VIREONIDA.

1. VIREO GRACILIROSTRIS, sp. 0.

V. similis V. magistro,et forsan proximus, sed forma graciliore,
coloribus dilutioribus, facie laterali pallide flavicante, et rostro
valde tenuiore et graciliore distinguendus. Long. tot. 5-7, culmin.
0°6, ale 2°5, caudex 2:25, tarsi 0°8.

Five specimens were procured, and after comparing them with
the series of Vireonide in the British Museum, there is no doubt
that the Fernando Noronha bird comes nearest to V, magister,
of which species the Museum has now a large series from the:
islands of the Bay of Honduras, presented by Messrs. Salvin and

Bill of V. gracilirostris. Bill of V. magister.

Godman. The yellow face and the slender bill distinguish it at
a glance from V. magister.

Fam. TyRANNIDZ.

2. Evatinra RIDLEYANA.

Elainea Ridleyana, Sharpe, P. Z. S. 1888, p. 107.

This species has been fully described by me (/. ¢.). Dr. Sclater
(Cat. B. Brit. Mus. xiv, p. 189) does not consider it to be very
different from 2. pagana, but the size of the bill is very marked
in the insular birds.

[This bird occurred only on the main island and Rat Island as
far as we saw, and was very common in the gardens and in the
woods. We saw only a few nests, and of these only one was
finished and contained an egg, which was destroyed in an attempt
to reach the nest. The egg was white with dark red spots.
The nest, which was about three inches across, was made of the
tendrils of Cucurbitacez and a few fine twigs, but lined thickly
(and in fact almost entirely constructed in some cases) with the
woolly down of the seeds of Gonolobus micranthus. It was
placed often in the bare branches of a Burra or Hrythrina tree,
or in a Cashewnut-tree.—H. NV. £. |

ZOOLOGY OF FERNANDO NORONHA. A479

Fam. CoLUMBID.

3. ZENAIDA MACULATA.

Zenaida maculata (V.), Sel. § Salv. Nomencl. Av. Neotr. p. 132 (1873).

Zenaida aurita, Gray, List Galline etc. Brit. Mus. p. 14 (1855).

Zenaida noronha, Gray, List Columbe, p. 47 (1856, descr. nulla).

The bird from Fernando Noronha is merely a small race of the
ordinary Z. maculata of the South-American continent, with a
slightly shorter wing (5°1-5°4 inches) and tail (2°75-3°2); but as
some Brazilian specimens are of the same dimensions, I do not
see how the idea of a small insular race can be maintained.

[This little Dove is exceedingly common on all the islands where
it can find food, and flies about from one island to the other,
singly or in flocks of from 2 or 3 to 30. It is very tame, and
even when fired at, or alarmed, usually goes but a short distance
before settling. The nest is loose in texture, about 6 inches
across, and built of small sticks of the Spermacoce, vetches, &c.,
and lined with roots. It is placed often in the bare branches of
_ a Spondias or Burra, with no attempt at concealment. The eggs
are two in number, white, blunt at both ends, and about 14 inch
long. One bird shot off its nest proved to be a male. The convicts
catch these birds both for eating and as pets, keeping them in
wicker cages. They are fed on the seeds of Cassias and other
Leguminose and Cucurbitacee, and probably the fig and other
succulent fruits.— H. NV. £. |

Fam. LArip”.

4, ANOUS MELANOGENYS, Gray; Sharpe, Phil. Trans. vol. 168.
p- 467 (1879).

Two adults and a young bird agreed perfectly with speci-
mens obtained on St. Paul’s Rock by the ‘ Challenger’ Expedition
and determined by Mr. Howard Saunders. ‘The young bird is
browner than the adult, and has the head sooty brown with
some white on the forehead, eyebrows, and occipital region.

[This Noddy was very common on the island, and is called
“ Viuva preta.” A specimen also flew on board the vessel as
we were going to Pernambuco from Europe, about a day’s steam
from Fernando Noronha. The species nests in small colonies on
the rocks in various spots, and also in trees in the Sapate. An
ege was obtained from a nest on St. Michael’s Mount; it was

480 MR. H. N. RIDLEY ON THE

oval and blunt at both ends, 2% inches long, and about 1 inch
through in the thickest part, chalky-white in colour, marked
somewhat sparingly with underlying ash-grey, and overlying
sienna. <A living young bird from the nest was brought to me,
but soon died. H. NV. R.]

5. Gya@is CANDIDA (Gm.); Sharpe, t.c. p. 465.

One adult and two young birds. The latter are white like
the old birds, but have much smaller bills. |

[This is a common bird in many parts of the island, nesting
in trees, especially those of the Sapate, where there is a colony
near that of the Anous. The bird is called “ Viuva bianca.”—
FL IN Pte

Fam. PELECANIDA.

6. PHAETHON HTHEREUS (L.); Sel. & Salv. Nomencl. Av.
p. 124.

Of this Tropic-bird two specimens were procured. It is
common on the island, nesting on the Peak and on other rocks
and cliffs. An egg was obtained on St. Michael’s Mount. The
birds were taken in snares by the convicts.

7. SULA LEUCOGASTRA.

An adult and a young bird. This species of Gannet, known
as “Mbebu,” is a common bird, nesting on cliffs on all the
islands. The young are pure white.

Besides these birds we saw several of which no specimens
were procured. Zachypetes aquila was abundant, nesting on St.
Michael’s Mount, and a small species of Albatros appeared several
times round the island, but kept well out of gun-shot. Three
species of Waders were seen :—One, a small Plover, of which we
twice saw a flock at San Antonio Bay, and once or twice single
birds flying along the coral-reefs ; a bird resembling a Yellow-
shank, grey and white, of which a pair appeared at San Antonio
at the end of our visit ; and a single specimen of a Sandpiper, at
the same spot and time. These wading birds were all very shy,
in marked contrast to the endemic species, which suggested that
they were migrants, and had come from the mainland, where they
are more cautious at the sight of man. ‘The last two species
appeared on the same day towards the end of our visit, which
confirmed the view that they were migrating.

ZOOLOGY OF FERNANDO NORONTIA. 481

REPTILIA.

By G. A. Bouteneaer, F.ZS.,
Assistant in the Zoological Department, British Museum.

Only three species were found, viz. a Gecko (Hemidactylus
mabouia, Mor.),a Skink (Mabuia punctata, Gray), and an Amphis-
bena, described below.

The Gecko is of a widely-distributed species, ranging over the
greater part of Tropical America and Africa.

The Skink was originally described from two specimens ob-
tained on Fernando Noronha by H.M.S. ‘ Chanticleer,’ but has
since been recorded from Demerara. The specimens brought
home by Mr. Ridley are 10 in number; two have 36 scales round
the body, the others 38; in one specimen the frontonasal touches
the rostral and in another the two shields form a narrow suture.

AMPHISBHZNA RIDLEYI, sp. n.

Under this name I propose to designate an Amphisbena of
which a specimen, stated to be from Porto Bello, West Indies *,
presented by Capt. Austin, R.N., has been in the British Museum
for nearly 50 years, and was referred by Gray, Strauch, and
myself to 4. ceca, D. & B. The same species has been found
by Mr. Ridley on Fernando Noronha, and on re-examining the
question I find that A. ceca, which occurs on various West-
Indian Islands, but which was unrepresented in the British
Museum when I published my Catalogue, must be regarded as
distinct from the one with which I have now the pleasure of
connecting Mr. Ridley’s name.

16 specimens were collected by Mr. Ridley. One has 180 annuli
on the body, one 181, one 182, three 183, two 185, two 186, one
187, one 188, one 189, one 190, one 195, aud one 196; two have
18 annuli on the tail, eleven 19, and three 20. The “ Porto-
Bello” specimen has 189 annuli on the belly, and 19 on the tail.
The number of annuli in five specimens of A. ceca (including
the type) recorded by Strauch are respectively 212415, 215413,
927+18, 280+16, and 247+15. Dumeéril and Bibron give
926-329+18. <A specimen from Porto Rico, which I owe to the

* T am unable to find such a place either in the West Indies or Northern
Brazil, but as the other of the two specimens presented by Capt. Austin as from
“ Porto Bello” belongs to a North-Brazilian species (Amphishena vermicularis),
T entertain little doubt that both were obtained in Brazil.

482 MR. H. N. RIDLEY ON THE

kindness of Prof. Litken, has 228+19. Considering that the
number 247+15 given by Strauch is taken from a specimen in
the Paris Museum, received from the Copenhagen Museum as
from the island of St. Thomas, where only A. fenestrata (Cope)
= antillensis, R. & L., is known to occur, as Prof. Liitken
kindly informs me, it is clear to me that the specimen with 247
annuli belongs to A. fenestrata. The number of annuli would
range, in A. ceca from 212 to 229, and in the present species
from 180 to 196. According to Strauch, the length of the labial
border of the first labial shield in A. ceca is about one half the
length of that of the second; on Peters’s figure of the type
specimen, as well as in the Porto Rico specimen before me, it is
about two thirds; in A. Ridley both are equal, or the former is
a little longer. The snout is longer and somewhat more pro-
minent, the tail thicker and more obtuse in A. ceca than in
A, Ridleyi. The ventral segments of the two median rows are
broader than long in the former species, the coloration of which
is also different. I have therefore no hesitation in establishing
a new species, which may be characterized as follows :—

Premaxillary teeth 5 or 7, maxillaries 5-5, mandibulars 8-8.
Snout obtusely pointed, shghtly prominent. Tail thinner than
the body, tapering. Rostral small, triangular; nasals forming a
short suture ; a pair of very large prefrontals, followed by a pair
of much smaller frontals ; eye hardly distinguishable through the
ocular ; a postocular, no subocular; three large upper labials,
the second and third forming a suture with the ocular; lower
border of second labial as long as or a little longer than that of
the first, in contact with the second lower labial only ; mental
quadrangular, followed. by a large seven-sided chin-shield, which
is much longer than broad ; three lower labials, second very large.
180 to 196 annuli on the body and 18 to 20 on the tail; the
divisions of the annuli longer than broad, nearly equilateral on
the middle of the belly, but nowhere broader than long; 16 to
18 divisions above, and 20 to 24 below the lateral line. Anal
shields six or eight. Preanal pores four. Uniform brown or
dark purplish brown above, pale brown inferiorly.

millim
Liemeth to-vent.: ia... ts 255 Se 250
UTD Sa cations, ack «SIG (iss Ravaie = 24:

metho

ZOOLOGY OF FERNANDO NORONHA. 483

PISCES.

By G. A. Bovuneneer, F.Z.S.,
Assistant in the Zoological Department, British Museum.

The following marine species were obtained :—

Apogon imberbis, L., Hemulon chrysargyreum,Ginther, Holocen-
trum longipinne, C.& V., Acantharus chirurgus, Bl., Dactylopterus
volitans, L., Gobius soporator, C. & V., Salarias atlanticus, C.& V.,
Salarias vomerinus, C. & V., Clinus nuchipinnis, Q. & G., Clinus
delalandii, C. & V., Gobiesox cephalus, Vacép., Pomacentrus
leucostictus, M. & T., Glyphidodon saxatilis, L., Rhomboidichthys
lunatus, L., Hemirhamphus unifasciatus, Ranz., Clupea humeralis,
C. & V., Murena pavonina, Rich., Murena vicina, Cast., Murena
catenata, Bl., and the new species described hereafter.

J ULIS NORONHANA, Sp. n.
ee eT at 27. 1. tr. 2.

eee

Length of head one third of the total (without caudal),
or a little less; depth of the body one fourth. Dorsal spines
shorter than the rays. The length of the ventrals is two thirds
or three fifths that of the pectoral, which is shorter than the
head. Caudalis truncate. Upper half of body and caudal
blackish, lower half yellowish white (in spirit) ; a whitish streak
along each side of the back, just above the lateral line; dorsal,
anal, pectoral, and ventral fins transparent, immaculate; a black
spot between the first and third dorsal rays.

Several young specimens, the largest of which measures 60
millim.

The nearest ally of this species appears to be J. lucasana.

MOLLUSCA.

By Epaar A. Suiru, F.Z.S.,
Assistant in the Zoological Department, British Museum.

The total number of Mollusca now known from Fernando
Noronha is 80, of which 72 are marine forms, 7 terrestrial, and
1 freshwater.

Previous to this expedition no land or fluviatile species had
been collected, and only 28 marine forms, all obtained by the
‘Challenger,’ have been recorded from this locality. Ten of

484. MR. H. N. RIDLEY ON THE

these were also collected by Mr. Ridley, who has now added 44
additional species to the list.

The general facies of the marine Molluscan fauna is quite of a
West-Indian type, as a perusal of the following pages will show;
and it will also be observed that some of the species have a much
wider and in some instances a very peculiar range. Of the land-
shells two are known West-Indian species, one has been recorded
from Brazil, Peru, and the island of Opara, and the remaining
four, up to the present, appear to be peculiar to the island.
One of these, however, Bulimus Ramagei, suggests a faunistic
similarity to Brazil, as the section of Bulimus to which it belongs
(Lomigerus), with one exception, occurs only in that country.

The single freshwater species suggests no relationship with
any particular region, and might exist anywhere, similar forms
being found both in the Old and New Worlds.

The following pages contain an account of the species obtained
by Mr. H. N. Ridley and his colleagues, after which is appended
a list of those recorded in the ‘Challenger’ Reports.

I. MARINE SPECIES.

1. Ocropus ruGosUs, Bose.

Hab. Mediterranean, Cape Verde Islands, West Indies, Rio
Janeiro, &c.

This species is common in pools at low water. After being
dried in the sun the arms are made into soup and eaten by the
natives. Mr. Ridley, however, informs me that it is comparatively
tasteless and of a soft gelatinous consistency.

2. Conus NEBULOSUS, Solander.

Hab. West Indies: Barbados, Cuba, Martinique, Sta. Lucia.

The operculum of a shell 65 millim. in lengtn is 17 long and
only 43 wide. It is thickened and carinate along the middle
beneath, the muscular scar occupying more than half the entire
length, and the nucleus is not terminal as stated by Messrs.
Adams * and Tryon f in their respective Manuals of Conchology,
but situated three millim. from the extremity. The growth at
first is regularly concentric, but subsequently, to suit the nar-
rowness of the aperture of the shell, the layers of increase are

* “Genera of Recent Mollusca,’ vol. i. p. 246.
+ ‘Structural and Systematic Conchology,’ vol. ii. p, 187.

ZOOLOGY OF FERNANDO NORONHA. 485

added at one end only, thus producing a long narrow operculum.
MM. Cross and Marie * have also noticed, in respect of C. impe-
rialis, CO. lividus, and O. rattus, that the nucleus of the operculum
is subapical, and doubtless it has a similar position in other
species. The description of the operculum therefore as usually
given in manuals and other works requires modification, and the
nucleus should be termed apical or subapical.

3. Conus Daucus, Hwass.

Fab. Barbados (Mus. Cuming) ; St. Domingo and Guadaloupe
(Kister) ; Cuba and Martinique (d’ Orbigny).

The single beach-rolled specimen has a very strongly marked
double zone of brown spots upon the middle of the body-whorl.
With this species I unite C. mammillaris, Green, C. castus, Reeve
(not C. castus of Weinkauff), C. archetypus, Crosse, and C. san-
guinolentus of Reeve.

C. Reevei, Kiener, placed by Weinkauff+ in the synonymy of
this species, is quite a distinct shell, which I regard as the same as
C. piperatus, Dillwyn, not C. piperatus, Reeve, which, as stated
by Weinkauff, is the same as C. erythreensis of Beck.

4, PLEUROTOMA (CRASSISPIRA) FUSCESCENS, Gray.

1843. Pleurotoma fuscescens, Gray, Reeve, Con. Icon. fig. 125.

1845. Pleurotoma nigrescens, Gray, Reeve, l. c. fig. 235,

1845. Pleurotoma paxillus, Reeve, fig. 285.

1850. Pleurotoma solida, C. B. Adams, Contrib. Conch. vol. i. p. 61.

Hab. Cuba (d’Orbigny); Jamaica (C. B. Ad. for solida); St.
Vincent (Reeve for nigrescens).

Pl. nigrescens and Pl. paxillus differ from the typical form of
the species in being very much smaller, PJ. solida being inter-
mediate in size.

In his ‘ Manual of Conchology ’ (vol. vi. p. 193) Tryon states
that Pl. nigrescens of C. B. Adams and Pl. nigrescens of Gray
are the same species. Having types of the former received from
Adams and Gray’s types also for comparison, I can state that
beyond a doubt they are distinct. Pl. ewprea, Reeve, is rather
an unsatisfactory species at present, and I am rather inclined to
believe that, as suggested by Tryon, it will prove to belong to
this species also.

* Journ. de Conch. 1874, pp. 333-359.
+ Conch.-Cab. p, 312, no. 53.

486 MR. H. N. RIDLEY ON THE

5. Murex (OcINEBRA) ALVEATUS, Kiener.

Hab. Panama (Reeve, Kobelt, Sowerby); West Indies (Zryon).

As suggested by Tryon *, I think there must be some mistake
with regard to the locality “ Panama” which has been assigned
to this species first of all by Reeve and afterwards by others. I
have never seen a specimen from that locality, and Mr. G. B.
Sowerby informs me that he has frequently received it with
collections from the West Indies, but never from the Pacific side
of Central America. J. erosus, Broderip, JL obeliscus, A. Adams,
Triton Cantrainei, Récluz, and probably IZ. pauperculus, C. B.
Adams, are perfectly distinct from the present species and from
one another. This is another example of Tryon’s rash and indis-
criminate “lumping” of species, which detracts so much from
whatever value may be attached to his work.

6. PIsanra Pusio (Linné).

Buccinum pusio, Reeve, Con. Icon. fig. 43.

Hab. Wonduras and St. Thomas (Coll. Cuming); Sta. Lucia
(d@’Orbigny as Purpura accincta) ; Ascension I. (Conry).

The specimens from Fernando Noronha are rather small, and

much more distinctly striated than certain examples from the
West Indies.

7. PurPpuRA HHMASTOMA, Linné.

The specimens obtained by Mr. Ridley constitute a well-marked
variety of this well-known species, both as regards form and
colour. They have the spire more elevated in proportion to the
length of the aperture, and only the two uppermost of the four
series of nodules on the body-whorl are distinct. The interior
of the aperture is greenish blue, reddish near the labrum, which
is bordered within with black-brown, upon which the fine orange
or yellowish lire are very distinct. The exterior of the shell is
purplish black, streaked and spotted with greenish white.
P. hemastoma is known from the West Indies, West Africa,
Mediterranean, Ailantic coasts of France, Spain, and Portugal.

8. CoLUMBELLA MERCATORIA, Linné.

Hab. St. Vincent, Grenada, Nevis, Cuba, Martinique, and
Sta. Lucia (Brit. Mus.).

With one exception the eleven Fernando Noronha shells are

* Man. Conch. vol. ii. p. 128.

ZOOLOGY OF FERNANDO NORONHA. 487

white, variegated with very dark brown or black. The single
specimen, which differs from the rest, is of a pinkish tint sparingly
marked with rich brown.

9. Oxtva LiTERATA, Lamarck.

Hab. West Indies, Gulf of Florida.

Two beach-rolled shells are all that were obtained. They have
the transverse lire on the inner lip extending over nearly the
entire length of the columella.

10. Oritva (OLIVELLA) NIVEA (Gmelin).

Hab. St. Vincent and other islands of the West Indies,
Venezuela, and Brazil.

The shell named by Mr. Watson * Oliva fulgida, Reeve, from
Fernando Noronha, does not belong to that species, but is a
prettily coloured example of O. nivea. O. fulgida differs from
Gmelin’s species in the form of the columella and basal cauda.
The columella of O. nivea is very peculiarly excavated, and this
may be seen by looking as far within the aperture as possible.
No such excavation occurs in O. fulgida, which also does not
exhibit the numerous oblique folds or lire on the columellar
margin of the aperture which distinguish O. nivea.

A second species is quoted with doubt by Watson from Fer-
nando Noronha, namely O. pulchella, Duclos. The two fragments
referred to this species seem to me to bear little resemblance to
Duclos’s figure; but I have no hesitation in considering them
specifically identical with the other specimen from the same spot
which I refer to O. nivea.

11. Levcozonta crnauLirera (Lamarck).

Hab. West Indies, Honduras, West Africa.

L. rudis, Reeve, is I consider quite distinct from this species.
With this exception I agree with Tryon in his synonymy, and
would even suggest the propriety of maintaining L. leucozonalis,
Lamk., as a variety of this species.

The specimens from Fernando Noronha have stout rounded
ribs, exhibit a distinct submedian white zone on the body-whorl,
and have the aperture inclining to orange.

Tryon questions the West-African habitat of this species, but
J am inclined to think it correct, as in the British Museum there
are three specimens from that locality presented some years ago

* Gasteropoda of the ‘ Challenger’ Exped. p. 224.

488 MR. H. N. RIDLEY ON THE

by a Mr. Lewis, together with other species which are undoubtedly
West-African forms.

12. LEUCOZONIA OCELLATA (Gmelin).

Hab. West Indies.

The specimens obtained offer no differences from ordinary
West-Indian examples.

13. Lartrus spaDICcEuS (Reeve).

A single young shell seems to belong to this species.

L. coneentricus, Reeve, L. brevicauda, Reeve, L. gracilis, heeve,
and the present species are very closely related.

14. Mirra BARBADENSIS (Gmelin).

Hab. West Indies, Barbados, St. Vincent, &e.

MM. tessellata, Kiener, which Reeve named JZ. picta, is perfectly
distinct from the present species, and well known as a South-
African shell. Tryon*, not possessing or not having seen the
species, at once concludes, from their general superficial resem-
blance, that it must be the same as IZ. barbadensis. The sculpture
of the two is quite different. JZ. barbadensis is ornamented with
raised spiral lines, whilst JZ. picta exhibits transverse punctured
strie. The character of the outer lip also is quite different.

15. Mirra (Pusia) ansutata, Sowerby.

Mitra ansulata, Sowerby, Thes. Conch. vol. iv. p. 26, pl. 373. fig. 474.

Mitra microzonias, Reeve (non Lamarck), Con. Icon. figs. 185, 202;
Sowerby, l. c. fig. 635; Kiener, Cog. Viv. pl. 28. fig. 89 (probably) ;
Tryon, Man. Conch. iv. p. 183, pl. 54. figs. 568, 569.

Hab. St. Thomas (Mus. Cuming) ; “ West Indies, Morch, Krebs,
and Swift,” fide Tryon.

This species is said to occur in Polynesia, but the British
Museum Collection affords no evidence in proof of this statement.

The shell from Fernando Noronha belongs to that form of the
species as figured by Reeve (fig. 185).

This is usually considered the I. microzonias of Lamarck, but
if it be compared with the figure of that species in the ‘ Ency-
clopédie Méthodique’ (pl. 374. fig. 8), it will be seen that it is
a much more slender shell. The true M. microzonias has also
been figured by Kiister (Con.-Cab. pl. 17. figs. 12,13), and Reeve
also correctly depicts it (Con. Icon. pl. xxx. fig. 242 on left) under
the name of J. leucodesma. Sowerby in his description of

* Man. Conch. vol. iv. p. 118.

ZOOLOGY OF FERNANDO NORONHA. 489

M. ansulata does not mention the presence of a second white
zone on the body-whorl as represented in his figure, but this does
occasionally exist. Tryon places this species in the synonymy of
M. dermestina, Lamk., together with JZ. cavea, Reeve, M. Adamst,
Dobrn, I. pulchella, Reeve, MW. pisolina, Lamk., WM. histrio,
Reeve, and WM. consanguinea, Reeve. A more ridiculous instance
than this of the “lumping” of species I have never seen. Tryon
never could have examined examples of these various forms,
for if he had he would not have united them; he must have
been misled by the figures, or perhaps a little jealousy of non-
possession may have influenced him.

Reeve’s IW. leucodesma he says is beyond a doubt the same as
M. pardalis, Kister. From this it is, in my judgment, perfectly
distinct ; and the statement that ‘‘ Reeve’s figure of IZ. pardalis
is a Columbella” is sheer guess-work. The shell figured by
Reeve is in the British Museum, and not only is it a Mitra, but
correctly identified by Reeve as WU. pardalis, Kiister. What
right had Tryon to make such a statement in the face of Reeve’s
description, in which he properly characterizes the shell as a
Mitra with four plaits on the columella? Numbers of similar
absurdities occur throughout this work of Tryon’s, which might
have been avoided if more judgment had been used and the love
of “lumping” been overcome.

16. Mareine tna sacirtata, Hinds.

Hab. Bahamas to Brazil.

M. fluctuata, C. B. Adams, from Jamaica, appears to be the
same as this species.

17. Triron Ripweyi, sp.n. (Plate. XXX. fig. 1.)

Testa late fusiformis, albida, obsolete trizonata, zonis supra
varices aurantiis ; anfractus normales, superne declives et leviter
concavi, ad medium biangulati, inferne constricti, costis longitu-
dinalibus circiter 7 (in anfract. ultimo subtuberculiformibus inferne
evanescentibus) instructi, liris spiralibus tenuibus (in anfr. pen-
ultimo 7-8) aliisque longitudinalibus tenuioribus concinne can-
cellati ; apertura ovalis, alba; canalis brevis, dextrorsum versus ;
columella alba, superne arcuata, vix tortuosa, tuberculis vel liris
transversis supra callum tenuem munita; labrum compresse
varicosum, intus liris duodecim in paribus ordinatis instructum.

Longit. 19 millim., diam. 10.

LINN. JOURN.—ZOOLOGY, VOL. Xx. 35

490 MR. H. N. RIDLEY ON THE

This species belongs to the same group as 7. gallinago, Reeve
(Con. Icon. fig. 5), and 7. testudinarius, Adams & Reeve, and
some others. Although possibly not adult, the single shell at
hand is in excellent condition, and affords all the necessary
characters distinctive of the species. The last whorl has two
varices, namely the labrum and one on the opposite side. The
nuclear whorls are broken off; but, judging from the top of the
first normal whorl, the apex would be comparatively small. The
uppermost of the lire on the columella is rather conspicuous, and,
together with the uppermost of those within the labrum, forms a
semicircular sinus above.

18. Triton PILEARIS, Lamarck.

Hab. West Indies, Red Sea, Ceylon, Philippine Islands, island
of Anna, &c. (Brit. Mus.).

This, like some other species of Triton, occurs at the West
Indies and in the Indian and Pacific Oceans.

19. Trion (EpPrpRomuvs) TESTACEUS, MWorch.

Hab. West Indies (Moreh).

This species is very like Z. obscwrus, Reeve, but differs in
having more convex whorls, a granulated columellar callus, and a
narrower labral varix which is also hollowed out behind.

20. CYPRHA CINEREA, Gmelin, var.

Hab. West Indies.

With this species I unite C. clara of Gaskoin, with which the
specimens from Fernando Noronha agree. ‘This variety is of a
longer and more cylindrical form than the type, has only traces
of the black dotting around the base, and no purplish stain
between the teeth. Sowerby’s figure (Thes. Conch. pl. 307. f. 91*),
badly copied by Tryon (Man. Conch.vol. vii. pl. 1. f. 8), does not
represent the variety clara ; but a fair representation of it is given
by Sowerby on pl. 316, figure 222. The colour, however, is not
pinkish, and no dotting occurs along the sides in the types de-
scribed by Gaskoin.

21. Cyprma (TRIVIA) PEDICULUS, Linné.

Hab. West Indies.

One of the specimens from Fernando Noronha is remarkably
small, measuring only 7 millim. in length.

ZOOLOGY OF FERNANDO NORONHA. 491

22. LiTTORINA TROCHIFORMIS, var.? (Plate XXX. fig. 2.)

Littorina trochiformis, Dillwyn, Philippi, Abbild. vol. ii. p. 143, pl. i.
ff. 12, 14, 15.

Littorina nodulosa, Watson (non Gmelin), ‘ Challenger’ Gasteropoda,
p- 577.

Testa parva, fusiformi-ovata, grisea vel nigrescens, albo-nodosa ;
anfractus 6-7, convexiusculi, superiores granorum seriebus tribus
ornati, striisque spiralibus elevatis paucis sculpti, ultimus in
medio obtuse angulatus, seriebus quatuor cinctus, ad basim albo
punctatus ; apertura nigra, fascia basali pallida ornata, inferne
subacuminata ; columella lata, purpurea, superne macula lutes-
centi notata.

Longit. 19 mill., diam. 10. Apertura 7 longa, 6 lata.

” 11 ” ” 9. ” 63 ” 5 ”

The above measurements of two specimens from Fernando
Noronha show the variation in the form of this variety. The
white tubercles are rather acute in some specimens, whilst in
others they are scarcely raised above the surface. On the body-
whorl there are two approximated series at the periphery and
two above, and at the base is a tesselation of white and dark
spots.

The shells quoted by Mr. Watson from Fernando Noronha are
certainly specifically the same as those obtained by Mr. Ridley,
and are, I think, almost specifically distinct from the ZL. nodulosa
of d’Orbigny. They have less angular whorls and less acute
nodules, of which there are two series on the body-whorl above
the two principal series at the periphery, whilst in ZL. trochiformis
(=nodulosa, d’Orb.) there is only a single series. The aperture,
also, of the Fernando shells is darker and none of them exhibit a
second pale zone at the upper part, which is nearly always visible
in the West-Indian species.

23. LiTroRINA ANGULIFERA (Lamarck).

Litorina angulifera, Philippi, Abbild. vol. ii. p. 223, pl. v. ff. 12-15.

Hab. West Indies, West Africa and Pacific (Phil.).

Only a single young specimen was obtained by Mr. Ridley ; it’
agrees in all particulars with West-Indian specimens.

24, Torin1a mrutors (Menke).
Hab. West Indies.
Both Philippi and Hanley, in their respective monographs, admit
this as a distinct species, but I am inclined to think with the
38*

492 MR. H. N. RIDLEY ON THE

former * that 7. cyclostoma, T. nubila, T. cylindracea, and the
present species are mere varieties of one and the same form.

25. [ANTHINA FRAGILIS, Lamarck.

Of the various species figured by Reeve, that which he has
identified as Lamarck’s I. fragilis (Conch. Icon. pl. ii. ff. 6a, 60)
closely resembles the shells from Fernando Noronha. They have
the same perpendicular columella and the same division of colour,

the “ deep-violet ” tint of the base terminating abruptly at the
periphery.

26. CERITHIUM ATRATUM (Born).

Hab. West Indies, Pernambuco, and Rio Janeiro (Brit. Mus.).

I regard the C. caudatum of Sowerby as undoubtedly belonging
to this species. “Sicily,” the locality assigned by Sowerby in the
‘Thesaurus Conchyliorum ’ and in Reeve’s ‘ Conchologia Iconica ’
to C. atratwm, is evidently incorrect.

27. MITRULARIA ALVEOLATA (A. Adams).

Calyptreea alveolata, A. Adams, Reeve, Con. Icon. vol. xi. pl. 3. tf. 8 a—0.

Hab. Galapagos Islands (Reeve); St. Kitts, West Indies
(Brit. Mus.).

The single shell from Fernando Noronha possesses all the
characteristics of the type from the Galapagos Islands. The
specimens from Fernando Noronha, assigned with doubt to JZ.
wncinata, Reeve, by Watson}, in all probability belong to the
same species as that collected by Mr. Ridley. They are, how-
ever, only young specimens, so that their determination is all the
more difficult.

28. Hiprponyx ANTIQUATUS (Linné).

Hipponyx antiquatus (L.), Fischer, Journ. de Conch. vol. x. p. 5, pl. i.
ff. 1-9 (anatomy); Crosse, Journ. de Conch. vol. x. p. 20; Morch, Malak.
Blatt. vol. xxiv. p. 98.

Hab. West Indies; islands of Ascension, St. Helena, and
Trinidad in the South Atlantic, Peru and California (Brit. Mus.).

The single specimen from Iernando Noronha has the spire

more recurved than any other specimen I have seen and it is
inclined to the left.

* Conch.-Cab., Solariwm, p. 26.
| ‘Challenger’ Gastropoda, p. 461.

ZOOLOGY OF FERNANDO NORONTA. 493

29. Hipponyx GRAYANUS, var.

Hipponyx Grayanus, Menke, Carpenter, Proc. Zool. Soc. 1856, p. 4;
Crosse, Journ. de Conch. 1862, vol. x. p. 23.

The distribution of this species appears to be very extensive.
Carpenter quotes it from Galapagos, Sandwich Islands, Panama,
S.W. Mexico, Mazatlan, and St. Vincent (W. Africa). Some
Specimens in the British Museum from St. Helena, wrongly
named H. radiatus, Quoy & Gaimard, by Jeffreys *, and two
specimens from Fernando Noronha agree in all respects with this
species except in the more excentric position of the apex, which
gives them a more capuliform appearance.

30. NERITA ASCENSIONIS, Gmelin.

In his monograph of Nerita in the Conchylien-Cabinet, Dr,
von Martens mentions only the island of Ascension and Guinea
as localities for this species. I had previously noted + the fact of
its occurrence at the island of Trinidad off the Brazilian coast,
and now I record its presence at Fernando Noronha, where it was
also obtained by the ‘ Challenger’ Expedition.

31. TurBo (Catcar) OuFerst, Troschel.

Trochus Olfersi, Zroschel, Philippi, Conch.-Cab. ed. 2, p. 126, pl. 22.
i 1.

Calear Olfersi, Fischer in Kiener’s Coq. Viv. p. 18, pl. 77. f. 1.

Trochus digitatus, Reeve (non Deshayes), Conch. Icon. pl. 5. f. 24;
Sowerby, Thes. vol. v. pl. 504. fig. 135,

Hab. Brazil (Philippi & Fischer).

The localities quoted by Reeve and Sowerby, namely Central
America and Panama, will doubtless prove incorrect.

Failing to recognize this species as 7. Olfersi, Sowerby has
placed that name among the synonymy of 7. imbricatus, which,
however, is a perfectly distinct shell. 7. digitatus of Deshayes,
as pointed out by Philippi, Fischer, and Carpenter, is identical
with the common 7. waguis, Wood, of the Californian coast.

32. Troonus (Eurrocuus) susuBiInus, Ginelin.

Hab. West Indies (Philippi §& Fischer).
The two specimens from Fernando Noronha are more widely

* Ann, & Mag. Nat, Hist. 1872, vol. ix. p. 264.
| Lbid, 1881, vol. viii. p. 431.

4.94 MR. H. N. RIDLEY ON THE

umbilicated than the shells figured by Reeve * and Fischer }, and
also differ in form, being wider at the base and more shortly
conical.

Fischer has already pointed out that the localities of Reeve
and Lamarck, Swan River and Mauritius, are probably incorrect.

33. Trocuus (Eurrocuus) aemMMosts, Reeve.

This I believe, as in the case of the preceding species, is another
instance of a wrong locality (Philippine Islands) assigned by
Reeve.

Two specimens from Fernando Noronha agree in every minute
detail with the types in the British Museum, and, as the sculpture
and lineations are so remarkable, the identity is beyond doubt.
The umbilicus, which is as large as that of 7. jujubinus, at once
distinguishes this species from 7. nobilis, with which Philippi f
questioned its relationship. The type of Hutrochus was named
E. perspectivus by A. Adams; but as that name was previously
used by Koch for another species belonging to the same group,
Pilsbury has renamed it H#. Adamst.

34. FIssURELLA CANCELLATA, Sowerby.

Fissurella cancellata, Sowerby, Conch. Il. sp. 38, pl. 72. f. 29.

Hab. West Indies, Honduras.

With this species I would unite Fiss. suffusa, Reeve, and F.
lentiginosa, Reeve. A third species of the same author, Ff. egis,
is also very similar, but the form is a little more elongate and the
radiating riblets are squamose at the points of intersection with
the concentric lire.

35. FIsSURELLA ALTERNATA, Say.

Fissurella alternata, Say, Journ. Acad. N. Sci. Philad. 1822, vol. ii.
p. 224; Reeve, Con. Icon. pl. xii. f. 84 (probably).

Fissurella larva, Reeve, J. c. f. 98.

Fissurella Dysoni, Reeve, 1. c. f. 86.

Hab. Maryland, &c. (Say); Bermuda, St. Johns, Honduras
(Brit. Mus.).

The sculpture of the three above-named forms is essentially

he same and the character of the orifice is similar, and all have

* Con. Icon., Zizyphinus, pl. 2. fig. 12.
t+ Kiener’s Coq. Viv., Zrochus, pl. 18. f. 2.
t Conch.-Cab., Trochus, p. 86.

ZOOLOGY OF FERNANDO NORONHA. 495

the interior at the apex indented with a transverse line or pit at
the larger end of the perforation, as described by Say.

36. ? FissURELLA BARBADENSIS, Gmelin.

Hab. West Indies.

There are two or three specimens from Fernando Noronha
which closely approach this species, but I do not feel absolutely
certain of the identification.

37. ? FIsSURELLA NUBECULA, Linné.

Hab. Mediterranean, Spain, Morocco, coast of Gambia, Cape
Verd Islands.

Several specimens from Fernando Noronha in some respects so
closely resemble this species that I hesitate to separate them.
The interior is of the same greenish tint, the orifice has a purplish
tint or is ringed with purple, but the outer surface is uniformly
darker than Mediterranean examples. Withregard to sculpture
it is difficult to say that any material difference exists, as
specimens from any given locality exhibit slight variations in the
fineness and number of the radiating striz, such as may be noted
in the series from Fernando Noronha.

38. ACMHA NORONHENSIS, sp.n. (Plate XXX. figs. 3, 3a.)

Testa ovata, postice latior, mediocriter elevata, nigrescens,
radiis pallidis picta, ad apicem, paulo ante medium situm, erosa,
nigra, radiatim tenuiter striata, lieisque incrementi sculpta;
pagina interna intra cicatricem nigricans, apicem versus callo
tenui sensim albicans, extra cicatricem fere ad marginem ceruleo-
albida, ad marginem anguste nigro limbata, antice ab apice usque
ad marginem radio lato obscuro et postice alio latiore picta.

Long. 24 millim., lat. 19, alt. 9.

This species has a smoother surface than A. subrugosa, d’Or-
bigny (=Lottia onychina, Gould), from Rio Janeiro. Like that
species, however, it has in the interior a broad obscure ray from
the apex to the margin in front and a broader one at the opposite
end. These rays, however, are more distinct in the present
species than in the Brazilian shell. The external radiating striz
being very fine, do not, as a rule, produce a crenulated margin,
but in some instances a slight crenulation occurs. The surface
within the muscular scar is almost black, forming a marked con-
trast to the pallid space between it and the black margin. The

496 MR. H. N. RIDLEY ON THE

shells found attached to rocks, when placed upon a flat surface,
rest upon the anterior and posterior margins only, so that the
sides are slightly raised.

39. Curton (IscunocHIToN ?) PECTINATUS, Sowerby.

Chiton pectinatus, Sowerby, Con. Ill. pl. 174. f. 146; Reeve, Con. Icon.
pl. 26. f. 133.

Hab. ——? (Reeve); West Indies (P. P. Carpenter in Brit.
Mus.).

The marginal scales are not at all well drawn by Sowerby,
being much too elongate.

40. Curron (IscHNOCHITON) CARIBBHoRUM, Carpenter.
(Plate XXX. figs. 5, 5a.)

Testa elongato-ovalis, vix carinata, varie picta, griseo-olivacea,
albo, rufo et olivaceo picta, vel purpurea, interdum nigrescens,
albo virgata, valvis terminalibus concentrice et rugose granoso-
striatis vel squamatis, centralibus liris tenuissimis granosis cur-
vatis flexuosis ornatis, areis lateralibus rugose granosis vel squa-
matis ; valva postica pone apicem centralem leviter concava ;
cingulum minute squamatum, squamis minutis elongatis ovalibus
indutum, pallide roseo-griseum, dilute nigro tessellatum.

Longit. 27 millim., diam. 9.

Hab. St. Thomas (Brit. Mus.).

The above appears to be a manuscript name attached to speci-
mens in Cuming’s collection which are identical with few shells
from Fernando Noronha. The colour is very variable, some
specimens, when viewed from a distance of twelve inches, appearing
olive-grey speckled with white ; others are of a pinkish cream-
colour speckled with red or blotched along the sides in front of the
lateral areas with black, as in some of the specimens from Fernando
Noronha. The granules or scales of the lateral area and on the
front and posterior valves are peculiarly flat and are somewhat
transversely arranged on the former and concentrically on the
latter. The central areas are finely punctured along the centre,
and become more and more coarsely granosely lirate as the sides
are approached. One example is almost entirely reddish purple,
and others are blackish with a broad pallid stripe down the middle
from end to end.

41, Cuitton (ACANTHOCHITON) ASTRIGER, Reeve.
Chiton astriger, Reeve, Conch. Icon. pl. xvi. f. 109.
Hab. Barbados.

ZOOLOGY OF FERNANDO NORONHA. 497

Reeve describes this species as “ smooth along the summit, very
closely finely striated on each side.” This is not at all accurate.
The central portion of the non-terminal valves has a defined
elongate subtriangular space which is sculptured with minutely
eranular lines, and the sides are densely but rather more coarsely
granulated. The figure (47) of the detail of sculpture of C. sp7-
culosa, Reeve, which I believe to be the same species, gives quite
as good an idea of the ornamentation as figure 109. The outer
margin of the mantle bears a fringe of the same glassy spicules as
compose the tufts.

42. Doris, sp.

A single specimen was obtained, which appears to belong to the
same species as an unnamed example in the British Museum from
the West Indies.

43. APLYSIA, sp.

An animal about an inch long is all that was found. It pro-
bably is not full-grown. No attempt has been made to identify
either this or the preceding, as both belong to difficult groups
requiring special study.

44, SIPHONARIA PIcTA, var.?* (Plate XXX. figs. 4-40.)

Hab. Rio Janeiro (d’ Orbigny).

The specimens from Fernando Noronha are externally blackish
with numerous white radiating coste. The inner surface also is
much darker than in the tvpe specimens from Brazil. 8S. hispida,
Gould, also from Rio Janeiro, appears to be the same species.
S. lineolata, d’Orbigny, from Cuba, is a larger form, and the only
species that appears to have been discovered in the West Indies.
One or two species are known from Bermuda (S. brunnea,
Hanley); and Say has described a form from East Florida.

45. Venus (ANAITIS) PAPHTA, Linné.
Hab. West Indies and Cape Verd Islands (Brit. Mus.).

46, CARDIUM SUBELONGATUM, Sowerby.
Hab. St. Thomas, West Indies.

47. Carpium MepiIum, Linné.
Hab. West Indies.

* D’Orbigny, Voy. Amér. Mérid.’ vol. v. p. 469, pl. 56. ff. '7-11.

498 MR. H. N. RIDLEY ON THE

48. SEMELE CORDIFORMIS, Chemnitz.

1766. Tellina reticulata, Linné?, Syst. Nat. ed. 12, p. 1119.

1795. Tellina cordiformis, Chemnitz, Conch.-Cab. vol. xi. p. 208, pl. 199.
f. 1941-2. :

1815. Tellina decussata, Wood, Gen. Conch. p. 190, pl. 43. figs. 2, 3.

1822. Amphidesma orbiculata, Say, Journ. Acad. Nat. Sci. Philad.
vol. ii. p. 307; Reeve, f. 13.

1826. Amphidesma radiata, Say, J. c. vol. v. p. 220; Reeve (as of
Riippell), f. 12.

1832. Amphidesma lenticularis, Sowerby, Proc. Zool. Soc. 1832, p. 200;
Con. Ill. f.9; Reeve, f. 39.

184]. Amphidesma reticulata, Sowerby, Con. Illust. p. 8; Reeve, f. 29.

1841. Amphidesma snbtruncata, Sowerby, 1. ¢. p. 7.

1845. Amphidesma Jayanum, C. B. Adams, Proc. Bost. Soc. Nat. Hist
vol. 1. p. 10.

1853. Semele sinensis, A. Adams, Proc. Zool. Soc. 1853, p. 95; Reeve,
f. 28.

1853. Semele luteola, A. Adams, l. c. p.95; Reeve, f. 42.

1853. Semele modesta, A. Adams, |. c. p. 95; Reeve, ff. 35 a-b.

1853. Amphidesma cordiformis, Reeve, Con. Icon. f. 30.

Hab. West Indies, Bermuda, Rio Janeiro, Ascension Island,
St. Helena, Cape Palmas, and Fernando Po (Brit. Mus.).

Having carefully studied a large series of specimens from the
above localities, the types of S. lentieularis, said to have been
obtained in West Colombia, also the types of S. chinensis,
S. reticulata (Sow.), S. subtruncata, S. luteola, and S. modesta,
also the specimens figured by Reeve, I am inclined to believe that
all, the above-named forms constitute but a single variable
species.

Some authors have considered this species to be the Zellina
reticulata of Linné; but I am rather inclined to think, with
Hanley, that there is not sufficient evidence to determine this
with any degree of certainty. The locality “ China,” from which
Adams named a form S. sinensis, I regard simply as one more of
the innumerable errors of “habitat” occurring in Cuming’s
collection.

49. CHAMA, sp. incert.

Several specimens of a species of Chama were taken on the
rocks ; but the surfaces are so eroded and water-worn, that it is
impossible to determine them.

ZOOLOGY OF FERNANDO NORONHA. 499

50. Myrintus exustus (Lamarck), Reeve.

This is a West-Indian species, and was also obtained by the
‘Challenger’ expedition at Fernando Noronha and Pernambuco*.

51. Anca IMBRICATA, Brugquiére.

This species was also obtained by the ‘ Challenger’ Expedition
at Fernando Noronha; and other examples were dredged near
Cape York, N. Australia. This seems a remarkable distribution ;

still, as far as I can discover, there appears to be no differ-
ence in the shells.

52. Arca (Acar) ApAmstt, Shuttleworth, IZS.?.—Arca Adamsi
in Cuming’s Collection. (Plate XXX. figs. 6, 6 a.)

Testa oblonga, subquadrata, sordide albida, inequilateralis,
antice curvata, postice oblique arcuata, inferne in medio levissime
sinuata, lineis elevatis radiantibus aliisque concentricis cancel-
lata ; umbones parvi, parum remoti, paulo ante medium collocati ;
area dorsalis angusta, utrinque acuminata; ligamentum mini-
mum, adamantiforme, transversim striatum; pagina interna
alba, radiatim plus minus substriata.

Longit. 12 millim., alt. 74, diam. 74.

Hab. St. Vincents, Jamaica, and St. Thomas (Brit. Mus.).

Two specimens bearing the above name occur in Cuming’s
collection, and others from Jamaica presented by Dr. P. P.
Carpenter are also similarly labelled; but I have not succeeded
in finding any description by Shuttleworth of this species.

It is closely related to A. lactea, Linn., A. solida, Sowerby, and
some others. The points of contact of the radiating and con-
centric lire are nodulous, and a little coarser than in either of
the above-named species. The muscular impressions are clearly

defined by a raised ridge which is continued upward towards the
umbones.

53. Lima sguamosa, Lamarck, var.

Hab. Atlantic, Pacific, and Indian Oceans.

The distribution of this species is given in my Report upon the
‘Challenger’ Pelecypoda. The specimens from Fernando No-
ronha seem intermediate between L. squamosa and L. multi-
costata, having fewer ribs than the latter, and more than typical
examples of the former.

A specimen 22 millim. long has twenty-six ribs, and another
example 16 millim. in length has but twenty-one.

* Vide ‘ Report on ‘ Challenger’ Lamellibranchiata, p. 272.

500 MR. H. N. RIDLEY ON THE

54. SPONDYLUS, sp. :

A few odd valves picked up on the shores are too much worn
to be identified with certainty. About a dozen forms have been
described as West-Indian ; and doubtless it is one or more of
these species which occur at Fernando Noronha.

II. TERRESTRIAL SPECIES.

1. Hetrx(OPHto@yRa?) QUINQUELTRATA. (P1. XXX. figs. 7—-7c.)

Testa discoidea, supra leviter convexa, inferne anguste umbi-
heata, tenuis, viridi-flavescens, nitida; anfractus 7, lente accres-
centes, convexi, incrementi lineis striati, supra peripheriam in-
distincte concavus, antice haud descendens, intus plicis tribus
inequalibus perlucentibus munitus, lamellisque duobus validis
parietalibus instructus ; apertura semilunata, parva; peristoma
tenue, haud expansum.

Diam. maj. 6 millim., min. 54, alt. 3.

Hab. Found, both living and dead, at the north end of the
island, also on Platform Island.

HH. entodonta, Pfeiffer, from Ecuador, is an allied form; but
has a flatter spine, more open umbilicus, and no parietal lire.

2. Butimus (Tomigrrvs) RaMaAGEI,sp.n. (Plate XXX. fig.8.)

Testa subovata, rimata, solida, fusca, zonis angustis albis trans-
versis (in anfr, ultimo quatuor) cincta; anfractus 5, convexiusculi,
lineis incrementi subrugosis, striisque tenuissimis spiralibus
sculpti, ultimus magnus, antice descendens, post labrum con-
tractus, scrobiculatus; apertura irregularis, longitudinis totius 4
paulo superans, dentibus quatuor inwqualibus (duobus parvis in
pariete aperturali, uno magno compresso in margine dextro, uno
tuberculiformi valido in margine columellari) munita; peristoma
album, valde incrassatum, leviter reflexum, marginibus callo craso

junctis.
Longit. 234 millim., diam. 16.
” 174 ” ” 125.

The above measurements show that considerable difference
exists in the size of specimens ; and it is a curious fact that the
smallest example, obtained from a native, is the only one which
appears in fairly fresh condition. All the rest were found by
Mr. Ramage imbedded in sandy mud on a raised reef at Tobacco
Point, and have a semi-fossilized appearance.

The only forms at all approaching that now described are the

ZOOLOGY OF FERNANDO NORONHA. 501

species of Zomigerus, all of which, however, have a much more
complicated oral dentition, and are of different form, with the last
whorl ascending, and other marks of distinctness.

In general form and texture of the shell it is not at all unlike
Pythia inflata, Pfeiffer ; but of course has not the expanded lip
or the same dentition as that genus.

Two only of the twenty specimens which I have exawined
exhibit any variation in the teeth of the aperture. These want
the two parietal denticles.

3. Burrus (Butimvtts) Ripieyi, sp.n. (Plate XXX. fig. 9.)

Testa parva, ovata, superne acuminata, umbilicata, fusca, ad
peripheriam luteo-lineata; anfractus 5-6, convexiusculi, lineis
incrementi striisque spiralibus tenuissimis sculpti, ultimus antice
haud descendens ; apertura ovata, intus fusco-carnea, linea pallide
mediana ornata, longit. totius 1 ad equans; peristoma undique
expansum, carneo-albidum, marginibus callo tenui (interdum
crasso) superne subtuberculiformi junctis.

Longit. 12 millim., diam. 6.

Hab. Living under bark of Mango-trees in the garden and on
the north side of the island; also found at the base of the Peak,
north side, under stones, and on Rat Island.

I do not know any species sufficiently near this form wherewith
to offer a comparison. It resembles somewhat in form certain
species of Partula ; it faintly recalls, chiefly on account of colour,
Bulimus Jacobi, from the Galapagos Islands; and the spiral
striation, although finer, somewhat resembles that of some of the
species of the genus Plecotrema.

4, PUPA SQLITARIA, sp. nu. (Plate XXX. figs. 10, 10a.)

Testa minuta, rimata, albida, ovato-cylindracea; anfractus 5,
convexi, striis incrementi tenuibus sculpti, sutura vix obliqua
sejuncti, ultimus pone labrum subvaricosus; apertura parva,
quinque-dentata ; dente valido duplici columellari, uno minore
etiam duplici in pariete aperturali, duobus parvis ab margine
dextro remotis, quinto minuto basali ; peristoma anguste expan-
sum, album, marginibus callo tenui junctis.

Longit. 23 millim., diam. 13.

Hab. Platform Island.

This species is a trifle less cylindrical than P. pellucida, Pfeifter,
a Cuban species ; but has the number of teeth and their arrange-
ment similar. The columellar tooth, however, is double, the

502 MR. H. N. RIDLEY ON THE

upper portion of it being most prominent ; the tooth above it upon
the body-whorl is single, bifurcating at the end. The three re-
maining teeth are rather remote from the margin of the aperture.
The anterior part of the last whorl just behind and parallel to
the labrum exhibits a longitudinal swelling or varix, towards
which the lip expands.

5. Srenocyra (OpEAS) ocronorpEs, C. B. Adams.

Hab. Jamaica, Cuba, St. Thomas.

This species is more strongly striated than S. swbula, has
rounder whorls, a deeper suture, and a larger apex.

Four specimens were obtained at Platform Island.

6. SrENoGyRA (OpEas) suBULA, Pfeiffer.

Hab. Cuba, Porto Rico, &e.

The specimens from Fernando Noronha agree exactly with
examples of this species from Porto Rico. They were found

beneath stones on the promontory between Chaloupe Bay and
S. Antonio Bay.

7. Srenoayra (OpEAsS) BECKIANA, var.

Hab. Island of Opara, Peru, Brazil.

The shells from Fernando Noronha, obtained at the same spot
as the preceding species, answer well to Pfeiffer’s description ;
but they are more strongly costulate than Brazilian specimens
with which I have compared them.

ITI. FRESHWATER SPECIES.

1. PLANORBIS NORONHENSIS, sp.n. (Plate XXX. figs. 11-11 8.)

Testa parva, valde compressa, superne in medio depressa,
inferne subplanulata, albida, tenui, subpelJucida; anfractus 4,
superne convexiusculi, striis incrementi tenuissimis sculpti,
inferne radiatim subplicati, ad suturam angustissime marginati,
ultimus infra medium obtuse carinatus ; peristoma tenue, mar-
ginibus callo filiformi junctis, superiore oblique arcuato.

Diam. maj. 5 millim., min. 44, alt. 1.

This species is about the same size and shape as P. Gilberti,
Dunker, and P. fragilis, Brazier, from Australia. The lower
surface, however, is flatter, and the curve of the lip different

Tin.o0c .Jotraw.Zo00n,. Vor. Pt.

_ MFisher del. Mintern imp.

t :

MOLLUSCA OF FERNANDO NORONHA.

ZOOLOGY OF FERNANDO NORONHA. 503

when viewed from above. It was very plentiful in the lake on
the south-west corner of the island.

List of Species obtained at Fernando Noronha by the
‘Challenger’ Expedition.

Acmea, sp. Scalaria hellenica, Forbes.
Littorina nodulosa, d’Orb. Siphonodentalium tetraschistum,
Nerita ascensionis, Gmelin. Watson.
Cerithiopsis, sp. Solarium, sp.
Columbella mercatoria, Linn. Stomatella nigra, Quoy & G.
Cylichna noronyensis, Watson. Utriculus canaliculatus (Say).
Fossarus ambiguus (Linn.). Xenophora corrugata (Reeve).
Marginella sagittata, Hinds. Chiton Boogii, Haddon.
Mitrularia uncinata (Rve.). Pectunculus pectinatus (Gmelin).
Nassa capillaris, Watson. Ervilia subcancellata, Smith.
Oliva fulgida, Reeve. Cardium medium, Linné,
pulchella (?), Duclos. Lucina pecten, Lamarck.
Phasianella, sp. Mytilus exustus, Fve.
Rissoa, sp. Arca imbricata, Brug.

Pecten noronhensis, Smith.

For the above species, see the Reports on the Gasteropoda,
Polyplacophora and Lamellibranchiata, by R. B. Watson, A. C.
Haddon, and E. A. Smith respectively.

EXPLANATION OF PLATE “XXX.

Fig. 1. Triton Ridleyi, sp. n.
2. Littorina trochiformis, var.
3,34. Acmea noronhensis, sp. n.
4-4 b, Siphonaria picta, var.
5. Chiton (Ischnochiton) carribeorun.
5a. Ditto. Central valve, magnified.
6,60. Arca (Acar) Adamsii.
7-7 c. Helix (Ophiogyra?) quinquelirata. 7c. Aperture, enlarged ; lip
broken away to show the teeth.
8. Bulimus (Tomigerus) Ramagei, sp. n.
9. » (Bulimulus) Ridleyi, sp. n.
10. Pupa solitaria, sp.nu. 10a. Aperture, enlarged.
11-11 b. Planorbis noronhensis, sp. n.

504 MR. H. N. RIDLEY ON THE

POLYZOA.

By R. Krrxparrick,
Assistant in Zoological Department, British Museum.

POLYZOA.
The specimens chiefly encrust shells, and are generally much
worn away.
1. AntTEA RECTA, Hincks.
. SYNNOTUM AVICULARE, Pieper.
. SCRUPOCELLARIA FRONDIS, D. sp.
4. CRIBRILINA RADIATA, Moll.

5. SMITTIPORA ANTIQUA, Busk.
(Mollia antiqua, Smitt.)

STEGANOPORELLA Smirtirt, Hincks.
MasticgorHora Durertretr, Audowin.

SCHIZOPORELLA UNICORNIS, Johnston.

LEPRALIA DEPRESSA, Busk.
(Escharella rostrigera, Smitt.)

10. LepraLia CLELIDOSTOMA, Smitt.

11. Ruyncnorora BIsPiInosa, Johnst.—(Encrusting Gorgonia
axis.)

12. CELLEPORA RIDLEYI, n. sp.

13. MicropoRELLA VIOLACEA, Johnst.—Enerusting Gorgonia
axis ; (both purple and white varieties).

14, Crista Hotpswortutir, Busk.

15. AMATHIA BRASILIENSIS, Bush.

Family CELLULARIIDE.
Genus SCRUPOCELLARIA,

SCRUPOCELLARIA FRONDIS, 0. sp.

Zocecia of medium size, alternate ; area oval, occupying nearly
half the front of cell; spines 2-4 on the outer side, 2 on the
inner ; lowermost spine on outer side bending over the top of

ZOOLOGY OF FERNANDO NORONHA. 5U5

the aperture and giving off processes from its upper border ;
operculum entire, large, oval, marked with concentric striz; on

Fig, 1.

Scrupocellaria frondis, n. sp.

some cells a small pointed avicularian cell projecting from the
front of the cell; lateral avicularia wanting; on dorsal surface
vibracula, small, flattened, obliquely placed; sete long. Occia
small, globose, vitreous, punctured.

The presence of the antler-like spine across the top of the area
is a marked character of S. frondis.

Loc. Fernando Noronha; Pernambuco.

Family CELLEPORID®.
Genus CELLEPORA.

CreLLEpoRA RipieyI, n. sp. (Fig. 2, p. 506.)

Zoarium loosely encrusting; zocecia decumbent, rectangular,
and flattened at the margins, heaped, somewhat ventricose sub-
vertical in the centre, separated by raised lines; orifice from
‘semicircular to subquadrate, with concave proximal margin; two
or three short processes surrounding the orifice; at base of an
anterior process a small avicularium facing inwards, with small
semicircular mandible. On the front of some cells a small avicu-
larian cell with small rounded mandible. Ocecium shaped like a
thick semi-disc, concave below, overhanging the mouth of the

LINN. JOURN.— ZOOLOGY, VOL. XX. 39

506 MR. H. N. RIDLEY ON THE

cell; mucronate processes on the upper surface; front wall
of ocecium with a semicircular membranous area on the front

wall.

——-

Cellepora Ridleyi, n. sp.

The ocecium of C. Ridleyi is remarkable in its shape and relations
to the zocecium ; also there is a curious resemblance between the
membranous area and the orifice of the zocecium.

Loc. Fernando Noronha.

CRUSTACEA.

By R. I. Pocock,
Assistant in the Zoological Department, British Museum.

Introductory Remarks.—The fauna is in all essential respects
allied to that of the mainland and of the Antilles. The following
wide-spread forms were, as might have been expected, met with:—
Grapsus maculatus, Letolophus planissimus, Hippa seutellata, » _
Alpheus Edwardsii, and Gonodactylus chiragra. There are two
new species of Alpheus, one of Panulirus, and one of Stenopusculus
(S. spinosus). The last mentioned genus has hitherto only been
known from-the island of Mauritius; its occurrence here, there-
fore, is of great interest. A new freshwater Ostracod was also

obtained.

ZOOLOGY OF FERNANDO NORONHA. 507

DECAPODA.

MAIOIDEA.
Family PeRIcERIDS.

Genus Micropueys, I/.-Hdwards.
1851. Microphrys, M.-Edwards, Ann. Sci. Nat. Zool. 3, xvi. p. 251.
1879. Microphrys, Miers, Journ. Linn. Soc. (Zool.) xiv. p. 664.
1881. Microphrys, A. M.-Edwards, Miss. Sci. Mex. (Crust.) p. 59.

Micropurys Bicornutus (Latreille).

1825. Pisa bicornuta, Latreille, Encycl. Méth. Hist. Nat. x. p. 141.

1872. Microphrys bicornutus, A. M.-Edwards, Nouv. Arch. Mus. Hist.
Nat. viii. p. 247.

1881. Microphrys bicornutus, id. Miss. Sci. Mex. (Crust.) p. 61, pl. xiv.
figs. 2, 3, 4.

Nine specimens, six males and three females (two with ova).

This species is common on the coasts of Florida, Mexico, and
of the West-Indian Islands. Occurred under stones and on
coral-reef.

Genus Mirurax (Leach).
1817. Mithrax (Leach), Latreille, Régne Animal, iii. p. 23.
1834. Mithrax, Milne-Edwards (in pt.), Hist. Nat. Crust. i. p. 317.
1879. Mithrax, Miers, Journ. Linn. Soc. (Zool.) xiv. p. 667.

Mirnrax verrucosvs, W.-Edwards.

1832-38. Mithrax verrucosus, M.-Edwards, Mag. Zool. vii. pl. 4.
1881. Mithrax verrucosus, M.-Edwards, Miss. Sci. Mex. p. 102.

Four specimens, two males and two females (one with ova).
The largest specimen (a male), with the following measurements
of carapace, width 42 mm., length 85 mm., differs considerably
from the others, of which the smallest (the female with ova) gives
the following measurements of carapace :—width 17 mm., length
15 mm. In the three small specimens ail the spines are sharper
and relatively longer, and the carpus of the chelipedes is armed
above with four or five minute spines in addition to the three
spines which adorn its anterior (interior) margin.
Brazil and the West Indies are localities given for this species.
Under stones at Morro do Chapeo.
, 39*

508 MR. H. N. RIDLEY ON THE

Mirnrax (TELEOPHRYS) CRISTULIPES (Stimpson).

1862. Teleophrys cristulipes, Stimpson, Ann. Lyc. Nat. Hist. vii. p. 190
pl. in. fig. 2.

1881. Teleophrys cristulipes, A. JL.-Edwards, Miss. Sci. Mex. (Crus-
tacea), p. 113, pl. xix. fig. 2.

Regions of the carapace defined by shallow sulci. Superior
surface of body and limbs tubercular, inferior surface smooth.

Carapace broader than long, beset with low, inconspicuous,
scattered tubercles which vary in distinctness. The principal
tubercles arranged as follows :—two or three on each half of the
rostrum between the superior orbital prominences, one on each
side of the middle line at the base of the rostrum, two in longi-
tudinal series on each side of the gastric region, several on the
branchial regions, and three on the anterior lateral margin of
each. Orbit furnished in front with a superior and an inferior
blunt prominence. Not furnished with spines or prominences
behind.

Rostrum short, broad, with upturned anterior margin, not deeply
bifid, marked above with central sulcus, and separated on each
side from the superior orbital prominence by a conspicuous de-
pression. Its anterior margin projecting slightly beyond the
middle of the basal segment of the antenne and slightly in front
of the inferior orbital prominence.

Chelipedes large ; merus tubercular above and furnished below
in front with three large, rounded, compressed teeth; carpus
furnished above with four or five tubercles, and with one blunt
tooth in front. Hand smooth; its distal portion compressed
above and below into a crest. Dactylus and pollex meeting only
at the apices. Dactylus furnished with a single tooth.

In the first pair of legs the merus is furnished above with two
longitudinal rows of prominences, the posterior row consisting of
lower rounded tubercles, which distally decrease in size, the an-
terior row of five higher, compressed, sharper teeth, which distally
increase in size; distal margin of the segment produced into five
rounded prominences, varying in size; the carpus, in addition to
three or four low tubercles on its centre, with its distal margin
furnished with a larger anterior and a smaller posterior tubercle,
and its antero-superior surface with a larger proximal and a
smaller distal tooth; propodos furnished above with two tubercles,
one near the centre, the other at its distal margin; claw. long,
curved, hairy below, with its distal portion serrate below.

ZOOLOGY OF FERNANDO NORONHA. 509

The arrangement of tubercles and teeth upon the second, third,
and fourth pairs of legs is nearly the same as the arrangement
upon the first pair, but the posterior row of meral tubercles
becomes progressively fainter from before backwards, and the
teeth of the anterior row become gradually modified in form and
number until, in the posterior pair of limbs, this row is formed
of four teeth, two larger and two smaller, the larger and smaller
alternating, and one of the larger being the most proximal of the
series. Width of carapace 7; mm., length 7 mm.

One male specimen was obtained.

To guide me in the identification of the Fernando-Noronha
specimen, which I refer to 7. cristulipes (Stimps.), I have had to
trust to the descriptions and figures of that species published by
Dr. Stimpson and by M. Alphonse Milne-Edwards, and to my own
examination of a single imperfect individual which was taken off
Cape St. Lucas (California), and presented to the British Museum
by the Smithsonian Institute.

Now, although with the above-mentioned figures and descrip-
tions the specimen from Fernando Noronha does not present
agreement in all points, yet, making allowance for possible errors
on the part of the artists, I should unhesitatingly have referred
this specimen to 7. cristulipes (Stmps.) were it not for the fact
that the points of difference between it and the specimen from
Cape St. Lucas are by no means inconsiderable.

In the Californian specimen the sulci defining the regions of
the carapace are conspicuously deeper, and the tubercles of the
same part, though exhibiting in the main the same arrangement,
are much larger. This is especially the case with regard to those
of the branchial region, the three low tubercles of the antero-
lateral margin in the Noronha specimen being represented
in the Californian specimen by three large upstanding teeth.
Again, with regard to the limbs, the merus of the chelipede in
the Californian specimen is furnished below in front with one
large compressed tooth and the pollex is armed with two
small teeth, these small teeth being scarcely represented in the
Noronha specimen. The other limbs present much the same
arrangement of teeth in the two specimens, but, as in the case of
the carapace, the teeth of the Californian specimen are relatively
larger than those of the Noronha specimen.

J am well aware that the differences thus set forth are ampiy
sufficient to justify the separation as distinct species of the spe-

510 MR. H. N. RIDLEY ON THE

cimens which they characterize; yet having but one example
from each locality, I am unable to determine the constancy of
the differences presented, and must consequently leave the decision
of the question as to the specific identity or distinction of the
two to those whom either the possession of a long series of
forms or a more perfect acquaintance with this group of Crus-
tacea places in a better position to judge than myself.

I am not aware that this species, or at all events any closely
allied form, has before this been recorded from the eastern coast
of America. Stimpson obtained it from Cape St. Lucas and M.
Alphonse Milne-Edwards has described it from the Bay of
Panama.

Mirnrax (Mirnracutvs) coronatus (Herbst).

1782. Cancer coronatus, Herbst, Naturg. der Krabben, i. p. 184, pl. xi.
fig. 63.

ak Mithraculus coronatus, A. Milne-Edwards, Miss. Sci. Mex.

(Crustacea), p. 106, pl. xx. fig. 1.

Eight males and seven females (three with ova) were taken.

This species occurs on the coasts of Brazil, Central America,
and of the West-Indian Islands. Its presence in Fernando No-
ronha has been previously mentioned by Mr. EH. J. Miers, two
small specimens having been obtained from that island during
the voyage of H.M.S. ‘ Challenger.’

CANCROIDEA.
Family CancRIDaZ.

Genus Carpiuius (Leach, MS.), Desmarest.

1825. Carpilius, Desmarest, Consid. gén. sur la classe des Crust., foot-
note, p. 104.

1834. Carpilius, Milne-Edwards (pt.), Hist. Nat. Crust. i. p. 380.

1865. Carpilius, A. Milne-Edwards (pt.), Nouv. Arch. Mus. Hist. Nat.
is p. 212.

1886. Carpilius, EZ. J. Miers, Brachyura of H.M.S. ‘ Challenger,’
p. 110.

CaRPILIUS CORALLINUS (Herbst).

1782. Cancer corallinus, Herbst, Naturg. der Krabben, i. p. 133, pl. v.
fig. 40.

1865. Carpilius corallinus, A. Milne-Edwards, Nouv. Arch. Mus. Hist.
Nat. i. p. 216.

Three specimens, two males and one female.

ZOOLOGY OF FERNANDO NORONHA. 511

This species is the West-Indian representative of the genus.
[These crabs are exported in wooden crates filled with dry leaves
to Pernambuco, where they are in great demand as food. We
were told they were land-crabs.—H. NV. R.]

Genus Actma, de Haan.
1850. Actza, de Haan, Crust. in Siebold, Fauna Japonica, dec. i. p. 18.

ActmA ACANTHA, Milne-Edwards.

1834. Cancer acanthus, M.-Edwards, Hist. Nat. Crust. i. p. 379.

1881. Acteza acantha, A. M.-Edwards, Miss. Sct. Mex. (Crust.) p. 245,
pl. xlui. fig. 1.

One minute specimen, a male, was obtained.

It is only comparatively lately that the locality of this species
has been made known by M. Alphonse Milne-Edwards, who _
received a specimen of it from Guadeloupe.

Genus Leprropivs, A. Milne-Edwards.

1863. Leptodius, A. Milne-Edwards, Ann. Sci. Nat., Zool. sér. 4, xx.
p. 283.

1873. Leptodius, A. Milne-Edwards, Nouv. Arch. Mus. Hist. Nat. ix.
p- 221.

1886. Leptodius, Miers, Brachyura of H.M.S. ‘ Challenger,’ p. 136.

LEPTODIUS AMERICANUS (Saussure).
1858. Chlorodius americanus, H. de Saussure, Mém. sur divers Crust.
nouv. du Mex. et des Antilles, p. 14, pl. i. fig. 5.
-1881. Leptodius americanus, A. Milne-Edwards, Miss. Sci. Mex.
(Crust.) p. 269.
Of this species eight specimens (5 males, 3 females with ova)
were obtained. It is found in the West Indies and Florida.

Genus Lopuactma, A. Milne-Edwards.
1862. Lophactza, A. Milne-Edwards, Ann. Sci. Nat., Zool. sér. 4,
Xvili. p. 43.
1865. Lophactza, A. Milne-Edwards, Nouv. Arch. Mus. Hist. Nat. i,
p- 245.
1886. Lophactza, HE. J. Miers, Brachyura of H.M.S. ‘ Challenger,’
p- 113.

LopHacTMA LOBATA, Milne-Edwards.
1834. Cancer lobatus, Milne-Edwards, Hist. Nat. Crust. i. p. 375.

512 MR. H. N. RIDLEY ON THE

1868, Lophactea lobata, A. Milne-Edwards, Nouv. Arch. Mus. Hist.
Nat. i. p. 249, pl. xvi. fig. 3.

A single male specimen. This is a West-Indian and Mexican
species.

GRAPSOIDEA.

Family OcyPpopDID4.

Genus Ocypona, Fabricius.
1798. Ocypoda, Fabricius (pt.), Ent. Syst. Suppl. p. 347.
1837. Ocypoda, Milne-Edwards, Hist. Nat. Crust. vol. i. p. 41.
1880. Ocypoda, Kingsley, Proc. Acad. Nat. Sci. Philad. p. 179.
1886. Ocypoda, Miers, Brachyura of H.M.S. ‘ Challenger,’ p. 237.

OcYPODA ARENARIA (Catesby).

» 1771. Cancer arenarius, Catesby, Hist. of the Carolinas, ii. p. 35,
pl. xxxv.

1880. Ocypoda arenarius, Kingsley, Proce. Acad. Nat. Sct. Philad.
p. 1&4.

1882. Ocypoda arenarius, Miers, Ann. Mag. Nat. Hist. ser. 5, x. p. 384,
pl. xvii. fig. 7.
Five adult specimens (four males and one female).

[Common in holes in the sand at Sueste Bay and Sambaqui-
chaba.—H. NV. R.]

Family GRAaPSID&.

Genus Grapsus, Lamarck.
1818. Grapsus, Lamarck (pt.), Hist. Nat. Anim. sans Vert. v. p. 247.
1880. Grapsus, Kingsley, Proc. Acad. Nat. Sci. Philad. p. 192.
1886. Grapsus, Miers, Brachyura of H.M.S. ‘ Challenger,’ p. 254.

GRAPSUS MACULATUS (Catesby).

1771. Pagurus maculatus, Catesby, Nat. Hist. Carolinas, ii. p. 36,
pl. xxxvi. fig. 1.

1880. Grapsus maculatus, Kingsley, Proc. Acad. Nat. Sci. Philad.
" p. 192.

This species has a very wide range, occurring upon the coasts
of the warmer temperate and tropical parts of the Indian, Pacific,
and Atlantic Oceans. It is exceedingly variable, and the extent
of variation is well shown by the specimens brought from Fer-
nando Noronha.

Five immature and two adult males were obtained ; the former

ZOOLOGY OF FERNANDO NORONHA. 513

are of a dark green colour with feeble indications of macule, the
latter red-brown with maculee well developed. [Very common on
the rocks all over the group, running briskly just above water-
mark and leaping from stone to stone.—H. NV. R.]

Genus Piaeusia, Latreille.

1806. Plagusia, Latreille (pt.), Gen. Crust. Ins. i. p. 33.

1837. Plagusia, Milne-Edwards, Hist. Nat. Crust. ii. p. 90.

1878. Plagusia, Miers, Ann. Mag. Nat. Hist. ser. 5, i. p. 148.
1886. Plagusia, Miers, Brachyura of H.M.S., ‘ Challenger,’ p. 271.

PLAGUSIA DEPRESSA (Fabricius).

1775. Cancer depressus, Fubricius, Syst. Ent. p. 406.

1782. Cancer squamosus, Herbst, Naturg. der Krabben, i. p. 260, pl. xx.
fig. 113.

1878. Plagusia depressa, Miers, Ann. Mag. Nat. Hist. ser. 5, i. p. 149.

[This ran about on the stones and rocks like the Grapsus.—
EN. BR.)

Genus Lrrotornuus, Miers.
1850. Acanthopus, de Haan, Faun. Japon., Crust., p. 29 (nom. preocc.).
1876. Leiolophus, Miers, Cut. New-Zeal. Crust. p. 46.
1878. Leiolophus, Miers, Ann. Mag. Nat. Hist. ser. 5, i. p. 153.
LEIOLOPHUS PLANISSIMUS (Herbst).

1804. Cancer planissimus, Herbst, Naturg. der Krabben, iu. Heft 4,
p- 3, pl. lix. fig. 3.

1878. Leiolophus planissimus, Miers, Ann. Mag. Nat. Hist. ser. 5, i.
p- 153.

A single specimen (female with ova) of this wide-spread form
was taken.

PORCELLANIDEA.

Family PorceLLaNnip..

Genus PerroxistHEs, Stimpson.
1859. Petrolisthes, Stimpson, Proc. Acad. Nat. Sct. Philad. x. p. 227.

PETROLISTHES MARGINATUS, Stimpson.

1862. Petrolisthes marginatus, Stimpson, Ann. Lyc. Nat. Hist. New
York, vii. p. 74.

I have had no opportunity of examining specimens of P. mar-
ginatus (Stimpson), and consequently not being certain of the

514: MR. H. N. RIDLEY ON THE

correctness of the identification of the specimens that I have re-
ferred to that species, I have thought it desirable to publish a
description of them which may, so far as is possible, furnish a
test as to the accuracy of the conclusion that has been arrived at.

Carapace and upper surface of limbs pubescent. Width of
carapace approximately equal to its length. Carapace smooth,
punctured ; its anterior half furnished laterally with a small, sharp,
upstanding spine. From this spine there extends backwards into
the posterior half of the carapace a granular ridge which serves
to separate the superior portion of the carapace from the lateral
portion. The frons is slightly depressed and is marked off from
the hinder portion of the carapace by a distinct ridge, which runs
transversely between the posterior margins ofthe orbits. Inthe
middle this ridge is interrupted by a conspicuous sulcus, which
extends to the central lobe of the frons. This lobe is rounded
anteriorly ; its lateral margins are nearly vertical to the remainder
of the anterior margin of the frons and approximately parallel to
the superior margin of each orbit, which is the lateral border of
the frons. The anterior half of this lateral border marked off
from the posterior half by being at a conspicuously lower level.

Basal segment of antenna furnished on the inner side with a
small acute spine.

Upper surface of chelipede covered with more or less squami-
form granules ; lower surface smooth. Anterior margin of upper
surface of meral segment produced into asharp process ; beneath
this, on the under surface, is a sharp spine, which may be bifid;
posterior margin of upper surface spined. Anterior margin of
upper surface of carpal segment furnished with three or four
sharpened processes; posterior margin spined and produced
distally into a spined process. The middle of the upper surface
bearing a longitudinal series of larger squamiform tubercles.
Inferior border of anterior surface of carpal segment granular;
rest of the surface smooth. Anterior and posterior margins of
manus and dactylus granular. A slightly curved series of larger
squamiform granules extending along the upper surface of the
mauus from its carpal to the middle of its dactylar joint. Con-
tinuous with this is a series running from the base to the apex
of the dactylus. Apex of dactylus and of thumb smooth and
curved.

Anterior and posterior margins of meral segments of second,

ZOOLOGY OF FERNANDO NORONIIA. 515

third, and fourth pairs of legs spined ; posterior margin of second
and third pairs produced distally into a small acute spine.

Colour (of specimens preserved in spirit of wine) red or yellow
above, with darker spots, reddish pink beneath.

Three specimens. Length and width of carapace in largest
specimen 14 mm.; length of manus and pollex 20 mm.

The specimens that I have here described and identified pro-
visionally as P. marginatus (Stmps.) are evidently closely allied to
P. asiaticus (Leach), the common Indo-Pacific form, and I am
doubtful if they should be regarded other than as varieties of
that species.

HIPPIDEA.
Family Hippips.

Genus Remirzs, Latreille.

1806. Remipes, Latreille, Gen. Crust. Ins. i. p. 45.
1837. Remipes, Milne-Edwards, Hist. Nat. Crust. ii. p. 204.

REMIPES SCUTELLATUS (Fabricius).

1793. Hippa scutellata, Fabricius, Ent. Syst. ii. p. 474.

1858. Remipes cubensis, H. de Saussure, Mém. sur Crust. nouv. du
Mez. et des Antilles, p. 36, pl. ii. fig. 19.

1878. Remipes scutellatus, Miers, Journ. Linn. Soc. (Zool.) xiv. p. 319.

The species occurs on the tropical coasts of the Atlantic.

Twenty-three specimens, two of which are females with ova,
were taken. [Very common on the sandy shores. When a
wave broke, these little crustacea were often seen running and
burying themselves in the sand as the water retired.—H. WV. L.]

THALASSINIDEA.

Family GEBIIDS.

Genus GEBIA.

1816. Gebia, Leach, art. Annulosa, Edinb. Encyel. vii. p. 419.
1837. Gebia, Milne-Edwards, Hist. Nat. Crust. ii. p. 312.

Gepra sPInIGEeRA, S. I. Smith.

1869. Gebia spinigera, Smith, Rep. Peabody Acad. Sci. p. 92.

Right specimens, one female with ova, were brought back.

The species was originally described from specimens obtained
upon the west coast of Central America.

516 MR. H. N. RIDLEY ON THE

ASTACIDEA.
Family PALINURIDE.

Genus Panuirus, Gray.

1847. Panul:rus, Gray, Cat. Brit. Mus. ( Crust.) p. 69.
1852. Panulirus, Dana, Crust. U.S. Expl. Exp. i. p. 519.

Panuirnvus EcHINATUS, S. L. Smith.

1869. Panulirus echinatus, Smith, Trans. Connecticut Acad. i. p. 20.

Five specimens were taken, two adult females, one with ova,
and one immature female, one adult male and one immature
male. ;

The specimens described by Smith were from Pernambuco.

PANULIRUS ORNATUS (Fabr.).

1798. Palinurus ornatus, Fabricius, Ent. Syst. Suppl. p. 400.

1837. Palinurus ornatus, M.-Edwards, Hist. Nat. Crust. 11. p. 296.

1867. Palinurus ornatus, Heller, Reise Freg. Novara, Crust. p. 99.

In1872 v. Martens,in his paper “Ueber cubanische Crustaceen,”
Arch. f. Naturg. xxxvili. p. 128, recorded the occurrence on the
eastern coasts of America of a Palinurus, which he questionably
identified as P. ornatus (Oliv. P), a species which appears to have
its head-quarters in the Indo-Pacific Seas.. From Fernando
Noronha, Mr. Ridley obtained one specimen of a Panulirus,
which I cannot separate by any important character from P. or-
natus (Fabr.); and in addition to this specimen there is in the
British Museum Collection one other from Panama, which is
also, I believe, referable to P. ornatus (Fabr.). It will thus be
seen that this form occurs in the Indo-Pacific Seas and upon the
east and west coasts of America.

It is perhaps of interest to note that the spipes upon the
carapace and upon the peduncles of the antenne appear to be
somewhat sharper, and relatively longer, in the American indi-
viduals than they are in the Eastern individuals that I have had
an opportunity of examining.

[Tolerably common, and collected from the rock-pools for
food.— H. N. FR. ]

PANULIRUS INERMIS, 0. Sp.

Carapace somewhat flattened above, with sides nearly vertical.
The right ard left portions of the upper surface meeting m the

ZOOLOGY OF FERNANDO NORUNILA. 517

middle line at a very obtuse angle. Carapace nearly smooth
frontal spines considerably shorter than the eye-stalks, slightly
incurved at the apices, armed above at the base with a single
Spine; one spine situated near the ocular margin of the carapace,
one in the anterior third of the supero-lateral margin, and a
third beneath the eye-stalk near the outer portion of the basal
antennal segment.

Antennal peduncle about two thirds the length of the
carapace ; basal segment armed externally with a single spine on
its anterior margin; second segment armed above with five spines,
two forming a longitudinal series externally, three forming an
oblique series internally ; third segment armed above with ten
short spines. Below, the three segments are smooth.

Antennular plate nearly horizontal, with rounded antero-
external angles not armed with spines ; the peduncle shorter than
peduncle of antenne ; segments of peduncle not spined.

Epistoma with a straight unspined anterior margin.

The first and fifth pairs of limbs simple, unspined. (Second,
third, and fourth pairs absent.)

Postero-external angles of the sternum prolonged into a sharp,
long spine.

Abdominal tergites smooth, punctured, not marked with a
transverse sulcus; inferiorly and laterally prolonged into a spine.
The posterior margin of the last dorsal plate furnished with two
long, sharp spines.

Proximal portion of telson furnished in the middle of its
upper surface with two spines and with its posterior margin
armed on each side with four spines.

Total length from anterior margin of carapace to posterior
margin of telson 27 millim. Length of upper surface of carapace
11 millim.

One specimen.

Judging from its size, the specimen from which the above
description has been taken is certainly immature. It, never-
theless, presents the characters of a true Panulirus, and differs
from all the specimens of that genus that I have examined in
the absence of spines from the basal plate of the antennule.
Dredged in Water Bay. About 10 fathoms depth.

518 MR. H. N. RIDLEY ON THE

CARIDEA.
Family PALm@MONIDA.
Genus ALPHEUS (Fabricius).

1798. Alpheus, Fabricius, Ent. Syst. Suppl. p. 380.

1878. Alpheus, Kingsley, Bull. U.S. Geol. Surv. iv. p. 189.

AuPHEeus Epwarpsit (Auwd.).

1809. Athanasus Edwardsii, Audouin, Explic. planches de Savigny,
Descript. de ? Egypte, Atlas, pl. x. fig. 1.

1818. Alpheus heterochelis, Say, Journ. Acad. Nat. Sci. Philad.i.p. 243.

1884. Alpheus Edwardsii, Miers, Rep. Crust. H.M.S. ‘ Alert,’ p. 284.

Twenty-nine specimens. This species is common in the
warmer parts of the Atlantic, Pacific, and Indian Oceans, and in
consequence of its wide range and of the variations to which
individuals are subject it possesses a long list of synonyms.

These synonyms may be found upon reference to the above cited
work of Mr. E. J. Miers.

_ALPHEUS MINOR, Say.
1818. Alpheus minus, Say, Journ. Acad. Nat. Sci. i. p. 245.
1837. Alpheus minus, Milne-Edwards, Hist. Nat. Crust. ii. p. 356.
1878. Alpheus minus, Kingsley, Bull.U.S. Geol. Geogr. Surv. iv. p. 190.
One specimen.
This species occurs upon the east and west coasts of America.
Kingsley records it from N. Carolina, Bermudas, Florida on the
east, and from Pearl Island Bay (Panama) on the west.

AupHEusS RIDLEY, n. sp.

Carapace and abdominal tergites smooth; carapace furnished
in front with a short pointed rostrum, which does not nearly
reach to the second segment of the antennular peduncle;
rostrum separated by depression from the ocular hoods, each of
which is furnished with a spine projecting in front as far as
the extremity of rostrum.

Antennular spine reaching nearly to the second segment of
the peduncle, which is the longest of the three, the third being
the shortest.

Antennal scale as long as antennal peduncle, longer than
antennular peduncle; basal segment of antenna furnished be-
neath with a strong spine.

Terminal segment of external maxillipede hairy.

First pair of legs very unequal in size. Dactylus of larger
hand closing vertically, with evenly rounded supero-anterior

ZOOLOGY OF FERNANDO NORONHA. 519

border, without accessory teeth ; its greatest length equal to one
half of the length of the superior margin of the manus. Anterior
margin of the “thumb ” on the inner side nearly vertical, forming
an obtuse angle with the inclined superior margin. Superior
and inferior margins of thumbs on the outer side nearly parallel .
in front united by a distinct anterior border, which below
curving forwards forms with the inferior border the tooth of the
thumb, which does not project so far forwards as the anterior
margin of the dactylus.

Upper margin of the manus with a very faint constriction in
its anterior half; right and left sides smooth, without depressions ;
lower margin with a very faint depression in its anterior half;
upper margin marked with sulcus, which in the middle of the
hand curving downwards and backwards runs to the carpal joint.
Carpus rounded above, not bearing a tooth; meros three-sided,
flattened below, not bearing a tooth above in front. Smaller
manus simple, without constrictions or depressions; dactylus,
thumb, and upper margin of manus approximately equal in
length; carpus furnished with a blunt tooth above, equal in
size to the carpus of the Jarger manus; meros resembling the
meros of the larger manus.

In the second pair of legs the first carpal segment is as long
as the second and the third together; third about half the length
of the second, equal in length to the fourth, shorter than the fifth,
which itself is shorter than the second.

Meros and carpus of third and fourth pairs of legs not spined.

In size and form resembling A. Hdwardsiz, but differing from it
in having the larger hand very lightly constricted above and below.
Moreover, there is a large black spine on each side of the telson.

ALPHEUS PANAMENSIS, Kingsley.

1878. Alpheus panamensis, Kingsley, Bull. U.S. Geol. Surv. iv. p. 192.

Carapace smooth, furnished in front with a strong rostrum,
which projects considerably beyond the spines of the orbital
hoods, almost as far as the second segment of the antennular
peduncle; separated by a depression from the ocular hoods, each
of which is furnished with a small sharp spine.

Lower margin of hood continuous below the spine.

Antennular spine reaching slightly beyond the margin of the
basal segment of the peduncle. Second segment of peduncle
longer than the third, as long as the first.

520 MR. H. N. RIDLEY ON THE

Antennal scale and peduncle as long as each other, and slightly
longer than the antennular peduncle. Basal segment of antenna
furnished below with a strong sharp spine.

First pair of legs very unequal in size. Dactylus of larger
hand closing vertically, its greatest length being more than half
the length of the superior margin of the manus; without acces-
sory teeth.

Anterior margin of the thumb on the inner side nearly

vertical, meeting the inclined superior margin at an obtuse
angle ; less than half the length of the superior margin. Thumb
on the outer side without a vertical anterior margin, the superior
margin meeting the inferior at an acute angle and forming the
tooth. :
Manus smooth, without constrictions or depressions, longer
than the carapace; superior and inferior margins nearly parallel,
Carpus rounded above, not bearing a tooth. Meros three-sided,
flattened below; superior margin produced in front into a blunt
process.

Smaller manus simple; dactylus and thumb approximately
equal in length to each other and to the manus.

Carpus furnished above on the inner side with a small pro-
jection. Meros resembling meros of larger limb, except that the
front process is smaller.

In second pair of limbs the carpal segments are 1, 2 and 5, 3
and 4.

First segment almost as long as the second, third, and fourth
together. Second segment a little shorter than the third and
fourth together, these being approximately equal; fifth as long
as the second.

Meros and carpus of third and fourth pairs of legs not spined.
Dactyli of limbs not bifid.

One specimen. If I am right in referring this species to
Al. panamensis of Kingsley, with the description of which it
agrees well, it is of interest to note that it occurs upon the
eastern and western coasts of America. Mr. Kingsley described
his specimens from Panama and Acajutla.

ALPHEUS OBESO-MANUS, Dana.

1852. Alpheus obesomanus, Dana, U.S. Expl. Exped., Crustacea, i.
p. 547, pl. xxxiv. fig. 7.

Carapace smooth, furnished in front with a short rostrum,

ZOOLOGY OF FERNANDO NORONHA. Vall

which does not reach nearly so far as the anterior border of the
first segment of the antennula, but a little beyond the ocular
hoods, from which it is separated on each side by a deep sulcus.
Ocular hoods not spined, but slightly produced in front.

Antennular spine short, not reaching to the front margin of
the first segment of the peduncle of the antennula. Second
segment of peduncle the longest of the three, the third the
shortest.

Antennal seale as long as peduncle of antennula, shorter than
peduncle of antenna. Basal segment of antenna without a
Spine.

Legs of first pair very unequal in size. Dactylus of larger
manus closing horizontally, about half as long as the upper
margin of the manus. The superior (outer) margin of the
thumb furnished with two large teeth, of which the posterior is
smaller, more slender, and with a blunt apex, the anterior having
a rounded margin.

Dactylus short, rounded. Dactylus and thumb very hairy.

The manus simple, cylindrical, without constrictions or depres-
sions, as long as the carapace and the two proximal segments of
the antennular peduncle. Carpus deep from above downwards,
rounded above, and not furnished with a tooth. Meros deep
from above downwards, three-sided, flattened below; upper
margin produced in front into a conspicuous process.

Smaller hand somewhat resembling the larger, except that it is
less twisted, less cylindrical, with dactylus and thumb straighter
and relatively longer. Carpus less deep, and furnished on the
upper inner margin with a distinct nodule. Meros less deep,
with upper tooth scarcely conspicuous.

In the second pair of legs the carpals are 2, 5, 4 and 3 and 1.
The first, third, and fourth segments approximately equal in
length, the fifth a little longer; the second as long as the third,
fourth, and fifth together. In the third and fourth pairs of legs
the carpus and meros are below furnished in front with a strong
spur.

Ten specimens. So far as I know, this species has not been
hitherto recorded from the American coasts. Its occurrence has
been mentioned in the Samoan Islands. (Atngsley), Fiji Islands
(Dana, Miers), and in Mauritius (Jeich ters).

LINN. JOURN.—ZOOLOGY, VOL. XX. 4.0

522 MR. H. N. RIDLEY ON THE

ALPHEUS ROSTRATIPES, 0. sp.

Carapace smooth, anterior margin crescentically excavated,
the sides of the excavation being formed by the ocular hoods,
which are anteriorly produced but not furnished with spines,
and not separated from the rostrum by a depression. Rostrum —
springing from the centre of the excavation, pointed, short, pro-
jecting slightly in front of the ocular hoods, but not reaching
the anterior margin of the first segment of the antennular
peduncle. Antennular spine reaching to the middle of. the
second segment of peduncle. Segments of peduncle short,
approximately equal in length; second segment furnished ex-
ternally with a small spine on its anterior margin. Antennal
scale as long as the antennular peduncle, much shorter than the
antennal peduncle. Basal segment of antenna furnished laterally
with a conspicuous spine.

One of the legs of the first pair absent. ‘The aoe of the
remaining one (the smaller?) closing vertically; long, longer
than the manus, curved, pointed blade-like, when closed ; crossing
the thumb. Thumb almost as long as dactylus, and at the base
twice as thick, gradually tapering to a sharp, upturned point,
meeting manus at an obtuse angle. Manus elliptical, simple, ©
without constrictions or depressions, furnished close to the
dactylar joint-on each side with two blunt teeth, those on the
outer side being obscurely marked. Carpus furnished on its
inner side with a small blunt tooth. Meros three-sided, flattened
below, superior margin produced in front into a conspicuous
projection. Carpals of the second pair of legs becoming pro-
gressively shorter in the following order :—1, 5, 2, 8, 4.

Carpus and meros of third ‘and: fourth pairs not furnished
below with a spine. Dactylus of fourth pair bifid, of third pair
absent.

ALPHEUS, sp.

(Too mutilated for identification.)

Carapace furnished in front with a small pointed rostrum,
which projects slightly in front of the ocular hoods, but not
nearly to the anterior margin of the basal segment of the
peduncle of the antennula; ocular hoods rounded and not spined.
Basal spine of antennula reaching to the anterior margin of the
basal peduncular segment. Second ‘segment of peduncle longer
than the third, approximately equal to the first. Antennal scale

ZOOLOGY OF FERNANDO NORONHA. 523

longer than the antennular peduncle, shorter than the peduncle
of the antenna. Basal segment of antenna not provided with a
spine.

Legs of first pair absent.

Carpals of the second pair differing upon the two sides, on the
right side the fifth segment being longer than the second, and on
the left side shorter. In each case the first is the longest, and
the third and fourth the shortest. ;

Carpus and meros of third and fourth pairs not produced
below into a tooth.

Dactyli of third, fourth, and fifth pairs simple.

ALPHEUS, sp.

(Too mutilated for identification.)

Carapace furnished in front with a short rostrum, which does

not project as far as the middle of the first segment of the
peduncle of the antennula, and is separated by a depression on
each side from the ocular hoods. Each ocular hood furnished
‘with a spine. Antennular spine short, sharp, not reaching to
the front margin of the basal segment of the peduncle. Second
segment of the peduncle the longest, the first and third approxi-
mately equal in length. Antennal scale as long as antennal
peduncle, a little longer than antennular peduncle. Basal seg-
ment of antenna furnished with a long, sharp spine, which pro-
jects as far as the middle of the second segment of the antennular
peduncle.

First and second pairs of legs absent. Carpus and meros of
third and fourth pairs not produced below in front into a strong
process. Dactyli of third, fourth, and fifth pairs bifid.

[The Alphet were taken in numbers from the holes in which
they hid by breaking up the coral-reef—H. N. R.]

Family PEN xID&.

Genus STENOPUSCULUS, Richters.

1880. Stenopusculus, Richters, Beitridge zur Meeresfauna der Insel Mau-
ritius und der Seychellen, von Mobius, Richters und v. Martens, p. 167.

STENOPUSCULUS SPINOSUS, Nn. sp.
? Syn. Stenopusculus crassimanus, Richters, ¢t. c. p. 168, pl. xviii. figs.
27-29,
Upper portion of cephalothorax sparsely spined ; spines in
4.0*

524 MR. H. N. RIDLEY ON THE

front of the cervical suture larger than those behind it.
Posterior margin of the cervical suture furnished above with
4 or 5 spines, and laterally with 3 or 4 larger ones. Postero-
lateral portions of cephalothorax almost smooth; antero-lateral
portions beset with spines arranged more or less in longitudinal
series. Anterior marginal excavation adjoining the basal an-
tennal segment armed with four spines.

Cephalothorax furnished in front with a pointed rostrum,
which starts upon the anterior half of the cephalic portion of the
carapace and reaches almost as far forwards as the front margin
of the antennular peduncle. Upon each side the rostrum ex-
tends horizontally over the basal portion of the eye. Furnished
above with eleven teeth, of which five are larger than the rest,
and below near the apex with one tooth.

Proximal portion of antennular peduncle furnished externally
with a strong curved spine; upper surface of peduncle with
three spines ; under surface with four on the inner margin and
one on the outer margin. |

Basal segment of antennal peduncle furnished above with two
spines externally, and with a laminate process internally ; second
segment covered by the basal segment, furnished below with
three spines; third segment with one spine externally and with
three internally. External margin of antennal scale with five or
six fine teeth, internal margin fringed; antennal scale somewhat
triangular, laminate, projecting slightly in advance of the
antennal peduncle, which is approximately as long as the anten-
nular peduncle.

Epistome furnished with four strong teeth.

Ischial segment of external maxillipede furnished distally with
three spines externally, and with one spine internally; meral
segment externally with three strong spines. Internal margin
of all the segments clothed with hairs.

Segments of first and second pairs of legs simple, more or less
cylindrical, unspined.

Meropodite of third pair of legs cylindrical, spined, with some
larger sharp spines near the distal extremity on the inner
surface. Carpopodite rounded below, flattened and hollowed
above; the hollowed portion with a few small spines, the rest
thickly spined ; spines on the outer surface larger.

Inner surface of the hand covered with small tubercles; outer

ZOOLOGY OF FERNANDO NORONHA. 525

surface almost smooth, with a few small tubercles near the upper
and under margins. Upper margin compressed into a serrated
keel; under margin also serrated. Anterior margin of hand
nearly at right angles to the axis of the pollex. Pollex upturned
at the apex, furnished on its occludent margin with a tooth
which closes behind the tooth of the dactylopodite. Upper
margin of dactylopodite serrated.

Fourth and fifth pairs of limbs resembling each other in being
slender and elongated, in having the propodite furnished below
with a series of fine spines and consisting of three segments, and
in having the dactylopodite bifid. But whereas the propodite of
the fourth pair consists of five segments, the propodite of the fifth
pair consists of but three. The number of divisions of these
segments, however, appears to vary upon the two sides.

Abdominal tergites smooth above; lateral portions narrowed,
somewhat pointed, and with margins more or less spined.

The outer and inner lamelle of appendages of the sixth
abdominal somite with a median longitudinal crest, serrate
exterior margin, and fringed inner margin. Outer margin of
inner lamella furnished below with a stronger tooth.

Telson with converging lateral margins, rounded posterior
margin; each lateral margin furnished with a central tooth;
posterior margin furnished with three teeth, one on each side

-and one in the middle. Upper surface of telson marked with two

longitudinal crests, each of which bears three spines arranged
longitudinally; the depression between the crests furnished proxi-
mally with four spines in two longitudinal series. Base of
telson bearing on each side one marginal spine.

Two specimens.

Length from apex of rostrum to posterior margin of telson
13 millim.; total length of upper surface of carapace (including
rostrum) 54 millim.; length of manus and pollex of third pair of
feet 8 millim.

This species seems to differ from St. crassimanus, Richters,
in the possession of a greater number of teeth upon the rostrum
and in the absence of a crest upon the abdominal tergites.

The three species which hitherto have, so far as I am aware,
composed the genus were taken at Touquets (Mauritius).

526 MR. H. N. RIDLEY ON THE

STOMATOPODA.

Genus Gonopactyivs, Latreille.

1825. Gonodactylus, Latreille, Encycl. Méth. Hist. Nat. x. p. 473.
1837. Gonodactylus, Milne-Edwards, Hist. Nat. Crust. ii. p. 528.
1880. Gonodactylus, Miers, Ann. Mag. Nat. Hist. v. p. 115.

1886. Gonodactylus, Brooks, Stomatopoda of H.M.S. ‘Challenger,’ p. 55.

GONODACTYLUS CHTRAGRA (Mabricius).

1793. Squilla chiragra, Fabricius, Ent. Syst. ii. p. 513.

1880. Gonodactylus chiragra, Miers, Ann. May. Nat. Hist. v. p. 115.

Fourteen specimens of this widely distributed species were
brought back. In the coral-reef.

MYRIOPODA.

By R. I. Pococg,
Assistant in the Zoological Department, British Museum.

The island does not seem to be rich in members of this group,
since four species only were obtained in it. ‘T'wo of these appear
to be new to science, one being referable to the genus Geophilus,
the other to the genus Spirobolus. The others are the two wide-
spread tropical species, Scolopendra morsitans (Linn.) and Para-
desmus gracilis (C. Koch).

CHITOLO DA.

SCOLOPENDRA MORSITANS (Linn.), emend., Kohlrausch, Arch. f.
Naturg. 1881, p. 104.

Thirteen specimens were taken. Common under dung and
stones, at the east end of the main island and base of Peak

Garden and elsewhere. The bite is about as bad as a wasp’s
sting.

GEOPHILUS RIDLEYI, n. sp.

Length 44 millim. Width about 1 millim. Posterior end of
the body slightly more slender than the anterior.

ZOOLOGY OF FERNANDO NORONHA. 527

Ochraceous, head-plate slightly darker.

Number of pairs of legs 73 (in the female).

Antenne hirsute, the distal end more so than the proximal.
Segments of the proximal half cylindrical, those of the distal half
narrowed proximally ; apical segment as long as the two pre-
ceding segments.

Head-plate with straight anterior margin, rounded lateral
margins, and concave posterior margin; sparsely clothed with
hairs, and almost destitute of punctures.

Frontal lamina coalesced with rest of head-plate.

Basal lamina about twice as wide as long, with abruptly con-
verging lateral margins and concave anterior margin. The pre-
basal lamina visible in the space left between .the concave
posterior border of the head-plate and the concave anterior
border of the basal lamina.

Maxillary sternite wider than long; its anterior margin slightly
excavated, but scarcely bidentate.

Maxillary feet largely visible from above, and projecting
slightly in front of the head-plate ; the segments on the inner
side furnished with hairs but not armed with teeth.

Dorsal plates conspicuously bisulcated.

Ventral pores occupying a circular area in the posterior half
of the sternites.

Legs sparsely clothed with longish hairs.

The anal tergite wide, but not covering the pleure; with
rounded postero-lateral angles, staight posterior margin, and
lateral margins slightly converging behind.

Anal pleure smooth, not furnished with punctures.

Anal sternite very wide at the base, gently converging lateral
borders, rounded posterior angles and straight posterior margin.

Anal pores conspicuous. Anal legs broken.

A single female specimen, found under a stone in the Sapate.

This species appears to be closely allied to G. occidentalis,
Meinert (Proc. Amer. Phil. Soc. xxiii. p. 220), from San
Francisco ; but differs in the absence of teeth from the segments
of the maxillary feet, and in the absence of pores from the anal

pleure.

528 MR. H. N. RIDLEY ON THE

DIPLOPODA.

PARADESMUS GRACILIS (C. Koch).

Two female specimens.

For the synonymy and an excellent description of this species,
see Dr. Robert Latzel’s ‘Die Myriopoden der 6st.-ungar.
Monarchie,’ 1. p. 162.

This very wide-spread form occurs in the East and West Indies
and Brazil.

It has been introduced, in connection with tropical plants,
into Europe; and I have examined many specimens of all ages,
which were captured in the conservatory of Mr. Alfred O.
Walker at Chester, andin the orchid-houses of Mr. Herbert Druce
at St. John’s Wood.

[It was very common in the garden.— H. N. #. |

SPIROBOLUS (s. 8.) NORONHENSIS, 0. sp.

Length about 30 millim. Number of somites 37.

Colour deep slate-grey or almost black; anterior half of each
somite (the first and last excepted) adorned above and below on
each side with a single reddish spot. Legs and labrum reddish.

Distal portion of the head-plate furnished with a faint median
longitudinal impression, upon each side of which, near the
margin of the labrum, are two setiferous punctures, one near the
middle line, the other near the external portion of the labral
excavation. Distal segments of antenne pilose.

First dorsal plate smooth, without striz; laterally, where the
anterior and posterior margins pass into one another, evenly
rounded ; furnished with a fine sulcus which runs from near the
ocular region of the head-plate, close to the antero-inferior -
margin, and terminates at the postero-inferior margin.

Foraminua repugnatoria situated, somewhat dorsally, in the
posterior portion of the somites. Posterior portion of each
somite smooth above; anterior portion marked with transverse
striw ; inferior and lateral portions marked with numerous longi-
tudinal striae. Somites not furnished with the ‘ scobina.’

Posterior somite smooth; produced behind into a blunt
rounded process, which extends slightly beyond the margins of
the anal valves. Anal valves with margins not compressed.
Margin of subanal plate rounded.

ZOOLOGY OF FERNANDO NORONHA. 529

The right and left moieties of the male copulatory apparatus
held together in front by a triangular plate. Below this plate
terminates in a rounded apex, which extends as far as the inferior
margins of the halves of the apparatus. Each upper angle of
this plate produced laterally and upwards into a relatively
slender bar, which curves round the superior portion of the
anterior lamina of its side. Hach anterior lamina simple, more
or less spatulate, with evenly rounded external margin and
slightly concave inferior margin. Viewed from the side, seen
to be considerably thicker above than below. Posterior lamina
irregularly quadrate, with even outer and inner margins which
below slightly converge; the inner margin distally produced
into a conspicuous rounded, noduliform process, which projects
slightly below the level of the concave inferior margin of the
anterior lamina, and is consequently visible when the copulatory
apparatus in its entirety is viewed from the front. Above and
externally, the interspace between the anterior and posterior
lamine is occupied by a small sclerite, with even margins and
rounded below, which, dilating above, forms the posterior margin
of the superior aperture of the sheath, of which the walls are
composed of the four lamin just described, and which contains
the protrusible portion of the copulatory apparatus. This pro-
trusible portion is articulated at its proximal end to a simple
rod, which is itself articulated to the upper extremity of the
anterior lamina. Protrusible portion curved almost through the
are of a semicircle, and composed of two segments. The distal
segment about twice as long as the proximal, membranous and
hollow behind, chitinous in front, with its posterior portion armed
with a simple small process.

A dozen specimens found under stones in the Banana planta-
tions at the base of the Peak.

In many points this species appears to resemble Sp. paraensis
(Humb. & Sauss.). But the absence of all knowledge of the form
of the copulatory apparatus of that species makes it impossible
for me to refer these specimens to it.

530 MR. H. N. RIDLEY ON THE

INSECTA, excepting Coleoptera.

By W. F. Kirsy, F.L.S., F.ES.,
Assistant in the Zoological Department, British Museum.

Notwithstanding the comparatively large proportion of new
species in the present collection, it would be a mistake to suppose
that very many will ultimately prove to be confined to the island
of Fernando Noronha. The greater part belong to Orders of
insects which are comparatively little collected or studied, and
among which large numbers of conspicuous species remain to be
described, even from the best explored tropical countries.

The few Lepidoptera in the collection were taken at an un-
favourable season of the year, and many are worn specimens.
They exhibit more decidedly West-Indian affinities than might
have been expected.

I prefix to the paper a full list of all the species obtained,
except a few which were worn, immature, or too scantily re-
presented for satisfactory identification. They are arranged
systematically under the orders and principal. families to which
they belong.

List of Species obtained.

Order ORTHOPTERA.

ForFIcuLip&. | GRYLLIDE,
1. Pygidicrana notigera, Stal. | 11. Scapteriscus abbreviatus, Scadd,
2. Labidura riparia, Pall. | 12. Gryllus assimilis, Yabr.
3, Anisolabis janeirensis, Dohin. | 13. forticeps, Sauss.
4. Antoni, Dohrn. | 14, Cicanthus (?) pallidocinctus, n. sp.
PHANOPTERID&.
Buatrip&. 15. Conocephalus vernalis, n. sp.
5. Phyllodromia poststriga, Walk. ——-, var, n. frater,
6. Ischnoptera lucida, Walk. 16. Ciucella (n. g.) furcifera, n.-sp.
7. Periplaneta americana, Linn. 17. Meroncidius viridinervis, n. sp.

8. Blatta incommoda, n. sp.
9. Leucopheza surinamensis, Linn.
10. Euthyrrapha pacifiea, Cog. 18. Stenopola dorsalis, Thund.

Locustipz&.

21.
22.

23.

24.
25.

26.

ZOOLOGY OF FERNANDO NORONHA.

531

Order NEUROPTERA.
ODONATA.

LIBELLULID&,

19. Pantala flavescens, Fabr.
20. Tramea basalis, Burm.

Order HYMENOPTERA.

TEREBRANTIA.
ENTOMOPHAGA.
CiHALCIDID.

Blastophaga obscura, n. sp.
Ganosoma dispar, n. sp.

EVANIDs.
Evania levigata, Lat.

ACULEATA.
HETEROGYNA.,
Formicip&.

Camponotus bimaculatus, Smith.
Pheidole omnivora, n. sp.

FossoreEs,
BemBiIcip&.

Monedula signata, Linn.

iw
“I

28,

bo
le}

30,

PoMPILID.£.

. Pompilus nesophilus, un, sp.

LARRID#.

Tachytes inconspicuus, n. sp.

DIPLOPTERA.

VESPID&.

. Polistes Ridleyi, n. sp.

ANTHOPHILA,

ANDRENIDA.

Halictus levipyga, n. sp.

31. —— alternipes, n. sp.

v2. —— atripyga, n. sp.

Order LEPIDOPTERA.

RuoOrPALOCERA.

LyCANID.

. Tarucus Hanno, Séo/l.

HETEROCERA.

Noctvz.

. Heliothis armiger, Hiidn.

. Anomis (?) dispartita, Walk.

. Anthophila flammicincta, Walk.
. Bolina bivittata, Walk.

. Thermesia gemmatalis, Hiibn.

GEOMETRES,

. Nemoria denticularia, Wa/k.
. Acidalia Fara, n. sp.

41,
42.

45.

49,

PyraLegs,

Pyralis manihotalis, Gwén.
Samea castellalis, Guwén.
Hymenia perspectalis, Hiibn.
. Phakellura hyalinata, Linn.
. Margaronia jairusalis, Wads.

. Acharana phzopteralis, Gwén.
. Pachyzancla detritalis, Guén.
. Opsibotys flavidissimalis, Grote.

CRAMBI.

Pitycipip&.

Mella zinckenella, 77ettschke.

532 MR. H. N. RIDLEY ON THE

Order HEMIPTERA.

HETEROPTERA. 52. Hereseus variegatus, n. sp.

Parsee! 53. Ligyrocoris balteatus, S¢,J.

o4. bipunctatus, n. sp.
50, Pentatoma testacea, Dall.
Lycx1pa&, VELIIDE.
51. Lygeus rufoculis, n. sp. 55. Rhagovelia incerta, u. sp.

Order DIPTERA.

DoLicnoPoDID&. Muscip&.
56. Psilopus metallifer, Walk. 58. Sarcophaga calida, Wied.
SyrrHipD&.

57. Temnocera vesiculosa, Fadr.

Description of New Species and Special Notes.

1. Py@rprcraNa NoTIaERA, Stal.
Pygidicrana notigera, St@/, Hugenie’s Resa, Zool. Ins. p. 299 (1858).
Flew into light.

2. LABIDURA RIPARIA (Pall.).
Forficula riparia, Pall. Reise, ii. Anhang, p. 30 (1773).
A cosmopolitan species.

3. ANISOLABIS JANEIRENSIS (Dohri).
Forcinella janeirensis, Dohrn, Stett. ent. Zeit. xxv. p. 285.

4. ANISOLABIS ANTONI (Dohrn).
Forcinella Antoni, Dohrn, Stett. ent. Zeit. xxv. p. 289 (1864).

These earwigs were common under stones in the main island.

5. PHYLLODROMIA PosTstTRIGA (Walk.).
Blatta poststriga, Walk. Cat. Blatt. p. 99, n. 69 (1868).
The locality of the typical specimen is unknown.

6. IscHNOPTERA LUCIDA, Walk.

Ischnoptera lucida, Walk. Cat. Blatt. p. 120, n. 39 (1868).

A single immature specimen, probably belonging to this species.
Taken under stones, base of Peak.

7. PERIPLANETA AMERICANA (Linn.).
Blatta americana, Linn. Syst. Nat. 1. p. 424, n. 4 (1758).
A cosmopolitan species. Common and introduced.

ZOOLOGY OF FERNANDO NORONHA. 5338

8. BLaATrA INCOMMODA, 0. sp.

Long. corp. 113 millim.

Female. FKerruginous brown; the thorax and sides of the
abdomen varied with black; legs and costal margin of the
tegmina testaceous. Pronotum rather long, moderately convex,
the sides converging in front, the hinder angles rounded off, and
the hind border convex. Tegmina broad, covering the whole
base of the abdomen, but ceasing at about two fifths of its
length.

Similar to B. orientalis, Linn., but much smaller, the tegmina
much larger, and the pronotum longer.

9. LEUCOPHHA SURINAMENSIS (Linn.).
Blatta surinamensis, Linn. Syst. Nat. i. p. 424, n. 3 (1758).
A cosmopolitan species. Under stones, base of Peak.

10. EurHyRRAPHA PACIFICA (Coq.).
Blatta pacifica, Coqueb. Illustr. Ins. i. p. 91, pl. xxi. f. 1 (1801).

11. ScapreriIscUS ABBREVIATUS, Scudd.

Scapteriscus abbreviatus, Scudd. Mem. Peabody Acad. Sci. i. p. 14, t. i.
ff. 8, 20 (1869).

Larve found in burrows in the sand under a Conferva (Entero-
morpha), in salt water, on the shore of San Antonio Bay, a little
above high water. Perfect insect in and about the yards of the
house.

12. GRYLLUS ASSIMILIS (Fubr.).

Acheta assimilis, Fabr. Syst. Ent. p. 280, n. 3 (1775).

A species widely distributed in America. This is the black
cricket mentioned by Webster and other visitors to the island.
It is very common in the central district on the paths, and
makes a great noise, especially about 4 o’clock in the afternoon.

18. GRYLLUS FoRTICEPS, Sauss.
Gryllus forticeps, Sauss. Miss, Sci. Mex. vi. p. 407 (1870).

14, GHCANTHUS (?) PALLIDOCINCTUS, U. sp.

Long. corp. 13 millim.

Male. Reddish brown, abdomen, hind knees, and hind tarsi
darker; head rather flattened, the palpi and the outside of the
scape of the long and slender antenne towards the base whitish.
Pronotum longer than broad, sides subparallel, more shining and
paler on the lower lateral border than above ; elytra brown, reti-
culated with darker nervures, about two fifths as long as the abdo-

5384 MR. H. N. RIDLEY ON THE

men, which is rather long and cylindrical, the incisions conspicu-
ously pale; cerci broken, but the remaining portions are bordered
on each side with very long and fine hairs. Legs short and
stout, and slightly compressed; all the femora enlarged, the
middle ones least so; hind tibiz spined from the base, with 3
large terminal spines on each side, and 3 or 4 larger spines alter-
nating with smaller ones beyond the middle ; first joint of tarsi
with 2 small and 1 large spine on each side.

Probably belongs to a new genus of Gcanthide, but tia, a
superficial resemblance to Gryllodes.

15, CoNOCEPHALUS VERNALIS, 0. sp.

Exp. al. circa 78 millim.

Bright grass-green ; wings hyaline, with bright green nervures ;
fastigium short, obtusely rounded, as in C. triops, Linn.; four front
femora unarmed. Male with face, antenne, eyes, and the whole of
the fastigium whitish ; tips of mandibles and palpi red ; tibie paler
than the ground-colour ; auditory apparatus and tarsi beneath
blackish ; tegmina yellowish along the costa, a buff streak at the
base above the subcostal nervure, and a white basal streak on the
left side above the median nervure. Female much less varied with
white; labrum white, mandibles more yellowish, auditory apparatus
and tarsi whitish, the latter brown below; ovipositor a little longer
than the abdomen, extending far beyond the knees, but shorter
than the closed wings; ovipositor paler towards the extremity,
aud slightly veined with reddish, the extreme tips of the blades
black.

This grasshopper was very common everywhere in the Main
and Rat Island. It makes a great noise at night. The brown
form, which flew about with it, was not so common.

Var. FRATER.

Exp. al. 75-83 millim.

Brown; the thorax apparently rather longer and narrower, an
effect which is caused by an indistinct pale or blackish line on
each side; frequently a pale streak behind each eye; mandibles
varied with pale reddish, the extreme tips black; tegmina
greyish brown, often with dark speckles above and below the
principal nervures and on the costa beyond the middle (similar
speckles are visible on one of the green specimens) ; wings hyaline,
with greyish-brown nervures. .

ZOOLOGY OF FERNANDO NORONHA. 535

Genus CicELLA, n. gen.

Affinities doubtful; nearest to Bargilis in the structure of
the legs, but more nearly approaching Elimea in neuration.

Vertex slightly convex, shaped nearly like an equilateral tri-
angle when viewed from above, the fastigium projecting about as
far as the scapes of the antenne, when the latter are recurved.
Eyes large, round, prominent. Antenne slender, filiform, at least
aslongasthe tegmina. Pronotum above flat, oblong, slightly nar-
rowed in front and rounded behind, excised laterally at the base
of the wings. Tegmina rather narrow, especially at the tips, but
hardly pointed ; shorter than the wings, the drums of the male
triangular, vitreous, nearly alike on both tegmina. Front legs
moderately stout, unarmed ; the tibie much swollen at the base
to receive the linear foramina, which are well marked on both
sides. Tarsi of all the legs similar, of nearly equal size, and about
as long as broad, the second joint lobate, the last very slender,
and nearly as long as the two preceding joints together. Middle
tibize with a few very short and small spines towards the extremity.
Hind legs long and slender, the basal half of the femora mode-
rately thickened, furrowed on the side, and carinated below;
tibiz with a double row of short and slender spines, and with two
small spines at the tip. Cerci of the male nearly as long as the
short abdomen, with long terminal forks, the longest slender and
almost sickle-shaped at the tip ; subgenital laminz short, concave
at the extremity. Ovipositor nearly as long as the abdomen,
moderately broad, strongly compressed, turned upwards, and
pointed at the tip.

16. GQicELLA FURCIFERA, 0. sp.

Exp. tegm. 38 millim.; exp. al. 42 millim.

Head, thorax, tegmina, tips and veins of wings, and greater
part of legs grass-green ; antennex and tibiew, abdomen and appen-
dages yellowish, a yellowish line on each side of the pronotum
above ; wings hyaline.

Hab. Pernambuco and Fernando Noronha. In all the speci-
mens from the latter locality the green colour has more or less
faded to yellowish brown. When fresh this insect is bright

green. It was common on Main Island, and especially on Rat
Island.

536 MR. H. N. RIDLEY ON THE

17. MERONCIDIUS VIRIDINERVIS, 0. Sp.

Exp. al. 65 millim.; long. corp. 38 millim.

Male. Brown, head smooth; labrum greenish; mandibles
black, except at the base; a depressed circle surrounding the
space of the antenne, and the fastigium projecting in a spoon-
shape between them ; scape pointed; flagellum broken: the basal
joints varied with lighter and darker brown; thorax strongly
granulated, a little speckled with black, and much raised behind,
where it assumes aslight greenish tint; tegmina brown, minutely
reticulated and spotted with dark brown, chiefly above and
below the nervures ; longitudinal nervures mostly green; in the
costal area the nervures are blackish towards the base, where
they anastomose a little; on the disk the transverse nervures are
brown or indistinctly green ; inner margin with alternate darker
and paler spaces: wings smoky hyaline, with reddish-brown
longitudinal and brown transverse nervures; hind margins
damaged, but probably browner than the rest of the wing; legs
indistinctly mottled; spines of femora mostly black on the
inner sides, hind femora with a black basal streak on the outside.

Somewhat resembles MW. indistinctus, Walk., but the wings
are shorter. :

A single specimen on a tree in the Sapate.

18. STENOPOLA DORSALIS (ZLhwnb.).

Truxalis dorsalis, Thunb. Nov. Acta Upsal. ix. p. 80 (1827).

Stenopola dorsalis, St@/, Recensio Orth. 1. p. 83 (1873).

The hind legs have not been described; they are reddish
brown, the middle of the femora being black on both surfaces,
the striations more or less marked with paler. The hind tibie
are armed, except on the basal third, with a double row of mode-
rately long and pointed spines, the intermediate space above is
clothed with long fine white hairs, and there is a row of much
shorter white hairs on the under surface also. The sides and
under surface of the hind tibie are generally dark green or
blackish ; at the tip there are two short spines on the outside,
and two long ones on the inside. There are apparently only
three joints to the hind tarsi: the first is three times as long
as broad, but is broad and flattened; the second is much ~
narrower, half as long again as broad, and produced into a long
tooth at the extremity beneath, and the terminal joint is very

ZOOLOGY OF FERNANDO NORONHA. 537

slender at the base, gradually enlarging to the pulvillus, and
about as long as the two preceding joints together.

The species appears to be common, and differs considerably in
size, the tegmina expanding from 30-40 millim., and the body
measuring from 18-20 millim. in length.

On both Main and Rat islands, but especially common on the
latter.

19. PANTALA FLAVESCENS (fabr.).

Libellula flavescens, Fabr. Ent. Syst. Suppl. p. 285 (1798).

Distribution. World-wide.

Very common everywhere on Main Island. The larve living
in puddles in the central district.

20. TRAMHA BASALIS (Burm.).

Libellula basalis, Burm. Handb. Ent. u. p. 852, n. 25 (1839).

Less common than the preceding. The abdomen when fresh
is dark crimson-red. A well-known South-American species.

21. BLaAsTOPHAGA OBSCURA, 0. sp.

Male. Long. corp. 2 millim. 7

Brown or yellowish brown, smooth, except afew short hairs
on the tarsi. Front tarsi apparently 3-jointed, middle and hind
tarsi 5-jointed; tarsal claws very strong, and front and hind
tibize ending in short strong spines.

This species resembles the description of B. brasiliensis, Mayr,
from Blumenau, but is considerably larger. I think it useless to
give a detailed description, for which a larger series, including
both sexes, and preserved in different ways, would be desirable.
The locality will probably serve to fix the species, especially as
true Blastophaga does not appear to be well represented in
America.

22. GANOSOMA DISPAR, L. sp.

Male. Long. corp. 14 millim.

Yellow, smooth; head forming a long oval, broad behind,
gradually narrowed in front, antenne inserted widely apart ;
legs of nearly equal size and structure, femora slightly thickened ;
tibie spinose on the outer edge, and terminating in a coronet of
short spines, none of which are conspicuously longer than the
rest; first joint of the tarsi longer than thick ; abdomen long,
tapering.

LINN. JOURN.—ZOOLOGY VOL. XX. 4]

538 : MR. H. N. RIDLEY ON THE

Differs from Ganosoma attenuatum, Mayr (¢ ), in not being
depressed, and (perhaps) in the long abdomen ; and from Tetra-
gonaspis gracilicornis, Mayr (2), in the much shorter joints of
the antenne.

Female. Long. corp. 2 millim.; ovipositor 44 millim.

Tawny-yellow, with a slight greenish-coppery reflexion (colours
perhaps altered by spirit) ; antenne 12-jointed, serrated and set
with very short hairs; brown, except the two basal joints;
scape as long as the three following joints, second joint rather
longer than the fourth, third (annulus) very small, fifth and
following gradually smaller, the last three joints forming a club ;
ovipositor more than twice as long as the body; veins of the
wings of nearly uniform thickness ; ulua as long as the ptero-
stigma, hardly curved or thickened, metacarpus about as long as
the radius. Head and thorax finely rugose. Legs yellow, the
femora slightly thickened.

Appears to approach most nearly Tetragonaspis* flavicollis,
Mayr, but that species has two annuli (ring-joints) to the
antenne. Except in the structure of the antenne, the single
specimen before me much resembles the figure of 7. gracilicornis,
Mayr, but the latter species has longer hairs on the antenne,
and the terminal joints do not form a club, to say nothing of
other differences.

23. Evanta Lmzviaata, Latr.

Evania levigata, Latr. Gen. Crust. Ins. iii. p. 251 (1807).

A cosmopolitan species. The larve of this genus are parasitic
in the egg-capsules of Blattide.

A single specimen taken in a house at Sambaquichaba. It is
very common in Pernambuco.

24. CAMPONOTUS BIMACULATUS (Smith). .

Formica bimaculata, Smith, Cat. Hym. B. M. vi. p. 50, n. 171 (1858).
Six specimens, all small workers. The species is new to the
Museum collection. Smith described it from St. Vincent’s.
Roger (Berl. ent. Zeitschr. vi. p. 285, 1862) identifies this species
with Fogmica ruficeps, Fabr. (Syst. Piez. p. 404, n. 32); but I am
not satisfied that this is correct, as Fabricius does not mention

the conspicuous pale spots on the second segment of the

abdomen. In the small workers the head is mostly black;

* This genus proved to be the female of Ganosoma.

,
:

ZOOLOGY OF FERNANDO NORONHA. 539

in the large workers it is red. Dr. Mayr records this species
from New Granada, and it is probably widely distributed in
South America.

Under the wood of a Burra tree in the Sapate.

25. PHEIDOLE OMNIVORA, 0. sp.

Soldier 43 millim.; worker 23-3 millim. in length.

Soldier red, shining (abdomen darker), sparingly covered with
raised white hairs. Head, without the mandibles, about as
broad as long, finely and sparingly longitudinally striated above
and in the middle, where it is depressed, behind; sides gradually
rounded behind the eyes. Mandibles very broad, hardly punc-
tured, blackish at the base and tips. Scape of the antenne
straight, gradually thickened beyond the middle, nearly one
third of the length of the antenne. Second joint about three
times as long as broad, joints 3-9 very slightly longer than broad,
joint 10 much thicker and twice as long as broad, joint 11 rather
shorter and thicker, and joint 12 forming a long pointed cone;
the hair on the antenne is thicker and closer than on any other
part of the body. Mesonotum with a hump on each side above,
and finely punctured above ; metanotum, which is armed with two
strong spines, more closely ; first node of the petiole with the
spine somewhat truncated at the extremity ; second node fully
twice as broad as the first, and with strongly projecting lateral
angles; abdomen finely punctured at the base.

Worker similar, but smaller and generally darker, the antennz
lighter ; the scape nearly as long as the remainder of the antenna,
slightly curved, but hardly thickened, and the terminal joint
thicker in proportion and less pointed than in the soldier. The
spines on the metanotum and on the first node of the petiole are
much shorter,-and the projecting sides of the second node are
rounded off.

This species much resembles Phetdole pusilla, Heer, in size
and general appearance, but differs widely in structure.

This was exceedingly abundant in the houses, making nests in
the earth between the bricks of the floor. It is very destructive.
devouring all kinds of food, and even ate up the insects we
captured, in the chip-boxes.

41*

540 MR. H. N. RIDLEY ON THE

26. Moneputa stenata (Linn.).

Vespa signata, Linn. Syst Nat. ed. x. 1. p. 574, n. 14 (1758).

A common South-American species.

Very common on the sand-hills, where it makes its burrow.

27. PoMPILUS NESOPHILUS, 0. sp.

Long. corp. 10 millim.; exp. al. 15 millim.

Female. Dull black, first three segments of abdomen, the
sides of the fourth, and the under surface of the hind tibie red.
Wings smoky, a little lighter and subhyaline on the hind wings
and towards the base of the fore wings.

Head large, eyes hardly extending to the base of the jaws,
face rather more strongly punctured than the vertex; clypeus
short, slightly emarginate ; labrum short, transverse ; eyes nearly
parallel, front ocellus forming the apex of a rectangle with the
hinder ocelli, the two latter rather wider apart than the space
between these ocelli and the eyes; second joint of the antenne
half as long again as the third. Pronotum falling in front to the
neck in a short rounded curve, rather broader than the meso- or
metanotum ; the metanotum is furnished with a small pro-
minence on each side near the base.

Second and third cubital cells of nearly equal size, the second
recurrent nervure striking the third cubital cell in the middle.

Spines of the legs as usual.

Much resembles P. sobrinus, Blanch., a Chilian species, in which,
however, the thorax is verdigris-green above, instead of black.

Taken flying over paths in the centre of the island, not common
and difficult to catch.

28. TACHYTES INCONSPICUUS, . 8p.

Long. corp. 6-8 millim. |

Black, clothed with a very fine silvery pile (that on the upper
part of the face with aslight golden appearance in certain lights),
otherwise most conspicuous on the sides of the abdomen, towards
the extremities of the segments ; thorax and scutellum shining,
with very numerous small punctures, not very close together:
metathorax rugosely-punctate; tegule testaceous; wings clear
hyaline, with a strong violet iridescence ; nervures dark brown.

Closely allied to 7 iridipennis, Smith, from Hga, but in that
species the thorax is longitudinally striated, and the tips of the
tarsi are ferruginous.

ZOOLOGY OF FERNANDO NORONHA. 541

29. PotistEs RIDLEY, n. sp.

Long. corp. 15-16 millim.; exp. al. 26-20 millim.

Worker. Varied with ferruginous red, dark brown, black, and
yellow ; clothed with very fine silky pubescence. Head red,
the vertex darker, the face and orbits lighter, base of the head
black ; antenne black in the middle above. Thorax black, the
prothorax and mesothorax red above, the hinder edge of the
prothorax paler, and the front edge narrowly bordered with
yellow ; the edges and central line of the mesothorax more or
less bordered with black above; on the pleura, beneath the
fore wings, is a conspicuous yellow spot; scutellum and post-
scutellum edged in front with yellow, the band on the former
hardly complete in the middle, and the space behind it more or
less red; metathorax finely and transversely striated, a deep
longitudinal channel in the middle, more strongly striated, and
edged with a yellow stripe on each side; on each side, above the
base of the hind cox, is another yellow spot. Abdomen dark
brown, finely pubescent, shading into black at the base, and
generally more or less red at the extremity. Legs red, coxe
and femora black, knees red or yellow, hind tibie more or less
black in the middle. Wings smoky hyaline, strongly tinged with
ferruginous along the costa of the fore wings; tegule ferru-
ginous.

Var. a. Face, head beneath, pectus, and coxe beneath yellow ;
femora striped beneath with yellow; first two segments of the
abdomen with a small yellow spot on each side above, and a
larger one near the base of the first segment beneath.

Very nearly allied to P. instabilis, Sauss., from Mexico; but
this is a redder insect, with the segments of the abdomen always
more or less bordered with yellow.

This insect is called here ‘‘ Marimboudo,” and is very common.
It makes its nest on the underside of an overhanging rock or
eaves of a house, or on the branch of atree. I have seen a
Cashew-nut tree containing an immense number of nests in
various stages of construction. The nest consists of a single
comb of cells of a triangular or oval outline, and attached by a
pedicel at the narrow end; a large one is about four inches in
length, and three across in the broadest part. The cells are
about three quarters of an inch deep, and a quarter of an inch
across. The insect stings slightly, but only when much irritated.

542 MR. H. N. RIDLEY ON THE
It plays a very important part in the fertilization of the flowers,
especially the Cucurbitacee.

30. HALIorus LEVIPYGA, n. sp.

Female. Long. corp. 10 millim.; exp. al. 16 millim.

Head and thorax dark green, slightly bronzed; abdomen
shining, shading into violet at the extremity of the segments ;
wings hyaline; legs black, clothed with pale hairs.

Head and face finely and closely punctured, sparingly studded
with whitish hairs, dark green, occasionally shading into bronzy
or violet in certain lights; clypeus green or violet-black, with
much larger and fewer punctures than the upper part of the
head; the extremity and the labrum black, the lower mouth-
parts dull bronzy green ; antenne black, ferruginous towards the
extremity beneath; thorax dark green, thickly punctured, most
finely on the prothorax, which shades into bronzy ; metanotum
and base of abdomen above densely clothed with whitish hairs ;
metanotum with longitudinally diverging strie in front, behind
deeply and longitudinally concave, with small punctures, the
intermediate spaces still more finely punctured; tegule tes-
taceous ; wings hyaline, with brown nervures and a slight violet
iridescence ; pectus dull green, densely clothed with whitish
hairs; propectus violet-black; legs black, with whitish hairs
above, and the tibiz and tarsi densely clothed with red hairs
beneath ; the knees (very narrowly), spines, and last joint of the
tarsi mostly reddish. Abdomen smooth and shining, the punc-
tuation being extremely fine, even under the microscope, and
densely clothed with short white hairs, except towards the
extremity of the first two segments; the segments greenish in
front and violet-black behind ; towards the extremity the lateral
bristles are longer; under surface of abdomen brown, the ex-
tremities of the segments slightly reddish.

31. HaLICcTUS ALTERNIPES, 0. sp.

3 2. Long. corp. 11 millim. ; exp. al. 138-14 millim.

Head and thorax dark green above ; abdomen green, smooth
and shining, the neighbourhood of the incisions more or less
violet ; wings hyaline, or slightly clouded in and below the radial
cell, and towards the extremity of the discoidal cell of the hind
wings ; legs reddish, the femora and tibie in the middle, and often
the last joint of the tarsi dark green or blackish ; elypeus tipped

ZOOLOGY OF FERNANDO NORONHA. 543

with tawny, and with more numerous punctures than in
Hi, levipyga; the punctures of the mesothorax and the lower
part of the metathorax larger; the channel of the metathorax
narrower; antenne black above, ferruginous below; cheeks
bronzed.

Closely allied to H. levipyga, but differs in colour and
punctuation. Appears to be a very common species.

32. HALICTUS ATRIPYGA, D. sp.

3 2. Long. corp. 5 millim.

Head and thorax clothed with whitish hairs, green,.very finely
punctured; head much bronzed; clypeus rather short, the tip
black ; metanotum longitudinally striated in front; abdomen
bronzy black, more violet in the incisions ; antenne black, ferru-
ginous beneath; head and thorax dull bronzy green beneath ;
abdomen beneath shading more into ferruginous brown; wings
hyaline, with brown nervures ; tegule pitchy; legs ferruginous,
the femora, and sometimes more or less of the tibie, blackish.

A small species, not closely allied to any other, and easily
recognizable by the strong bronzy colouring of the back of the
head.

These Halicti were taken in the flowers of the Melons and the
Oxalis, and play an important part in the fertilization of the
flowers in the island.

33. Tarucus Hanno, Stoll.

Papilio Hanno, Stoll, Suppl. Cram. pl. xxxix. ff. 2, 2 B (1790).

Rusticus adolescens Hanno, Hiibn. Sammi. ex. Schmett. i.

Many specimens, taken between July 25 and August 31.

The Brazilian form of this widely distributed and variable
species. It is generally larger than the more northern form of
the species, and stands in the British-Museum collection under
the MS. name of 7. monops, Zell. The specimens expand from
174, to 23 millim.

This little Blue was very common on Rat and Main islands.
The Catachrysops trifracta, Butl., said to have been caught on
Rat Island by the ‘ Challenger ’ Expedition, we did not meet with ;
and it seems possible that there was some mistake in the labelling
of this specimen, as the genus is only known from the East
Indies.

544 MR. H. N. RIDLEY ON THE
34. HEnIoTHIs ARMIGER (Hubn.).
Noctua armigera, Hiibn. Eur. Schmett., Noct. f. 370.
A. cosmopolitan species.

A single specimen was caught flying in the daytime on East
Hills.

35. ANOMIS (?) DISPARTITA, Walk.

Anomis (?) dispartita, Walk. Cat. Lep. Het. xiii. p. 990, n. 8 (1857).

Described by Walker from Jamaica and St. Domingo; the
specimens from Fernando Noronha most nearly resemble that
‘ obtained in the former locality.
Taken on the wing at night. Main Island.

386. ANTHOPHILA FLAMMICINCTA, Walk.
Anthophila flammicincta, Walk. Cat. Lep. Het. xxxii. p. 801 (1865).
The types are from St. Domingo.

37. Bonina BrviTrata, Walk.

Bolina bivittata, Walk. Cat. Lep. Het, xiii. p. 1156, n. 23 (1857).

A common species. The specimens in the British Museum
are from Honduras, Venezuela, St. Domingo, and Jamaica. The
single specimen from Fernando Noronha has a narrower band
than any of the others; but it would require a series to show
whether this character is constant in the locality, or a mere
individual variation.

38. THERMESIA GEMMATALIS (Hiibn.).

Antisarsia gemmatalis, Hiibn. Zutr. ex. Schmett, i. p. 26, n. 77, ff. 153,
154 (1818).

Common among the wild beans on Main and Rat Islands
(August 17-25).

09. NEMORIA DENTICULARIA, Walk.

Nemoria (?) denticularia, Walk. Cat. Lep. Het. xxii. p. 536, n. 20 (1861).

The locality of the type specimen is unknown; but it is pro-
bably a wide-ranging species, as there are specimens in the
British Museum from Corrientes and Goya.

40. AcIDALIA Fara, n. sp.

Exp. al. 16 millim.

Male. Grey, thickly dusted with brown, the first line com-
mencing at about one third of the length of the wing with a dark
shade curving to a conspicuous black spot, thence curving sharply
inwardly till it terminates in a more conspicuous dark acute
angle projecting outwardly just above the inner margin; itis not

ZOOLOGY OF FERNANDO NORONHA. 545

continued on the hind wings. The second line commences at two
thirds of the length of the wing in another dark shade on the
costa, which runs obliquely to another black spot, below which it
is continued in grey festoons across both pairs of wings, forming
more than a half-circle on the basal side of a conspicuous black
spot on the hind wings. The third line is the most conspicuous,
and is formed ofa series of black spots connected by a grey line
on the fore wings, which curves inwards twice, a little above the
middle, and again above the inner margin. On the hind wings it
forms a nearly continuous black line, curving outwards at one -
fourth of the distance from the costa, and again, in a wider and
more regular curve, below the middle. At the base of the fringes
runs a row of black dashes, and between these and the third line
are a series of indistinct greyish markings. Underside grey-white,
with the central spots indicated, but not conspicuous. Head
black above; body and legs grey; abdomen rather indistinctly
banded with black above, and with a blackish space at the base of
the anal tuft; femora tawny beneath, front femora and tibie
blackish above; antenne tawny, alternating with whitish above.

Allied to A. retractaria from Florida, but darker and more
distinctly marked.

41. PyRALIS MANIHOTALIS, Guén.

Pyralis manihotalis, Guen. Spéc. Gén. et Lép., Delt. et Pyr. p. 121
(1854).
Described by Guénée from Cayenne.

42. SAMEA CASTELLALIS, Guén.
Samea castellalis, Guén. Spéc. Gén. Lép., Delt. et Pyral. p. 195 (1854).
A widely distributed species throughout America.

43. HYMENIA PERSPECTALIS (Hubn.).
Pyralis perspectalis, Hubn. Eur. Schmett., Pyr. f. 101.
A widely-distributed species.

44. PHAKELLURA HYALINATA (Linn.).

Phalzna-Geometra hyalinata, Linn. Syst. Nat. i. (2) p. 874, n. 279
(1767).

This little moth was very common on the Main Island, especially
about the Peak, among the wild melons, &c. (August 17-20).

A cosmopolitan species.

45. MarGARONIA JAIRUSALIS, Walk.
Margaronia jairusalis, Walk. Cat. Lep. Het. xviii. p. 524, n. 15 (1859),
Originally described from Venezuela.

546 MR. H. N. RIDLEY ON THE

46. ACHARANA PH HOPTERALIS (Guwén.).
Botys pheopteralis, Guén. Spéc. Gén. Lép., Delt. et Pyr. p. 349 (1854).

A common and widely-distributed species in Tropical America.

47. PACHYZANCLA DETRITATIS (Guén.).
Botys detritalis, Guén. Spéc.Gén. Lép., Delt. & Pyr. p. 347 (1854).

Widely distributed in South America.

48. OPSIBOTYS FLAVIDISSIMALIS (Grote).

Botis flavidissimalis, Grote, Canad. Ent. ix. p. 105 (1877).

Described by Grote from Texas; but there are also specimens
from South America in the British Museum.

49. MELLA ZINCKENELLA (Treitschke).

Phycis zinckenella, Treitschke, Schmett. Eur. ix. (1) p. 201 (1832).

Many specimens. A cosmopolitan species. ‘There are spe-
cimens in the British Museum from S. Europe, 8. Africa, and
Australia. It is a somewhat variable insect, but is easily
recognized by the broad pale costa and the huge palpi.

50. PENTATOMA TESTACEA, Dall.

Pentatoma testacea, Dall. Cat. Hem. p. 250, n. 43 (1851).

Several specimens were met with in various spots in the Main
and Rat Islands.

51. Ly@Us RUFOCULIS, 0. sp.

Long. corp. 9-11 millim,

Bright red, including the eyes and ocelli; head above, between
and in front of -ocelli, blackish ; antenne and legs blackish and
setose ; coxee reddish at base, corium more or less varied with
blackish, outer edge narrowly black, membrane brown, blackish
in 2. Head and thorax rugose, the former truncate and
bicarinate in front. Rostrum black, extending to the extremity of
the posterior coxe. Corium fully developed in 6; in @ about
half as long as the abdomen, and rather pointed.

An extremely common species, allied to L. anticus, Walk., in
which, however, the head and thorax are unicolorous red, and the
tegmina are reddish brown.

This highly-coloured Bug occurred in large numbers upon the
ground in the Sapate under bushes of Jacguinia; but was local,
only found in a few spots.

ZOOLOGY OF FERNANDO NORONHA. 547

52. HEREUS VARIEGATUS, 0. sp.

Long. corp. 7 millim.

Head and front of pronotum dull red; posterior lobe of pro-
notum testaceous, with very large separated punctures. Corium
blackish, with rows of punctures; outer edge of corium testaceous,
with two black spots, one at its extremity, the other a little
before ; the veins, a large triangular spot on the inside, and three
smaller spots between this and the margins are also testaceous ;
scutellum with a converging testaceous line on each side; mem-
brane blackish, with a testaceous blotch at base andtip. Antenne
finely pubescent, the two basal joints testaceous, the third
biackish, the fourth black, with the base white. Legs testaceous,
front and hind femora and all the tibize strongly spined. Body
~ red beneath, pectus darker. Rostrum testaceous, extending to
the base of the hind coxe.

Allied to H. percultus, Dist.

Taken at light.

53. LigyrocoriIs BALTEATUS, Stal.

Ligyrocoris balteatus, Stal, Vet.-Akad. Handl. (2) xii. p- 146, no, 4
(1874).

Flew into light.

54. LIGYROCORIS BIPUNCTATUS, N. sp.

Long. corp. 4 millim.

Head, pronotum, and scutellum black ; hinder lobe of pronotum
with two short, reddish, longitudinal stripes in the middle, a red-
dish spot on each side in front and one at each angle behind;
antenne testaceous, the last jomt black. Corium testaceous,
with rows of black depressed punctures in the middle; the veins
on the inner edge black, running into a large irrregular apical
black border ; the outermost vein broadens out just before reach-
ing it and encloses a very conspicuous oval white spot ; membrane
greenish, with two brown, diverging, curving lines in the middle,
two brown spots at the base, and one on each side of the curve.
Under surface of the body apparently black; legs testaceous,
femora more or less black ; but the specimen is carded in such a
manner as not to allow of a proper examination of the under
surface or of the legs and rostrum.

548 MR. H. N. RIDLEY ON THE

55. RHAGOVELIA INCERTA, D. sp.

Long. corp, 24 millim., lat. 1 millim.

Blackish brown ; the orbits, front of prothorax, whitish, slightly
tawny; abdomen with a whitish pile on the sides and less con-
spicuous beneath ; base of antenne, femora above, and legs beneath
tawny ; closed hemilytra whitish (or slightly tawny) at the base
between the nervures, but with no other pale markings except
the conspicuous long-oval white spot before the tip.

A shorter and broader species than any at present represented
in the British Museum ; but with most general resemblance to
Microvelia vagans, White. It is possibly a variety of the widely
distributed and variable Velia collaris, Burm.

On grasses in the lake.

56. PsILopus METALLIFER, Walk.
Psilopus metallifer, Walk. List Dipt. B. M. iii. p. 647 (1849).

Flying over Cucurbitacez on the Peak.

57. TEMNOCERA VESICULOSA (Fubr.).
Syrphus vesiculosus, Fabr. Syst. Anil. p. 226, n. 11 (1805).

Flying over herbage in the sun, and also taken on the flowers
of the cabbage in the garden.

58. SARCOPHAGA CALIDA, Wiedem.
Sarcophaga calida, Wiedem. Aussereurop. zweifl. Ins. ii. p. 366, n.-24
(1830) ; Walk. Ins. Saund. Dipt. p. 326 (1856).

COLEOPTERA.

By Cuas. O. Watrrnovusse, F.E.S.,
Assistant in the Zoological Department, British Museum.

The number of species obtained was 24. As might naturally
be expected, many of them are Brazilian species or are species
with a wide geographical range. One of the Heteromerous
genera, which I have named sthetus, is almost certainly
indigenous. Perhaps the most interesting species is a Longicorn
of the genus Acanthoderes. At first sight I took it to be a pale

ZOOLOGY OF FERNANDO NORONHA. 549

variety of A. yaspidea, a common Brazilian species, but on closer
examination the apex of the elytra was found to be different. If
this species should hereafter prove to be peculiar to the island, it
will be a somewhat remarkable fact.

A few of the species the determination of which would have
been attended with great labour, I have left unnamed, as I feel
sure they are introduced species.

BRACHELYTRA.
BuLonucuus, sp.
A single specimen, closely resembling B. formosus, Gray., of
Brazil, but smaller, 24 lines in length. It was found in a decom-
posing rat.

TROGOPHL@US, sp.

A single example, closely resembling 7. pusillus, Grav., of Eu-
rope, but a trifle larger, with the two basal impressions of the
thorax strongly marked.

NECROPHAGA.

DERMESTES FELINUS, Fabr.
A widely-distributed species.

EPuURamA ?

A single specimen of a very small species (17 millim.) somewhat
resembling Hpurea limbala, but a little narrower, with the mar-
gins not expanded, light brown, finely pubescent, closely and
very finely punctured; each elytron having two fine, widely
separated, impressed striz. Abdomen covered by the elytra.

PADPICORNI A.

PHILHYDRUS MARGINELLUS, Fabr.
A widely-distributed species.

DAOTYLOSTERNUM ABDOMINALE, Habr.
A widely distributed species.

LAMELLICORNTA.
ATENIUS, sp.
Three examples belonging to two species. No doubt intro-
duced.

550 MR. H. N. RIDLEY ON THE

STRATEGUS ANTAUS, Fabr. |

The prothorax and elytra of this North aud Central- American
species. [Its larvee were found under rubbish in the Sapate with
the fragments of the perfect insect, which we did not succeed in
taking alive.—H. N. R.]

SERRIOCORNIA.

HYPporRHAGUS MARGINATUS, Fabr.
A single example of this West-Iudian species in the flowers of
an acacia in the garden.

MAaLACODERMATA.

XYLOPERTHA, Sp.

Three specimens. Probably an introduced species. Four milli-
metres in length; reddish yellow, shining, with the rough ante-
rior part of the thorax brown and the apex of the elytra pitchy.
The elytra finely punctured at the base, strongly punctured pos-—
teriorly ; the posterior declivity also strongly punctured, with
three nodes on its upper margin on each elytron.

[These came to light in our rooms.—H. N. R.]

HETEROMERA:

Eprrracus Baresi, Maklin.

Elongatus, ellipticus, modice convexus, parum nitidus, parce
flavo-pubescens, crebre punctatus ; elytris striato-punctatis, striis
postice obsoletis. .

Long. 73 millim. |

The head is closely and rather strongly punctured. The thorax
two fifths broader than long, broadest a little behind the middle,
very slightly narrowed behind, a little more in front; aistinetly
and moderately strongly punctured ; the punctures on the disk
separated from each other by one or one and a half times the
diameter of the punctures; the punctures towards the sides
larger and closer together, giving a slight rough appearance ; the
punctures at the anterior margin are finer. The elytra are at
the base a little wider than the base of the thorax, very slightly
widened to rather behind the middle and then arcuately narrowed
to the apex; moderately finely but distinctly punctured, the

ZOOLOGY OF FERNANDO NORONHA. Bod

punctures not very close together; with several short lines of
rather larger punctures, which are most distinct towards the
margin. Antenne and legs pitchy.

Numerous examples of this species from the Amazons are
labelled in Mr. F. Bates’s collection with the name “ H. Batesii,
Makl.,” but the species does not appear to be described.

Briapstinus RIpLEYI, n. sp.

Elongatus, oblongo-ovatus, parum nitidus, fusco-brunneus,
flavo-pubescens ; thorace crebre evidenter punctato ; elytris
punctato-striatis, interstitiis lateralibus et ad apicem convexius-
culis, subtiliter vix crebre punctatis; antennis, tibiis tarsisque
piceis.

Long. 5 millim.

Antenne with the third joint elongate, about one fourth longer
than the second, the fourth a trifle shorter than the third, the
fifth, sixth, and seventh about as long as broad, the eighth, ninth,
and tenth slightly transverse. The head is moderately strongly
punctured, but the punctures are not very close together; the
epistome is moderately emarginate, rather more closely punctured
than the head. The thorax is evenly convex, broadest at posterior
angles, scarcely sinuate at the sides, narrowed at the anterior
third ; moderately strongly punctured, the intervals between the
punctures about equal to the diameter of the punctures; the
anterior angles moderately prominent and acute; the base rather
strongly bisinuate. The elytra a little broader than the thorax ;
somewhat strongly punctate-striate; the striz near the suture
scarcely impressed at the base ; the punctures in the striz mode-
rately large and close together ; the punctures on the interstices
are fine but distinct, the spaces between them being about once
and a half the diameter of the punctures.

I have ventured to describe this species as it appears to be new,
although probably introduced.

BLAPSTINUS, sp.

Several specimens of a species closely resembling B. pulveru-
lentus, Esch., but with the strize of the elytra more impressed.
There are several North-American species closely allied to this
with which I am unacquainted, and it is not improbable that it is
referable to one of them.

MR. H. N. RIDLEY ON THE

Le) |
On
bo

AESTHETUS, l. gen.

General characters of Cyrtosoma. Mentum small, narrowed at
the base, truncate in front; ligula somewhat round; labial palpi
short and stout, the apical joint rather large, ovate. Labrum
nearly twice as broad as long, nearly straight in front, the angles
rounded. Head transversely impressed between the eyes. Epi-
stome not separated from the forehead by a distinct line, consider-
ably advanced in front of the insertion of the antenne, obliquely
narrowed in front. Thorax evenly convex, the sides gently arcuate.
Scutellum very small, short triangular. Elytra oblong-ovate, very
convex, but somewhat flattened dorsally; their epipleura very
broad and flat, gradually narrowed to the apex of the elytra.
Wings absent. Prosternal process considerably produced pos-
teriorly, acuminate, horizontal. Mesosternum sloping, slightly
concave. Metasternum very short; intermediate and posterior
coxal cavities separated by a very narrow space. Antenne mo-
derately long and slender, the third to seventh joints elongate,
the apical joints a trifle broader. All the tibie slightly curved.

ARSTHETUS TUBERCULATUS, 0. sp.

Piceo-niger, nitidus ; capite crebre sat fortiter punctato, epi-
stomo convexo, subtiliter punctulato ; labro piceo-flavo ; thorace
creberrime punctato, latera versus tenuiter ruguloso; elytris
opacis, fortiter striatis, striis impunctatis, interstitiis sat convexis,
singulis serie tuberculorum minutorum instructis ; antennis, palpis
tarsisque piceis, tibiarum apice intus, tarsisque subtus fulvo-
pilosis.

Long. 9-11 millim.

The antenne are somewhat slender; the second joint scarcely
longer than broad, the third three times as long as the second ;
the fourth to seventh joints elongate, each a trifle shorter and
broader at its apex than the previous one; the eighth, ninth, and
tenth joints pilose, broader and shorter than the seventh, but
not transverse ; the eleventh oval. The thorax is evenly convex,
very gently arcuate at the base, emarginate in front, moderately
rounded at the sides, finely margined all round (except at the
middle of the front margin), the posterior angles are very slightly
projecting, the anterior angles slightly obtuse. The punctures on
the disk are close and distinct, at the base and at the sides they
are very fine and obscure. On each side of the disk the surface

ZOOLOGY OF FERNANDO NORONHA. 553

_is finely longitudinally rugulose, but this is very slight in the

larger examples. The interstices of the elytra are closely and
very finely punctured, each interstice having a line of rather
closely placed, minute, shining tubercles. Epipleura of the elytra
dull. Under flanks of the prothorax dull and closely longitudi-
nally striated. Sterna and abdomen shining, finely punctured.

The two smaller examples have the thorax relatively narrower
than in the larger examples, and the rugulose surface more marked
and more extended. These differences are no doubt sexual.

These were found under stones and bark in the woods of the
Sapate.

CoPIDITA, sp.

Several examples of a species which I am unable to determine.
Yellowish, usually with slight grey shade on theelytra. Length
6-7 millim.

Those captured were attracted by a light.

aac HTD. A.

Brucuus Porosus, Sharp.

Two imperfect male specimens, which may be referable to B.
porosus, Sharp (Biol. Cent.-Amer., Coleopt. v. p. 490), from Guate-
mala, the type of which (unique) is a female. The brown colour
is rather more suffused over the elytra, and the punctures are not
quite so large. The pectinations of the antenue are very long.

RHYNOCHOPHORA.

SITOPHILUS ORYZA, L.
Introduced. [It is very destructive to the maize-grains, so that
in the store-rooms the maize is covered with a thick layer of sand

- to prevent their attacks.—H. N. &.]

XYLOPHAGA.
Tomicus ?
Two immature specimens belonging to this or an allied genus.
Very pale yellow ; 13 millim. in length.

PYONARTHRUM ? SETULOSUM, 0. Sp.

Oblongo-ovatum, brunneum, sat nitidum, convexum ; thorace
latitudine paulo breviore, convexo, postice paulo angustiore, ante

LINN. JOURN.— ZOOLOGY, VOL. Xx. 42

554 MR. H. N. RIDLEY ON THE

medium oblique angustato, creberrime subtiliter punctalato ;
elytris thoracis basi perparum latioribus, subtiliter striato-punc~
tatis, interstitiis parce subtilius punctatis, parce pubescentibus,
seriatim squamulato-setosis ; antennis pedibusque sordide tes-
taceis.

Long. 2 millim.

Head distinctly visible from above ; concave in front in one sex.
Eyes coarsely granular, widely separated above, but very slightly
separated below. Antenne testaceous ; funiculus 6-jointed (or
possibly 7)*; the first large, subglobose; the following joints
very short and transverse, gradually increasing in width; club
large, 3-jointed, oval, pubescent. The thorax has a well-defined
margin separating the under flanks. The surface (seen through
a microscope) is finely coriaceous, moderately finely punctured,
the intervals between the punctures about equal to the diameter
of the punctures; sparsely pubescent, the hairs at the front
margin slightly thickened. The strie of the elytra are lightly
impressed, but scarcely so on the disk; the punctures in the striz
moderately fine and close together, the punctures on the inter-
stices rather smaller and moderately widely separated. Anterior
tibie rather broad, with four or five small obtuse teeth on the
outer side, and two larger ones, one at one third io the apex,
the other apical. ‘Tarsi slender.

This insect agrees in the majority of its characters with Pyc-
narthrum gracile, Kichh. (Mém. Soc. R. d. Sci. Liege, viii. 1878,
p- 104). The anterior tibize are, however, evidently different :
“tibise anteriores apice extus rotundate.” The structure of the
antennee appears to be the same, but the club is ovate and not
acuminate. The elytra are punctate-striate and not crenate-
striate, and the punctures are round and not subquadrate, &e.

It appears to be related to Cnestnus, Horn, but the anterior _

cox are not so widely separated. ~ “7

[It was bred from the bark of the endemic fig-tree, from a
specimen out of the garden of the Residency.—H. NV. £.]

PLATYPUS PARALLELUS, Ff.
. Two examples of this Brazilian species,

* The joints after the first are so confused that even with the antenna
mounted in balsam I am not quite certain of their number.

f
>

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«
F
:
-
-

Cr
Cr
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ZOOLOGY OF FERNANDO NORONHA,

LONGICORNIA.

ACANTHODERES RIDLEY, n. sp.

Latus, depressus, omnino albo-griseo-pubescens ; elytris pone
medium macula laterali inconspicua ornatis ; apice mucro brevi
instructo. :

Long. 113-16 millim.

Form and general appearance of A. jaspicea, Germ., but a little
more depressed and with the elytra rather more obtuse at the
apex. The colour is pale whitish grey, generally with some sandy-
yellow shade on the base of the antenna, disk of the thorax, and
on parts of the elytra. The front of the head is paler, with some
conspicuous black punctures, especially between the eyes. The
thorax has the usual median raised line and slight swelling on
each side of the disk ; there is no black at the sides; there is a
line of very distinct black punctures along the basal margin, and
a similar (but less regular) line along the front margin, and there
are some other punctures scattered over the surface. The elytra
have the usual costa distinct, slightly sinuous as in A. jaspdea.
On the shoulder a few small tubercles may be traced through the
pubescence. Some examples have scarcely any trace of spots,
but most have a not very conspicuous pale fuscous spot behind
the middle near the side, and behind this there are generally
numerous black punctures which are surrounded by a brown
shade; usually a short oblique brown line may be seen at the
apex of the costa. Apical mucro shorter and less acute than in
A. jaspidea. Abdomen with a slight grey shade in the middle,
and a line of black dots on each side. Tibiz unicolorous, or with
a very slight pale brown spot near the apex. Some examples
have a slight oblique brown spot on the.disk, rather before the
middle.

[These flew into light in the evening, and were very plentiful.
They made a loud squeaking noise when caught.—H. JN. PR. |

TRYPANIDIUS ISOLATUS, n. sp.
Dense pallide griseo-pubescens ; thorace guttis minutis non-
nullis ornato ; elytris nigro-punctatis.
Long. 138-15 millim.
Closely allied to 7. dimidiatus, Th., but relatively a little nar-
rower and quite differently coloured. The pubescence is very
42*

556 ‘MR. H. N. RIDLEY ON THE

pale grey, some parts being a trifle paler than others; and there
is a slight mixture of pale yellowish-brown pubescence, especially
on the elytra. On the underside the pubescence is more sandy
yellow, leaving the middle of the sterna and abdomen dark. The
thorax. has the usual line of large punctures at the base; a very
slight raised line behind the middle of the disk; the lateral tooth
small and acute. The elytra have the very slight costa near the
suture a little less raised at the base than in 7. dimidiatus, and
the apical truncature of each elytron is not straight but has the
angles rounded. The black punctures are arranged as in 7. di-
midiatus, but extend to the apex. There is a small pale spot at
the base close to the scutellum, and a scarcely noticeable brown
spot near the suture a little way from the base. One example
has a slight whitish mark on the suture just before the middle,
and a moderately broad whitish band near the apex, somewhat
similar to the band in 7. dimidiatus, but broader nearer the apex
and less angular near the suture, where it is only carried up toa
level with its origin on the margin.

This species is, in many respects, intermediate between 7 di-
midiatus aud T. melancholicus.

[These also came to light with the preceding, but were rarer,
only two being taken.— Hl. WV. &.]

PsEUDOTRIMERA.,

ScYMNUS, sp.

Two examples of a species resembling the Indian §. weranipe-
linus, Muls., but a trifle smaller (13 millim.) ; uniform brownish
yellow, with golden pubescence ; punctuation of the elytra close,
fine but distinct ; metasternum very strongly punctured.

THYSANURA and COLLEMBOLA.
By H. N. Ripuzy, M.A., F.LS.
Iapyx Saussuru, Humbert, Rev. et Mag. Zool. Sept. 1868,
p. 351, pl. ii. figs. 1-5.
A single specimen of what I believe to be a young example of
this little animal was obtained under a stone in the Sapate woods.

ee ae

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fw

ZOOLOGY OF FERNANDO NORONHA,. 557

It corresponds closely to the figure given by Humbert, excepting
that it is only 10 millim. in length instead of 22 millim., and
that the antenne are as hispid as those of I. solifuga, Halliday.
In Humbert’s figure the antenne are quite glabrous, but as he
does not mention this among the differences between his species
and J. solifuga, it is possibly an error of the draughtsman. The
number of joints in the antenne is almost equal to that of
I. Saussurit, and more than that of J. solifuga; but the animal
is but little larger than the type specimens of J. solifuga in the
British Museum. Another point of difference, though very
slight, is in the forceps. In both the above quoted species there
is a secondary tooth on the inner edge of each chela, besides
several smaller rounded papille; now in the specimen from
Fernando Noronha this secondary tooth is nearer the apex
of the chela than is either of the others.

Distribution. The distribution of the whole genus is very little
known at present, as specimens are not often collected. J. soli-
fuga occurs in South Europe, Algeria, and Madeira (the var.
Wollaston). I. Saussurii was obtained in Mexico at Orizaba.
Species are also recorded, but not described, from the United
States and Calcutta.

Dr. Grassi, in p. 1 of “ Progenitori degli Insetti e dei Miria-
podi,” gives J. Saussurit as from Brazil; but does not say whether
he has seen Brazilian specimens; and in his list of species mentions
it merely from Mexico, evidently using “ Brasili”’ as a synonym
for Mexico. It is more than probable that, if sought for, it will
be found to occur also on the mainland of Brazil.

LepismMa LEat, n. sp.

Though it was to be expected that house-inhabiting Lepismas
would occur here, I sought for them in vain until just as we were
leaving, when a single large Lepisma, perhaps disturbed by the
packing-operations, appeared. The specimen was somewhat
damaged in capturing it; but as it seems to be undescribed, and
is a very curious animal, I describe it as it is.

Corpus 19 millim. longum (setis exclusis), griseum. Caput
parvum, subrotundatum, Antenne filiformes (fracte). Oculi
minimi, nigri, pone basin antennarum positi. Prothorax 3 mm.
longus, margine superiore recto, inferiore excavato, marginibus
lateralibus productis; mesothorax et metathorax similes sed
breviores. Pedes coxis valde crassis, brevibus, oblongo-ovali bus,

558 MR. H. N. RIDLEY ON THE

subtus duabus setis longis; articulis secundis longioribus multo
tenuioribus pubescentibus, supra spina crassa armatis; tertiis
tenuioribus, equilongis, subtus setiferis ; tarsis multo brevioribus.
setiferis, uncis parvis duobus terminalis. Inter coxas tres
squame, ovales, obtuse, quam coxas minores. Abdominis
segmenta subequalia, glabra. Segmentum ultimum breve. In
medio penultimi segmenti appendices due breves, complanate,
acute; post eas duz laterales, breves, teretes, hispid, tune
duz longs multo longiores et tenues hispide, tunc due longe
graciles hispide, et in medio appendix longissima unica, crassior,
setosa, articulata et annulata.

The abdomen was filled with some bright green substance,
which was emitted from the mouth when touched—apparently
ereen paint nibbled off the shutters.

The most nearly allied species to this which I have seen was
obtained in Socotra, and is now in the British Museum. The
breadth of the thorax is greater than in Z. saccharina, but the
head is not concealed as in some species.

LEPISMA CORTICOLA, N. sp.

Parva, 1 cm. longa, angusta, metallica plumbea, dorso arcuato
nec complanato. Caput parvum, rotundatum ; oculiad basin an-
tennarum, rubri. Antenne graciles, annulate, hispide; articulus
basalis maximus, reliqui breves, plurimi, crassiusculi. Palpi
maxillares 5-articulati, articulis breviusculis. Palpi labiales
breves, clavati; articulus basalis brevis sectus, secundus longior,
tertius brevissimus conicus, quartus rotundatus brevis. Thorax
aneustus, quam abdomen vix latior. Prothorax quam meso-
thorax longior, metathorax brevior, marginibus omnium ciliis
rigidis munitis. Pedes longiusculi, hispidi; cox breves, late
nec crass; secundo paullo angustior et brevior; tarsi longius-
culi, trivnguiculati; squama inter coxas prothoracis ovata acuta
magna, altere minores. Abdomen breviusculum; appendices
segmenti ultimi graciles, hispid, breviuscule, subeequales.

In rotten wood and under stones in the Sapate and the’ base
of the Peak. It also occurred on the mainland at Pernambuco
in similar localities.

The chief peculiarity of this Lepisma is its very rounded back,
resembling that of a Machilis rather than that of a typical
Lepisma. It is a small active species occurring singly, of a dark
leaden-grey colour. The scales resemble those of Z. saecharina
in outline, but are more notched at the upper edge, and seem

Pee ea eT eee

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ZOOLOGY OF FERNANDO NORONHA. 559

also to have more numerous ridges. The thorax is not much
broader than the abdomen, and the margin does not extend much
beyond the feet.

Macuitis, sp.

A single specimen of a very small brown Machilis was taken
under a stone at the base of the Peak; but, by an accident, the
specimen was destroyed, and we met with no others.

SEIRA MUSARUM, 0. sp.

Minuta, gregaria, in vita metallica, 1 mm.longa. Caput rotun-
datum, hispidum; oculi in maculis nigris fascia obscura spe
connectente. Antenne breviuscule, violacesx, hispide; articulo
basali brevissimo, secundo et tertio subsequali basali duplo longi-
oribus; articulo quarto triunciali, longissimo. Collum distine-
tum. Segmentum secundum corporis (prothorax) latum, mar-
ginibus rotundatum ; tertium brevius, quartum multo brevius,
quintum longius, sextum quinto subzequale, septimum sexto triplo
longius, terminalia brevia. Corpus in speciem ferme glabrum,
insquamosum, flavescens, segmentorum basibus et marginibus
ceruleo-purpureis. Pedes hispidi, primi breviusculi, secundi
longiores, tertii longissimi, graciles.

Very abundant between the wet bases of the petioles of the
bananas, at the base of the Peak.

It is very nearly allied to 8. Buskii, Lubb., which was de-
scribed from specimens found ina hot-house in England, and pro-
bably introduced with tropical plants. It differs in the absence
of hairs round the neck and on the body, longer hind legs, and
also in coloration. The spring resembles that of S. Buskii, and
is rather hispid. The neck is very distinct. These small Col-
lembola have been much neglected by collectors, and it is most
probable that this species was introduced in the bananas.

ECHINODERMATA*.
There were not many species of Echinoderms found on the
island, but the following were obtained :—
CIDARIS TRIBULOIDES, Lamk.
Very plentiful on the north side of the island in coral-recf
pools, near Sambaquichaba and Morro do Chapeo.
* The species were determined for me by Prof. Jeffrey Bell.

560 MR. H. N. RIDLEY ON THE

DIADEMA SAXATILE, L.
Two specimens from pools at Morro do Chapeo.

TRIPNEUSTES ESCULENTUS, Leske.
Very plentiful in rock-pools in Sponge Bay, sometimes almost
filling a small pool. .«

OpHIURA CINEREA, I. & Tr.
Common under stones, north side.

OPHIACANTHA sp.
A very small specimen with the preceding.

OPHIOCOMA PUMILA, Luth.
Young specimens.

OPHIOCOMA ECHINATA, 4g.
A single specimen from Portuguese Bay. Quite unknown to
our guide, so it is probably rare here.

OLIGOCH ATA.
By W. Buaxtanp Benuam, D.Sc.

On February 2, 1889, I received a tube of small worms, which
had been collected by Mr. H. N. Ridley in the island of Fer-
nando Noronha, with the request that I would identify them.
I gladly undertook to do so, and obtained permission to open
the worms, if necessary ; for it is now admitted that in most
cases it is almost impossible to pronounce with certainty on the
genus of an Earthworm from external characters alone; and
although in some cases external characters may point to some
particular genus, yet it is not always safe to rest content with
such an indication, and we must examine the internal anatomy
in order to be sure of the point.

The tube which I received contained six small, ill-preserved
worms, one of which was a Polychete, which I did not further
examine. Of the remaining five, the first (which I will call A)
was of rather an earthy-brown colour and measured 4 inches
in length; the second and ,third were similar in colour and
general appearance, but were only about 2 inches long; the
fourth (B) was of a much darker tint, and was reddish brown in

ZOOLOGY OF FERNANDO NORONHEA. 561

colour, somewhat like that of Luwmbricus terrestris ; it measured
52 inches ; the fifth turned out to be merely the anterior portion
of a similar specimen.

The Worm A.—The first feature which struck me was the
quincuncial arrangement of the set in the posterior region of the
body ; anteriorly the sete are in couples.

The body-wall, being somewhat transparent, allowed me to
distinguish through it paired light-coloured bodies, or “ pyriform
sacs,’ lying in the ventral region posteriorly. The most
anterior somite of the body is very elongate, and carries the
mouth terminally, the prostomium being absent. These features
recalled the genus Urocheta of Perrier; but somewhat similar
characters are found in other Earthworms: thus the pyriform
sacs have been described by myself in Urobenus* ; and the scat-
tered condition of the sete, though not identical with the arrange-
ment noticed, closely resembled, and might easily be confounded
with, what obtains in Diacheta (Benham).

Turning then to the clitellum, I found it to cover the somites
XIV. to XXII. or XXIII.; it is not complete on the ventral surface ;
and both anteriorly and posteriorly is, as is often the case,
more feebly developed. One peculiar feature, however, about
the clitellum, which therefore recalled Urocheta, is the fact
that the intersegmental grooves are deep and noticeable; the
glandular structure not being continuous from somite to somite,
as is the case in most other EKarthworms.

I could see no pores, or external apertures, of the genital ducts
or nephridia; I therefore opened the worm, in the ordinary way,
by a median dorsal incision, in order to satisfy my suspicion as
to its belonging to the genus Urocheta.

The septa are thin, with the exception of four situated anteriorly,
which are greatly thickened, namely those forming the posterior
wall of somites VI., VII., VirI., and x. (the septum between Ix. and x.
is absent). Such thickened muscular septa are not unusual in
Earthworms; but whether their position is constant in a given
species is by no means certain. Perrier has not helped us to
settle the matter, since in his figure be represents only four such
septa, whereas in the text he speaks of five of them. However,
Beddard ft, in a species of this genus from Australia, describes four,
having the same position as in the worm under consideration.

* Quart. Journ. Micr. Sci. vol. xxvii.
t Proce. Roy. Soc, Edinb. xiv. 1887, p. 160.

562 MR. H. N. RIDLEY ON THE

The alimentary tract, the vascular system, the nephridia, all
exhibit the characters peculiar to or present in Urocheta.

There is but a single pair of seminal reservoirs, which have a
greater extent than in U. corethrura (F. Miiller *); for in the
present specimen that lying on the left side passes through eight
somites, that of the right side passes through twelve somites,
commencing in somite xII., where are situated also the ciliated
rosettes.

I was unable to trace the sperm-ducts ; I could find no ovary ;
I did not look for testes, as this would have necessitated some
damage to the worm, which I was anxious to injure as little as
possible.

There are three pairs of spermathece ; each is a very elongate,
thin-walled sac, enlarged distally, and lying respectively in somites
vit., vi1.,1x. The chief difference between the two species of
Urocheta that have received names lies in the different position
of the spermathece. In U. corethrura they lie in somites
VIIl., 1x., x.; in U. dubia (Horst) they are found in somites
VI., Vil., viit.; in Beddard’s specimen from Australia they
have the same position as in the present specimen.

Such is a brief sketch of the anatomy of the worm A, from
Fernando Noronha; it is sufficient, however, to identify it’ as
belonging to the genus Urocheta; but as to the species—
whether it belongs to any of those already described or
requires a new name—lI feel rather difident of expressing an
opinion. In most points it agrees closely with U. corethrura ;
but in the position of the gizzard (in somite vi. instead of vit),
in the position of the spermathece, and in the fact that the sete
are not bifid, the two forms differ. On this last point I think no
great stress can be laid, as Beddard recognizes no bifidity in
his Australian specimen; and I agree with him so far as the
present specimen is concerned, which differs also from Horst’s
species, U. dubia, in the position of the spermathece; in
fact, with regard to these organs, the present and Beddard’s
specimen are intermediate between Horst’s and Perrier’s
species. But are we justified in establishing a new species on
such slender grounds, and from an examination of a single speci-
men? I think not, and prefer to leave the specimen unnamed,
and to regard it as belonging to Perrier’s species, of which it
may be a variety; for we are at present ignorant as to how far

* See Perrier, Arch. d. Zool. expér. et gén. iii. 1874.

. = eae Sas

ZOOLOGY OF FERNANDO NORONHA. 568

variation may occur in Karthworms; since with the exception of
Beddard’s paper .on Perionyx (Journ. Linn. Soc., Zool. 1886,
p- 808), we know absolutely nothing of the subject, and the
present specimen forms a step between U. corethrura and
U. dubia.

The two small worms resemble the specimen A in colour and
in external characters; the clitellum is, however, undeveloped,
so that they are probably young specimens of the same worm.

- The worm B is longer than A, and of a somewhat different
colour, being of a rather more reddish or viclet-brown tint.

The body-wall is transparent, and showed white pyriform sacs
through it much more distinctly than is the case with A.

The worm is, however, so soft that no sete protrude, and I
was unable to satisfy myself as to their exact arrangement
posteriorly; anteriorly they are paired; posteriorly they are
scattered, but whether regularly or not I cannot be positive.

The clitellum occupies somites xiv. to xx1x., and is thus
rather more extensive than in A; but the worm is so soft that
it is difficult to count with accuracy the somites, as some of
the rings may be merely annuli. Thus far, then, we have no
indication as to its genus ; but on opening it, the arrangement of
the septa, seminal reservoirs, and spermathece are seen to agree
with what is found in A.

This second worm is. therefore Urocheta, and doubtless the
same species as the preceding.

It will be seen that I have made no morphological studies of
these worms, nor sought to do more than identify them. Indeed,
they were too badly preserved to be of any use histologically,
and I should not have felt justified in sectionizing them even if
they had been in good condition.

The fact that these worms belong to the genus Urocheta,
which has been already described from Brazil and some of the
neighbouring islands, lends considerable support to Mr. Ridley’s
supposition that they have been imported from the mainland in
the mould in which cultivated plants were brought to the island.
In conclusion I must express my best thanks to Mr. Ridley
for allowing me to examine and identify them.

564 MR. H. N. RIDLEY ON THE

PORIFERA.
By H. J. Carrer, F.R.S.

Dry Specimens.
These were all too much beach-worn for specific distinction.
The Nos. correspond with those on the Specimens.

. Potyruerses, Duchassaing et Mich.*

. HIRCINIA.

wo bo

. Cuatina.—Spicules fine, slender, acerate.
4. Hrrcornra.—F ine structure.
5

. Eusponeata (‘“‘ best Turkey Sponge ” of commerce).

6. Eusponara.— Bearing Polytrema miniaceum.

7. Hrrcrnta.—Skeletal structure partially filled with the
filaments of Spongiephaga communis.

Wet Specimens.

Most of these are too fragmentary for specific distinction,
although possessing the natural characters which they pre-
sented when taken from their habitat.

8. PoLyrHERSES.—-Two coarse pieces alone; the rest on pieces
of a fine Hircinia.

9. Eusponaia (“ best Turkey Sponge” of commerce”).—Three
or four discoloured pieces.

10. CHonpRILLA NucULA, NSdt.

11. Gropta.—? G. Tumulosa, Bk.—Siliceous balls spherical.
Zone-spicule trifid; arms simple, undivided, extending upwards,
outwards, and lastly horizontally. Bearing Polytrema.

12. CHONDROPSIS ARENIFERA, Cart. (Ann. & Mag. Nat.
Hist. 1886, vol. xvii. p. 122).—Acuate spicules, sometimes blunt
at each end.

* It should be remembered that ‘“ Polytherses” is a Hercinia in which the
soft parts have been replaced by a structure composed of the filaments of
Spongiophaga communis, Cart., which is of world-wide occurrence, but of which
the nature is still unknown.

ZOOLOGY OF FERNANDO NORONHA. 565 -

13. CHattna?P species.—Dark, dirty; fragments still bearing
traces of their natural red-purple colour. Extending hori-
zoutally ; throwing up thick ridges; scattered over with short,
erect, tubular vents of different lengths. Fibre tough, charged
abundantly with comparatively large, acerate spicules. Several
pieces, some accompanied by a portion of Chondropsis arenifera.

14. Evsponera (“fine Turkey sponge” of commerce).—_
Typically good, but small specimen; presenting the charac-
teristic, crinkled surface. Colour black above, light sponge-
yellow below. Bearing Polytrema.

15. CHonpritta nucuna, Sdt.—Typically good specimen,
growing over sand-detritus mixed with Polytrema.

16. CHONDRILLA PHYLLODES, Sd¢t.—Antilles. Spicules of two
kinds, viz., pin-like skeletal, and spinispirular flesh-spicule.
Closely allied in this respect to Spirastrella cunctatrix, Sdt.
Colour grey or violet. Consistence gelatinous, firm. Three
typically good specimens growing over sand-detritus mixed with
Polytrema covered with white Melobesia.

17. CHONDROPSIS ARENIFERA.—Black on the surface from a
layer of brown pigmental cells. Growing over a black Stelletta
(P species), also bearing a cortical layer of dark brown-black
pigmental cells mixed with small stellates. Zone-spicule trifid.
Arms simple, straight, extended upwards and outwards.—Four
large pieces.

18. CHonpriIttaA NucuLA, Sd¢t.—Small, but typically good

specimen.

19. Sywascripta.—Common form. Globular, radiated cal-
careous spicule. Colour purple-white. Two pieces.

20. CHaLINA P species.—Same as No. 18. One piece bearing
a bit of Hircinia.

21. Evsronata (“ best Turkey sponge ” of commerce).—Three
small pieces.

92. ? Ancyonium or Hyprorp Zoopuyte.—Digitate, reptant ;
colour yellowish ; consistence soft.

23. AcrINIA ? sp,—Now lead-colour.

566 MR. H. N. RIDLEY ON THE

24. AncyonrtuM.—Congregated, short, columnar individuals ;
constricted circularly throughout the column.

25. Evsponata (“ Honeycomb sponge” of commerce).—Coarse
cavernous structure.

26. Evsponeta.—Ditto.

27. Geopra, same as No. 11.—Fragments of skin and body-
substance only.

28. DonarIa LyNCURIUM.—Four specimens; the largest ? in.
in diameter.

29. Potyruerses, with skeletal structure of Hircinia pro-
truding.

30. Eusponeta (‘‘ Honeycomb sponge ” of commerce).—
Coarse cavernous structure.

31. SuBeRites MAssa, Sd¢.—Spicule of one form only, simply
pin-like, with subglobular head. Light fragments. Surface
warty. Colour yellowish.

32. AXINELLA P species.—Form of specimen globular, about
an inch in diameter, composed of radiating, erect, tough fibre,
charged throughout with projecting tufts of simply acuate
curved spicules; of one form only.

33. TerHyaA CRANIUM.—Two small, discoloured, black frag-

ments.

34. CHONDROPSIS ARENIFERA.—Specimen triangular, elongate ;
4 inches long. Bearing Polytrema and Melobesia.

35. Evusponera (“ Honeycomb sponge” of commerce ).—
Coarse, cavernous structure. Three pieces. Colour black above,
light sponge-yellow below.

36. CHONDROPSIS ARENIFERA.—Good, but small typical speci-
men.

37. ? HymENrIActpon sanauinEa, Bk.—Small insignificant
specimen. Spicule of one from only, viz. pin-like and slightly
curved. Colour now yellowish.

38. LEUCONIA sSAccULATA, n. sp. (Calcisponge)—Form of
specimen sacculated, consisting of four or more inflations

1
4
}
Z
«
’
"
;

— ——_—

ZOOLOGY OF FERNANDO NORONTIA. 557

projecting from a common cavity. Colour white. Entire speci-
men about an inch long, and ? of an inch in diameter (1 x 2X #
inch); broken out on one side, if not the point of attachment.
The uppermost or principal division ending in a peristomatous
mouth, which can only be seen with a microscope, hence to the
unassisted eye looks “naked”; ;2, in. in diameter. The same
on the summit of each inflation, but reduced to the size of a pin’s
head. Surface of the body smooth, composed of intercrossing
arms of quadriradiate spicules only, between which are the pores.
Vents as just described, leading into a general cloacal cavity
corresponding in its inflations with those of the body. Surface
of the cloaca scattered over with holes of very different sizes,
very irregularly situated in a layer of minute quadriradiates
whose fourth arm is much smaller than the rest, curved towards
the -oral orifice and projecting plentifully above the surface of
the cloaca. The spicules of three kinds, viz. 38-radiate, 4-
radiate, and linear cylindrical acerate.

Wall of the body about ;% in. in diameter, composed of.
three layers of spicules, viz. :—1, consisting of comparatively large
quadriradiates whose shaft projects inwards and whose other
three arms are spread out horizontally over the surface; arm
about z. by zt> in. in its greatest diameters; 2 (the middle
‘substance of the wall), consisting of 3- and 4-radiates mixed
irregularly, whose arms are about the same size as that of the
quadriradiates of the first or external layer; 3, or internal
layer, forming the surface of the cloaca, and consisting of minute
or infinitely smaller quadriradiates, whose shafts are directed
outwards; the other two arms horizontal, and the fourth, or
“spine” as it has been called, which is much the smallest, pro-
jecting above the surface in the way mentioned. Peristome con-
sisting of palisading spicules about 45 in. long, very fine and
straight with abruptly pointed ends, crossed and kept in position
by the spreading arms of the quadriradiates of the third, or cloacal,
layer, here much enlarged. Wall permeated by branched canals,
which commencing on the pores on the surface end in the holes or
apertures on that of the cloaca.

Loc. Island of Fernando Norenha.

39. Geop1a.—Fragments of skin and body-substance. Same
as No. 11, &e.

568 MR. H. N. RIDLEY ON THE

Summary of Specimens, arranged according to the Author's
Classification (Ann. Mag. N. H. 1875, vol. xvi. p. 48).

Order I. CaRnosa.
Family GUMMINIA.

CHONDRILLA NucULA, Sdé. 10, 15, 18.
CHONDRILLA PHYLLODES, Sd¢. 16.

Order Il. C ERATINA.
None.

Order II]. PSAMMONEMATA.

Husponara (‘‘fine Turkey sponge” of commerce). 5, 6, 9,
14, 21.

Evusronera (“ Honeycomb sponge” of commerce). 25, 26,
30, 35.

Hirota. 2, 4, 7.
POLYTHERSES. 1, 8, 29.

Order lV. RHAPHIDONEMATA,
CHALINA. 38, 18, 20.

Order V. ECHINONEMATA.

AXINELUA. 32.

Order VI. HOLORHAPHIDOTA.

HiYMENIACIDON SANGUINEA, Bk. 37.
SuBpERITES MASSA, Sdé. 31.

DonAaTIA LYNCURIUM. 28.

CHONDROPSIS ARENIFERA, Cart. 12, 17, 34, 36.
Geropia, 11, 27, 39.

TETHYA CRANIUM. 383.

Order VII. HEXACTINELLIDA.
None.
Order VIIT. CanoaREa.

LEUCONIA SACCULATA, n. sp. 38,

ZOOLOGY OF FERNANDO NORONHA. 569

Remarks.

Of the present collection, it may be stated that the facies is
West-Indian. Chondrilla phyllodes, Sdt., has as yet only been
chronicled by that author, and that, too, from the “ Antilles”
(Grundziige Spong.-F. atlantisch. Gebietes, p. 26). “ Poly-
therses”’ was the name given by Duchassaing and Michelotti
to this transformed, sponge-like body which they dredged in the
Caribbean Sea, and whose constituent parts have been above
noticed. Both sorts of the Officinal Sponge, viz. the “ Best
Turkey ” and the “ Honeycomb” of Commerce, are plentiful, as
they are in the West Indies generally. The Calcisponge,
Leuconia sacculata, is a new species.

MADREPORARIA.
By Prof. P. Marrin Duncan, M.B. (Lond.), F.B.S., &e.

Section MADREPORARIA APOROSA.
Family AstR HID, Hd. & H.
Subfamily AstR#IDH REPTANTES.

ASTRANGIA SOLITARIA, Leswewr, sp., Verrill, Bull. Mus. Comp.
Zool. No. 3; Pouwrtalés, 1871, lll. Cat. Mus. Comp. Zool. p. 79.

A specimen much covered with Nullipores. The corallites
smaller than the Florida types, and the so-called ‘“ pali” not
united before the second and third cycles of septa. The form is
allied to Astrangia Dane and to A. Michelint. It may be con-
sidered as a small variety of the Floridan species.

Subfamily AstRmIDH CMHSPILOS AM.
Mussa, sp.—A worn specimen of a species.

Subfamily AstTRMHIDH AGGLOMERATMH FISSIPARANTES.

Favia conrerta, Verrill, 1867, Zrans. Connect. Acad. vol. 1.,
in Hartt’s Collection of Corals from the Abrolhos Reef, Brazil,
p. 355 (1868).

This species is remarkable for its elongated calices due to
fissiparity, and for their closeness. Often a ridge separates the

LINN. JOURN.—ZOOLGGY, VOL. XxX. “or

570 ZOOLOGY OF FERNANDO NORONHA.

neighbouring calices, instead of some vestige of interspace and
cost. ;

Verrill and Pourtal¢és noticed the alliance of the species with
the genera Goniastrea and Meandrina. In a small specimen
the Goniastroid appearance is striking, and the coste between
the calices are only visible at one spot. There are several rolled
specimens, and, as Pourtalés remarked, they resemble Goni-
astroids very much.

Favia ANANAS, Lamarck, sp.
The specimens have the usual well-developed columella, the

costs are very visible in one, and the fourth cycle of septa is
incomplete.

Favria pDEFoRMATA, Hd. & H., Hist. Nat. des Corall. vol. 1.
p. 434.

A somewhat worn and broken coral, very Coeelorian in appear-
ance, and with large calices, some long and serial in appearance,
but really the result of fissiparity, appears to be a specimen of
this form, the habitat of which has hitherto been unknown. The
walls are close, but in many places their former separation can
be shown. It is a very erratic species, and better specimens are
required.

A very worn specimen of the same species was also obtained.

These two specimens came from Rat Island. They were
washed up on the south-east corner with a number of Sponges.

Section MADREPORARIA FUNGIDA.
Family PhestoFUNGID &, Duncan.
Revision of the Genera of Madreporaria, Journ. Linn. Soc. vol. xviii. 1884, p. 133.

SIDERASTR#A SIDEREA, Hllis § Soland., sp.
This specimen, and indeed all the rolled ones besides, have smaller
calices than the Caribbean type, but that is the only distinction.

They greatly resemble Siderastrea stellata, Verrill, from the ~_

Abrolhos Reef.

Remarks.

This little Coral fauna has the Abrolhos Reef homotaxis, and
the species are fairly intermediate between those of the same
genera of that reef and of the Caribbean Sea.

|
4
.

INDEX,

Acantharus chirurgus, B/., 483.
Acanthias vulgaris, isso, 446,
Acanthoderes, 548.
jaspidea, Germ., 549, 555.
Ridleyi, Waterh., 555.
Acanthopus, de Haan, 513.
Acari found in Great Britain, On some
unrecorded Parasitic, by A. D.
Michael, 400.
Acartia, 149.
denticornis, Brady, 149.
laxa, Dana, 149.

Acarus, 286, 287, 288, 292, 295, 296.
domesticus, de Geer, 287.

Acetes indicus, MZ.-Hdw., 112.

Acharana pheopteralis, Guén., 931,
546.

Achelous Whitei, 4. M.-Edw., 110.

Acheta assimilis, Fabr., 533.

Acidalia Fara, Kird., 531, 544.

retractaria, 545.
Acmea, 503.
noronhensis, Smith, 495, 508.
subrugosa, d’ Orbigny = Lottia
onychina, Gould, 495.
Acontiophorus angulatus, Thomps., 153,
155, 156.
scutatus, 154.
Acta, de Haan, 511.
acantha, M.-Hdw., 511.
areolata, var., Dana, 109.
Riippellii, Krauss, 109.
spongiosa, Dana, 109.

Actinia, sp., 565.

Actinopora regularis, d’Ordb., 280.

Actumnus setifer, de Haan, 110.

Aculeata, 531.

Adamsia palliata, 45.

/Bgotheles, 388, 351.

Novee-Hollandiz, 340.

®olosoma, ftnote, 99.

Ainidea, Baly, 27.

facialis, Baly, 57.
Austhetus, 548, 552.

tuberculatus, Waterh., 552.
Aetea recta, Hincks, 504.
Aglaophenia tubulifera, Hincks, 460.

Agonus cataphractus, Linn., 446.
Ainos of Japan, half-breeds not per-
manent (Gulick), 268.

Alcedo, 337.

Alcyonella, 61, 62.

Alcyonium, 565, 566.
digitatum, 44, known as dead

man’s fingers, 44,

Alpheidee, 112.

Alpheus, Fabr., 506, 518, 522, 528.
comatularum, Hasw., 112.
Edwardsii, Aud., 112, 506, 518,

519.
heterochelis, Say, 518.
minor, Say, 518.
minus, Say, 518.
minus, var. neptunus, Dana, 112.
obeso- -manus, Dana, 520.
panamensis, Kingslen y, 519, 520
Ridleyi, Pocock, 518.
rostratipes, Pocock, 522.

Alsinastrum, 82.

Amathia brasiliensis, Busk, 504.

Amathilla homari, Fabr., 456.

Amathina imbricata, G. B. Sowerd., 398,

400.
tricarinata, 398,

Amblypygus, 55.

Ampeline, 301.

Ampelis cedrorum,

(Shufeldt), 306-318.

Ampelis, 306, 307-318, 352, 354, 355,

o87.

morphology of

cedrorum, 305-309, 314, 315, 352,
382, 387, 392 393.
garrulus, 301, 305.
Ampelisca macr ocephala, Liilly., 456.
tenuicornis, L//7., 456.
Amphidesma cor difor mis, Reeve, 498.
Jayanum, OC. B, Adams, 498.
lenticularis, Sowerb., 498.
orbiculata, Say, 498.
radiata, Say, 498.
reticulata, Sowerb., 498.
subtruncata, Sowerb., 498.
Amphipoda, 46, 47, 118, 465.
Amphisbeena, 474, 475, 476, 481.
43*

572

Amphisbena antillensis, 2. § Z., 482.
ceea, D. § B., 481, 482.
fenestrata, Cope,= antillensis, R. &

L., 482.
Ridleyi, Bowl., 481, 482.
vermicularis, D. & B., ftnote, 481.

Amphiura, 44, 47.

Chiajei, Forbes, 44, 458, 470.
filiformis, O. F. Miill., 44, -458,
470.

Anacharis, 82.

Anapagurus levis, Z’oimps., 454, 469.

Anatomical notes upon the Nestling
Trochilus (Shufeldt), 385.

Anatomy of Antrostomus apart from
the Skeleton, Observations on the
(Comparisons with Chordeiles) (Dr.
R. W. Shufeldt), 348-352.

of certain Caprimulgi (Shufeldt),
338-352.

—— of the North-American Hirun-
dinide (Dr. R. W. Shufeldt), 352-
36).

of the Pelvic Limb in Antros-

fomus and Chordeiles (Suufeldt), 546.

of the Perignathic Girdle and
of other parts of the Test of Dis-
coidea cylindrica, Lamk., sp., On
the, by Prof. P. M. Duncan, and W.
Percy Sladen, 48.

Andrenidez, 531.

Anisolabis Antoni, Dohrn, 530, 532.

janeirensis, Dohrn, 530, 552.
Anomia ephippium, Linn., 450, 467.
patelliformis, Lénn., 450, 467.

Anomis dispartita, Walh., 551, 544.

Anomura, 111.

Anonyx denticulata, 46.

Anous, 480.

melanogenys, Gray, 479.
preta, native name of, 479.
Anser ceerulescens, 138.
canadensis, 137, 138.
hutehinsi, ftnote, 137.
hyperboreus, 138.
leucopareia, ftnote, 137.
occipitalis, ftnote, 137.

Antedon rosacea, Linck, 45, 458.

Antennularia ramosa, Lamk., 460.

Anthophila, 531.

flammicincta, Walk., 531, 544.

Antisarsia gemmatalis, Hiibn., 544,

Antrostomine, 384.

Antrostomus and Chordeiles, On the
Pterylographical tracts of (Dr. R
W. Shufeldt), 339-348.

Antrostomus, 339-352, 384.

carolinensis, 383, 384.

vociferus, 302, 339, 340, 342, 384,
398.

Viuva

INDEX.

Antrostomus apart from the Skeleton,
Observations on the Anatomy of
(Comparisons with Chordeiles) (Dr.
R. W. Shufeldt), 343-352.

Ants, Bees, and Wasps, Observations

on, by Sir John Lubbock, 118.

oe honey-dew from Aphi- |

es 121.

On the Colour-sense of Lae J.
Lubbock), 122.

mimicry among, 121.

and seeds of Melampyrum pra-
tense (Lubbock), 133.

Aphides, ants collecting
from (Lubbock), 121.

Aphrodita aculeata, Linn., 45, 458,
469.

Aplysia, sp., 497.

Apogon imberbis, Linn., 483.

Aporrhais pes-pelecani, Linn., 46, 448, ‘
466. .

honey-dew

Arachnids, 438.

Arca Adamsi, Shuttlew,, 499.
imbricata, Bruguiére, 499, 503.
lactea, Zinn., 499.
solida, Sowerb., 499.

(Acar) Adamsii, Shuttlew., 499,
503.

Arcania pulcherrima, Hasw., 111.

1l-spinosa, Adams & White = A.
pulcherrima, Hasw., 111.

Arcturus longicornis, Sowerb., 46, 456,
469.

Argentina sphyrena, Linn., 448.

Artotrogidee, 147, 155.

Artotrogus Normani, Brady § Roberts,

154.

Arvicola agrestis (field-vole), Myocoptes
tenax, and Goniomerus musculinus,
Mich., parasitic on, 403, 406.

Ascidia ‘mentula, O. fF, Miil., 448,

466.
scabra, O. #'. Miill., 448, 466.
virginea, O. F. Miill,,
466.

Astacidea, 516.

Astarte compressa, Mont., 450, 467.
elliptica, Brown, 450, 467.
sulcata, DaC., 450, 467.

Asterias rubens, Linn., 458, 470.
violacea, O. F. Miill., 458, 470.

Astreide, Hd. & H., 569.
agglomerate fissiparantes, 569.
ceespitosee, 569.
reptantes, 569.

Astrangia Dani, 569.

Michelini, 569.
solitaria, Lesweur, 569.
Astropecten irregularis, 45.

448,

| Ateenius, sp., 549.

a eS ae ae

INDEX. 5/3

Atemeles, 119, 120.
emarginatus, 119.
Atergatis floridus, Rwmph, 109,
integerrimus, Lamk., 109.
Aulacophora, Chevr., 1, 2, 27, 156, 163,
175, 178.
abdominalis, Fabr., 2, 13, 14, 17.
albicornis, Chupuis, 9.
albofasciata, Baly, 2, 4, 6, 7.
analis, Weber, 176, 177, 184, 188.
anchora, Redt., 26.
antennata, Baly, 3,4, 21, 22.
approximata, Baly, 2, 4, 12.
argyrogaster, Montrowz., 178, 188.
atripennis, Fabr., 2, 4, 8.
atripennis, Hope, 5.
austro-caledonica, Monfrouz., 179,
188.
Batesi, Jac., 26.
bicolor, Weber, 3, 4, 19, 20, 177.
bipartita, Baly, 176, 183, 188.
Boisduvali, Baly, 2, 4, 10, 23.
coffe, Hornst., 3, 4, 17, 18, 20,
cornuta, Baly, 5, 5, 15, 16.
costatipennis, Baly, 5, 26.
dorsalis, Boisd., 11.
Downesi, Baly, 3, 4, 20.
Duboulayi, Baly, 3, 4, 22.
Duvivieri, Baly, 5, 24.
excavata, Baly, 3, 5, 18.
excisa, Buly, 5, 25.
Fabricii, 2, 5, 14.
femoralis, Motsch., 16.
flaviventris, Baly, 2, 4, 15.
foveicollis, Kiister, 3, 5, 13, 16,
Ve
fraudulenta, Jacoby, 176, 182, 185,
188.
frontalis, Baly, 176, 181, 188.
impressa, Habr., 18.
instabilis, Daly, 2, 4, 10.
lata, Baly, 2, 4, 8, 9, 10, 13.
levifrons, Baly, 176, 185, 188.
Lewisii, Baly, 5, 24, 179, 188.
luteicornis, Mabr., 3, 5, 21, 22,
25.
melanoptera, Boisd., 2, 4, 7.
melanopus, Blanch, 2, 4, 5.
Mouhoti, Baly, 5, 25.
nigripennis, Mo?sch., 2,4, 5, 24.
nigrivestis, Boisd., 3, 20, 24.
nigroscutata, Baly, 2, 5, 14, 177.
nigrosignata, Baly, 4, 12, 23.
occipitalis, Baly, 176, 179, 184,
185, 188.
octomaculata, Baly, 3, 4, 17.
Olivieri, Baly, 176, 184, 188.
- orientalis, Hornst., 176, 180, 188.
palliata, Schadler, 176, 180, 188.
palustris, Perroud, ftnote, 3, 178.

Aulacophora Perroudi,
188.

postica, Chapuis, 2, 8, 4, 12.
propinqua, Badly, 2, 3, 4, 11.
pygidialis, Baly, 2, 4, 7.
robusta, Duviv., 16.
rosea, Fabr., 2, 4,9.
rubrozonata, Blanch., 2, 4, 11.
semilimbata, Baly, 4, 24.
sexnotata, Chapuis, 19.
similis, Oliv., 3, 5, 16, 17.
simplicipennis, Clark, 21.
smaragdipennis, Duv., 23.
Stevensi, Baly, 3, 5, 22.
testacea, Fabr., 2, 3, 138.
tetraspilota, Baly, 3, 4, 17.
unicolor, Jac., 3, 23.
varians, var. B?, Chapuis, 18.
Viridipennis, Chap., 23.
Wallacii, Baly, 3, 5, 15.
Wilsoni, Baly, 177, 188.

Aulophorus, 100.
vagus, Leidy, 95.

Aves of Fernando Noronha, by R.

Bowdler Sharpe, 477.
Axinella, sp., 566, 568.

Baly, 177,

Balanus hamiri, Asc., 456.

Baly, J. S., Descriptions of a new Genus
and of some new Species of Galeru-
cine, also Diagnostic Notes on some
of the older described species of
Aulacophora, by J. 8. Baly, 1.

—-—, Descriptions of some Genera and
Species of Galerucine, 156.

Bargilis, 535.

Bathyporeia pilosa, Lindstr., 456.

Bees, Ants, and Wasps, Observations
on, by Sir John Lubbock, 118.

Belonuchus formosus, Grav., 549.

Bewbicide, 531.

Benham, W. B., Oligocheta of Fer-
nando Noronha, 560.

Bill-trout, ftnote, 71, 72.

Biological terms, definition of (Gulick),
200.

3irds and Mammals of Hudson’s Bay

Company’s Territory, and of Arctic
Coast of America, by Dr. J. Rae,
136.

Blapstinus, 551.
pulverulentus, Hsch., 551.

Ridleyi, Waterh., 551.

Blastophaga, 5387.
brasiliensis, Mayr, 537.
obscura, Kird., 581, 537.

Blatta americana, Linn., 475, 582.
incommoda, Kird., 530, 533,
orientalis Linn.. 533.
pacifica, Coqueb., 533,

574

Blatta poststriga, Walk., 532.
surinamensis, Linn., 538.

Blattide, 530.

Blenniidz, 38.

Blenny, Common, 382.

Viviparous, 41.

Blowfly (Calliphora erythrocephala),
On the Structure of the Retina of
the, by B. T. Lowne, 406.

(Calliphora erythrocephala), On
the Structure and Development of
the Ovaries and their appendages in
the, by B. T. Lowne, 418.

Blue-winged goose, 138.

Bolina bivittata, Walk., 531, 544.

Bolocera tuedizx, Johnst., 460.

Boreophausia inermis, Kroyer, 456.
Raschi, WZ, Sars, 456, 469.

Botys detritalis, Guén., 546.
flavidissimalis, Grote, 546.
pheopteralis, Guén., 546.

Boulenger, G. A., Pisces of Fernando
Noronha, 483.

, Reptilia of Fernando Noronha,

Bousfield, E. C., The Natural History
of the Genus Dero, 91.
Brachelytra, 549.
Brachiopoda, 452, 462, 464, 468, 471,
472.
Brachyura, 109.
Brissopsis lyrifera, Forbes, 45, 458,
470.
Bruchida, 553.
Bruchus porosus, Sharp, 553.
Buccinum pusio, Reeve, 486.
undatum, Linn., 448, 466.
Buceros rhinoceros, 347.
Bugula turbinata, Alder, 452.
Bulimus, 484.
Jacobi, 501.
Ramagei, 484.
Ridleyi, 476.
(Bulimulus) Ridleyi, Sith, 476,
501, 508.
(Tomigerus) Ramagei, Smith, 500,
503.
Burra-tree, 478.
Byblis kallarthrus, Stedbing, 118.

Calanide, 147.
Calanus_ finmarchicus,
147, 148, 456.

gracilis, Dana, 147,
pavo, Dana, 148.
propinquus, Brady, 147.
tonsus, Brady, 147.
valgus, Brady, 147.

Calappa lophos, Herbst,

Haan, 111.

Giimner, 146,

var. y, de

INDEX.

Calappide, 111. |

Calcar Olfersi, Fischer, 493.

Calcarea, 568.

Callianassa subteranea, 45.

Callionymus lyra, Linn., 446.
maculatus, Raf., 446.

Calliphora erythrocephala, 406, 418.

Calliphoria vomitoria, 125.

Callisoma crenatum, Spence Bate, 456.

Calocaris Macandreze, Bell, 45, 454,

469.

Calycella fastigiata, Adler, 460.

Calyptrea alveolata, A. Adams, 492.

Campanularia angulata, Hincks, 460,
volubilis, Linn., 460.

Camponotus zthiops, 121.
bimaculatus, Smith, 531, 588.
ebeninus, Mmery, 121.
foveolatus, Mayr, 121.
lateralis, 121.
ligniperdus, 123.

Canada goose, 137, 188, 189. -

Cancer acanthus, M.-Kdw., 511.
arenarius, Catesby, 512.
corallinus, Herbst, 510.
coronatus, Herbst, 510.
depressus, Fabr., 518.
lobatus, Milne-Edw., 511.
pagurus, 45.
planissimus, Herbst, 5138.
sguamosus, Herbst, 518.

Cancride, 109, 510.

Cancroidea, 510.

Candace, 149.
brevicornis, Thomps., 142, 155.
nigrocincta, Thomps., 148, 155.
pachydactyla, Dana, 149.
truncata, Dana, 148, 149,

Canna Warscewiczi, 264.

Caphyra Archeri, Walk., 110, 116, 117.

Caprimulgi, anatomy of certain (Dr. R.

W. Shufeldt), 3388-352.
Caprimulgidee, 299, 300, 306, 384.
Caprimulgine birds, 299, 300, 306, 327,

336-389.

Caprimulgine Cuckoos, 318, 338.

Caprimulgus, 322, 341, 342, 343, 345,

346, 347, 349.
europaeus, 340, 344, 346.
forcipatus, 340.
longipennis, 340.
psalurus, 340.

Carcinoplacide, 110.

Cardium aculeatum, Linn., 450.
echinatum, Linn., 450.
fasciatum, Mont., 450, 467.
medium, Linn., 497, 503.
minimum, Phil., 450, 467.
subelongatum, Sowerb., 497.

Caridea, 518.

INDEX.

Caridion Gordoni, Spence Bate, 456,
Carnosa, 568.
Carpilius, Desm., 510.

corallinus, Herbst, 510. -
Carter, H. J., Porifera of Fernando

Noronha, 564.

Cashewnut-tree, 478.
Cassia, 474, 479.
Catachrysops trifracta, Butl., 543.
Cathartes aura, Linn., ftnote, 315.
Cathartide, ftnote, 315.
Catometopa, 110.
Cellaria fistulosa, Linn., 452.
Cellepora, 505.

avicularis, Hincks, 452.

pumicosa, Linn., 452.

ramulosa, Linn., 452.

Ridleyi, Kirkp., 504, 505, woodcut

506.

Celleporidzx, 505.

Cellulariidz, 504.

Centronotus, 39, 40, 41, 42.
gunnellus, 39, 40, 41, 446.

Centropages brachiatus, 146, 150.
hamatus, Li/d7., 456.
typicus, 146, 150.
violaceus, 150.

Centropodine, 337.

Cephalopoda, 465.

Cerapus abditus, Zemplet., 456.

Ceratina, 568.

Ceratocarcinus dilatatus, A. M.-Edw.,

Ceratoplax ciliatus, S¢émps., 110.
Cerithiopsis, sp., 505.
Cerithium atratum, Born, 492.
caudatum, Sowerh., 492.
Cerotoma, 156.
Cheetogaster, ftnote, 99.
Cheetognatha, 437.
Chetura, 363, 364, 366, 368, 371,
372, 376, 380, 383, 384, 390.
pelagica, 302, 365, 3870, 371, 372,
Sr o10, Off, 018, 389, 390,

394.
Vauxi, 383.
Chalcididx, Subfamily Eucharine ;

with descriptions of several new
Genera and Species of Chalcidide
and Tenthredinide, A Synopsis of |
the Genera of, by W. F. Kirby, 28.

Chaleididx, 28, 35, 531.

Chalcidins, 35.

Chalcis amenocles, Walk., 36.
Kassalensis, W. F'. Kirb., 36, 37.
restituta, Walk., 36.

Chaleura, W. F. Kirb., 30, 33.
deprivata, Walk, 30, 37.

Chalina, sp., 460, 564, 565, 568.

Chama, sp., 498.

570

Chara, 82.

Characters of the Genus Lophopus,
with description of a new Species
from Australia, On the, by 8S. O.
Ridley, 61.

Charidea, 157, 187.

fortunei, Baly, 158.
punctato-striata, Wotsch., 157, 158,
137;

Cheiloxena, 163.

Cheirocratus assimilis, Zil/jeb., 456.
Sundevalli, Rath., 456.

Chelidon, 354, 361, 363, 364.
erythrogaster, 305, 561, 395.

Chilopoda, 526.

Chilostomata, 280.

Chirocerus, 32.
fureatus, Brullé, 32.

Chiton astriger, Reeve, 496.
Boogii, Haddon, 503.
marginatus, Penn., 450, 467.
pectinatus, Sowerb., 496.
(Acanthochiton) astriger,

496.
(Ischnochiton) caribbeeorum, Caz-
pent., 496, 503.

) pectinatus, Sowerd., 496.

Chlorodius americanus, H. de Sauss.,
HrT:

Chondrilla nucula, Sdz¢., 564, 565, 568.
phyllodes, Sd¢., 565, 568, 569.
Chondropsis arenifera, Cart., 564, 565,

566, 568.

Chordeiles, Anatomy of Antrostomus
compared with (Shufeldt), 343-352.

——- and Antrostomus, on the Ptery-

Reeve,

graphical tracts of (Dr. R.
Shufeldt), 389-345.
Chordeiles, 322, 334, 335, 338, 339-

352, 384.
acutipennis, var. texensis, 384,
33D.
texensis, 302, 335, 339, 341, 344,
346, 350.
, var. Henryi, 346.
virginianus, 302, 385.
, var. Henryi, 302,
Chordeilinx, 384.
Chorioptes, 404.
Chrysodomus antiquus,
466.
Chrysomelide, 156.
Cicada septemdecim, 227, 228.
tredecin, Fz/ey, 228,
Cidarida, 58.
Cidaris, 58.
tribuloides, Lams., 559.
Ciona intestinalis, Linn., 448, 466.
Cirolana hirtipes, M.-Edw., 456, 469.
Clamatores, 301, 310, 316.

Linn,, 448,

576

Clavagella aperta, 400.
minima, G. B. Sowerb., 400.
Clibanarius vulgaris, Herbst, 112.
Clinus delalandu, C. & V., 483.
nuchipinnis, Q. ¢& G., 483.

Clivicola riparia, 305, 362.

Clupea humeralis, C. ¢ V., 483.

Clypeastroids, 59.

Cnesinus, Horn, 554.

Coccus Persice, 435.

Coccyzus americanus, 333.

minor may be Geococcyx, 388.

Cochlostyla (Axina) Gloynei, G. B.
Sowerb., 398, 400. |

Coelenterata, 460, 462, 464.

Ceeligena clemencize, 303.

Celophyma granulata, Hagen., 276.

striata, /euss, 276.

Ceelopleurus, 53, 58.

Ceenobita perlata, W.-Mdw., 112.

Coleoptera of Fernando Noronha, by
Chas. O. Waterhouse, 548. |

Colour-sense of Ants, On the (Sir J.
Lubbock), 122.

Columbella, 489.

mercatoria, Linn., 486, 503.
Columbide, 479.
Compound Eyes and Ocelli in Ants,
Bees, and Wasps, On the function of
the (Lubbock), 125.
Conchexcetes conchifera, Hasw.,
114)
Conger vulgaris, Cwv., 448.
Conilera cylindracea, Mont.,
469.
Conocephalus triops, Linn., 534.
vernalis, Kirb., 5380, 534.

, var. n, frater, Kird.,

534.

Conus archetypus, Crosse. 485.
castus, Reeve, 485.
daucus, Hwass, 485.
erythreensis, Bech, 485.
imperialis, 485.
lividus, 485.
mamumillaris, Green, 485.
nebulosus, So/., 484.
piperatus, Dillwyn, 485.
rattus, 485.
Reevei, Kiener, 485.
sanguinolentus, Reeve, 485.

Copepoda of Madeira and the Canary
Islands, with Descriptions of New
Genera and Species, by i. C. Thomp-
son, 145..

Copepoda, 47, 145-155, 444.

Copidita, sp., 553.

Copilia mirabilis, Dana, 158,

Coracina cephaloptera, 368.

Corbula gibba, Olivi, 452, 467,

108,

456,

539,

INDEX,

Corella parallelogramma, O. F. Miili.,
448, 466.

. Corvide, 310.

Coryceide, 147, 153.

Coryceeus, 1538.
limbatus, Brady, 158.
pellucidus, Dana, 153.
speciosus, Dana, 153.
varius, Dana, 153.
venustus, Dana, 158.

Corystes cassivelaunus, 45.

Cottus bubalis, Huphr., 446.
Lilljeborgii, Collett, 446.
scorpius, Linn., 446.

Crassatella, 399.
japonica, G. B. Sowerb., 399;

400.

Crambi, 531.

Crangon, 45.

Allmani, Kinahan, 454, 469.
bispinosus, 45.

echinulatus, WW. Sars, 454, 469.
spinosus, Leach, 454,
trispinosus, 45,

vulgaris, 45.

Cremastogaster lateralis, 121,
scutellaris, 121.

Cribrella oculata, Linck, 458, 470.

Cribrilina radiata, Mold, 504.

Crinoid, 45.

Crisia denticulata, Lamk., 454.
eburnea, Linn., var. aculeata, 454.

, var. producta, 454.
Holdsworthii, Busk, 504.

Cristatella, 62.

Crossaster papposus, Linch, 458,

Crotalaria, 477.

Crotophaga, 338.

Crustacea of Fernando Noronha, by

R. I. Pocock, 506.
from Singapore, Notes on a Col-
lection of, by A. O. Walker, 107.
Crustacea, 45, LO7-117, 438, 445, 454,
462, 464, 465, 468, 471, 472. ;

, Sessile-eyed, 46, 74, 83.

, Stalk-eyed, 45.

Crustaceans, Sessile-eyed, 46, 74, 83.

Cryptodon croulinensis, Jeffr., 450,

467.
ferruginosus, Forbes, 450, 467.
flexuosus, Mont., 450, 467:

Cryptopodia fornicata, Mabr., 109. —

Cuckoos, 338, 337, 338.

Cuculidex, 337.

Cuculinz, 333, 337, 353.

Cuculus eanorus, 337.

Cucumaria Hyndmanni, Thovps., 458,

470.
Cucurbitacese, 474, 473, 479.
Cupularia, 280.

INDEX.

Cuspidaria abbreviata, Forbes, 452, 468.
euspidata, Olivi, 452.

Cyclometopa, 109.

Cyclopide, 147, 151.

Cyclostomata, 275, 277, 278.
Cyclostomatous Bryozoa, On some
Ovicells of, by A. W. Waters, 275.

Cylichna noronyensis, Watson, 503.

Cylindrecium dilatatum, Hincks, 454.
Cymbasoma, Thomps., 154.
Herdmani, Thomps., 155.
rigidum, Thomps., 154, 156.
Cynipide, 28.
Cynips adscendens, Fadr., 28.
Cynorta, Baly, 161.
apicipennis, Paly, 162, 187.
facialis, Baly, 162, 187.
ocellata, Baly, 161, 187.
Cynthia echinata, Linn., 448, 466.
Cyperus, 106.
Cyprea cinerea, Gmel., var., 490.
clara, Gaskoin, 490.
(Trivia) pediculus, Linn., 490.
Cyprina islandica, Linn., 450, 467.

Cypseli and Trochili, Morphology of

certain (Shufeldt), 865-387.
, anatomy of the head, 375-

, anatomy of pelvic limb, 373-
375

ee , axial skeleton, 377-879.

—— -——-, comparison of pectoral
limbs, 369-373.

, external form and _ pterylo-

graphy, 365-369.

, heart and carotids, trachea,
viscera, &c., 380-383.

Cypselidz, 299, 300.

Cypseloides niger and Nyctidromus
albicollis, var. Merrilli, Notes on (Dr.
R. W. Shufeldt), 385.

Cypseloides, 583, 384.

erythrogaster, 384.
niger, 300, 3838, 384.
Cypselus, 3853-391.
alpinus, 374.
apus, 355, 366, 368.
erythrogaster, 384.
Cyrtosoma, 552.
Cythere, Buird, 47.
minna, Baird, 47.
pellucida, Baird, 47.

Dacelo, 337.
gigantea, 337, 388.
Dacelonine, 88s.
Dactylopterus volitans, Linn., 488.
Dactylopus tisboides, Brady, 47.
tisboides, Claus, 152.

Dactylosternum abdominale, fabr., 549,

577

Daphnia, 122.
Day, F., On the Loch-Leven Trout
(Salmo levenensis), 71.
Decapoda, 507.
Deep-Water Fauna of the Clyde Sea-
area, On the, by W. E. Hoyle, 442.
Defrancia, 281.
diadema, Gold., 284.
disticha, Hagenow, 284.
reticulata, Hagenow, 284.
Delavalia robusta, Brady & Roberts,
152.
Dendronotus arborescens, Mii//., 448.
Dentalium entale, 46.
entalis, Linn., 450, 467.
Dermacarus, Haller, 287.
Dermestes felinus, Fabr., 549.
Dero. The Natural History of the
Genus, by E. C. Bousfield, 91.
Dero, Oken, 91-107.
acuta, Bousf., 108, 105,
digitata, 95, 96, 103.
furcata, Oken, 92, 98, 95, finote
97, 102, 103, 105, 107.
latissima, Bousf., 101, 102, 108,
104, 107.
limosa, Letdy, 95, 103, 105, 107.
Mileri, Bows/., 101, 104, 107.
obtusa, Perrier, 95, 103, 104, 105,
106

palpigera, Grebincky, 95, 105.
Perrieri, Bousf., 95, 108, 104, 107.
philippinensis, Semper, 95, 105.
Rodriguezti, Semper, 95, 105.
Descriptions of some Genera and Spe-
cies of Galerucine, by J. 8. Baly, 156.
Deviation from an average (Gulick),
193.
Diacantha, 168.
Diacheta, Benham, 561.
Diadema, 58.
saxatile, Linn., 560.
Dias longiremis, Lidljeb., 146, 149, 456.
Diastopora lineata, Macg., 284.
obelia, Johnst., 454.
Diastylus, 45, 47.
Diczeum, 353.
Dicoryre conferta, A/der, 460.
Dicrurus, 357.
Dinarda, 120, 121.
dentata, 120.
Diogenes avarus, var., Heller, 112, 18,
Luh
granulatus, Miers, 113.
miles, Fabr., 112.
Diosaccus tenuicornis, Claus, 152.
Diphasia attenuata, Hincks, 460.
fallax, Johnst., 460.
pinaster, H//. §° Sol., 460.
tamarisca Hill, J Sol., 460.

578

Diplopoda, 528.

Diploptera, 531.

Diptera, 582.

Direction in Ants, Bees, and Wasps
(Lubbock), 127.

Discocavea, 281.

Discoidea cylindrica, Lamk. sp., On
the Anatomy of the Perignathic
Girdle and of other Parts of the Test
of, by Prof. P.M. Duncan and W.
Percy Sladen, 48.

Discoidea, 48-61.

conica, 54, 55, 60.

eylindrica, Laimk., 48-61.

Dixoni, 60.

Fayrina, 60.

Forgemolli, 60.

infera, 60.

Jullieni, 60.

minima, 60.

subuculus, 60, 61.
Discoporella, 280.

californica, Busk, 282.

echinata, Macg., 282.

Holdsworthii, Busk, 285.

reticulata, Macg., 284.
Discotubigera, 284.

lineata, Macq., 284, 285.

Disporella, Gray, 280.

Divergence, cumulative, through cumu-
lative segregation (Gulick), 212-
222.

Divergent Evolution through Cumu-
lative Segregation, by Rev. J. T.
Gulick, 189.

Doclea muricata, Herbst, 109.

ovis, Herbst, 109.
tetraptera, Walk., 109, 114, 117.

Dogs, Experiments on the Sense of
Smell in, by G. J. Romanes, 65.

Dolichopodide, 552.

Donatia lyneurium, 566, 568.

Donax anatinus, 46. __

Dorippe astuta, Hadr., 108, 111.

sima, M.-Hdw., 108, 111.

Dorippidee, 111.

Doris, sp., 497.

Dosinia exoleta, Linn., 450.

lincta, Pult., 450.

Dotilla myctiroides, 1/.-Hdw., 111.

Dragonflies, 475.

Drepanopus furcatus, Brady, 149.

Dromia Rumphii, ? Fabr., 111.

vulgaris, M.-Hdw., 111.

Dromiidea, 111.

Duncan, Prof. P. M., Madreporaria of
Fernando Noronha, 569.

, and W. Percy Sladen, On the

Anatomy of the Perignathic Girdle

and of other Parts of the Test |

INDEX,

of Discoidea cylindrica, Lamk., sp.,

Ebalia tuberosa, Penn., 454, 468.
Kehinide, 51, 58.
Echinocardium cordatum, 45.
flavescens, OF F. Miill., 458, 470.
Echinocyamus pusillus, 45.
Echinodermata, 458, 462, 464, 466, 470,
471, 472, 559.
Echinodermata of Fernando Noronha,
559.
Echinoidea, 45, 48, 57.
Echinonemata, 568.
Ecehinoneus, 55.
Eehinus, 58.
esculentus, Linn., 458, 470.
miliaris, P. Z. S. Miill., 458.
Ketinosoma atlanticum, Br. g Rob.,
456.
Edora, 47.
Egeria longipes, Herbst, 109.
Elainea pagana, 478.
Ridleyana, Sharpe, 478.
Eledone cirrosa, Lamk., 448.
Elimea, 535.
Emarginula crassa, Sowerb., 448.
reticulata, Sowerd,, 450.
Entalophora intricaria, Bush, 278, 28).
Entomophaga, 531.
Entomostraca, 47.
Ephemeride, 85.
Epimeria cornigera, J. C. Fabr., 456.
Epitragus Batesii, MWaéklin, 550, 551.
Epureea limbata, 549.
Eremophila alpestris, ftnote, 309.
Hrinaceus europzxus (Hedgehog), 404.
Symbiotes tripilis, Wich., parasitic
on, 404.
Eriphia levimana, Latr., yar, Smithii,
M‘Leay, 110.
Eriphiide, 110.
Ervilia subcancellata, Smith, 503.
Erythrina-tree, 478.
Escharella rostrigera, Smitt, 504.
HKucalanus attenuatus, Dana, 148.
setiger, Brady, 148.
Eucheta norvegica, Boeck, 456.
prestandree, Philippi, 148.
EKucharinx, List and Synopsis of
Genera of, 28, 33, 37.
Kucharis, Latr., 28, 33.
contingens, Wa/k., 37.
deprivata, Walk., 30.
festiva, Fabr., 29.
Jurcata, Fabr., 32.
Tello, Walk., 29.
Kollari, Forst., 28,
rapo, Walk., 29.
volusus, Walk., 31,

INDEX. 579

Eucharis zalates, Walk., 30.
Eucharissa, 37.
Eucharissine, 37.
Eudora truneatula, 45.
Eumenia Jeffreysi, McJnt., 458, 469.
Eupagurus bernhardus, Linn., 454,
468.
excavatus, Miers, 454.
Prideauxii, Leach, 454, 469.
pubescens, Kréyer, 454, 469.
Eurynome aspera, Penn, 454, 468.
Euspongia, 564, 565, 566, 568.
Euterpe gracilis, Claus, 151.
Euthyrrapha pacifica, Ci oqueb., 530, 533.
Eutrochus, 494.
Adamsi, Pilsbury, 494.
perspectivus, A. Adaiis, 494.
Evadne Nordmanni, Lov., 456.
Eyvania, 475.
levigata, Latr., 531, 538.
Evaniide, 531.
Evolution, Divergent, not explained by
Natural Selection (Gulick), 202-212.
Experiments on the Sense of Smell in
Dogs, by G. J Romanes, 65.

Eyes and Ocelli in Ants, Bees, and
Wasps, On the Function of the Com-
pound (Lubbock), 125.

Fauna of the Clyde Sea-Area, On the
Deep-Water, by W. E. Hoyle, 442.

Favia ananas, Lamk., 570.

conferta, Verrill, 569.
deformata, Hd. §& H., 570.

Fernando Noronha, Notes on _ the
Zoology of, by H. N. Ridley, 473-
570.

Field-vole (Arvicola agrestis), Gonio-
merus musculinus, Mich., 406, and
Myocoptes tenax, Mich., parasitic on
403.

Filigrana implexa, Bers, 458, 469.

Filisparsa orakeiensis, S7o/., 279, 280.

Fissurella aegis, Reeve, 494.

alternata, Say, 494.
barbadensis, Gmelin, 495.
cancellata, Sowerb., 494.
Dysoni, Reeve, 494.
larva, Reeve, 494.
lentiginosa, /eeve, 494.
nubecula, Linn., 495.
suffusa, Reeve, 494.

Flustra, 44.

foliacea, Linn., 452.

securifrons, Pallas, 452.
Forcinella Antoni, Dohrn, 532.

janeirensis, Dohrn, 582.

Forfieula riparia, Pall., 532.

Forficulide, 530.

Formica, 120,

Formica bimaculata, Smith, 538.
congerens, 120.
exsecta, 121.
fusca, 118, 119, 121, 182, 138,
134,
gagatus, 121.
pratensis, 128.
pressilabris, 121.
rufa, 120, 121, 126.
rufibarbis, 121.
ruficeps, Fabr., 538.
sanguinea, 118-121, 128.
relation between, and
118.
Formicidae, 531.
Fossarus ambiguus, Lin2., 503.
Fossores, 531.
Fredericella sultana, 64.
Freshwater Polyzoa (Ridley), 61, 64.
Fusus (Sipho) gracilis, DaC., 448, 466.

its slaves,

Gadus xglefinus, Linn., 446.
Esmarkii, Vi/ss., 446.
luseus, Linn., 446.
merlangus, Linn., 446.
minutus, Linn., 446.
morrhua, Linn., 446.
Galathea Andrewsi, 45.
dispersa, Spence Bate, 45, 454,
469.
elegans, Adams §& White, 112.
nexa, Embleton, 454, 469.
squamifera, Leach, 45, 454, 469.
Galatheidex, 112.
Galbulide, 388.
Galearia, Brullé, 32.
violacea, Brullé, 32.
Galene bispinosus, Herdsz, 110.
Galerites cylindricus, Lams’., 48.
Galeruca, 156, 157.
analis, Olivy., 184.
Buquetii, Guér., 170.
luteicornis, Fabr., 21.
melanoptera, Boisd., 7.
multicostata, Jacoby, 157.
Galerucella, 157, 186.
Galerucine, On some Species of, by J.
S. Baly, 1; table of spec., 2
——, Descriptions of some Genera and
Species of, by J. S. Baly, 156.
Galerucine, 156, 163.
Gammarus, 74, 475.
locusta, Linn., 456.
pulex, 83.
Ganosoma attenuatum, Mayr, 538.
dispar, Kirb., 531, 537.
Gasteropoda, 503.
Gebia, Leach, 515.
spinigera, S. 1. Smith, 515.
Gebiidex, 112, 5r5.

580

Gebiopsis Darwinii, Mzers, 112.

Gecko, 474, 475, 481.

Gelasimus vocans, Zinn., 110.

Generation, Independent, Effects

Selection and (Gulick), 202-212.

Geococeyx, 325, 334, 336, 337, 344,

388.
californianus, 300, 325, 333, 385,
337, 338, 388.

Geodia, 564, 566, 567, 568.
tumulosa, Bh., 564.

Geometres, 531.

Geophilus, 526.
occidentalis, Meinert, 527.

Ridleyi, Pocock, 526.
Glaucidium, 328. |
Glyciphagus, 285-298.
Crameri, Michael, 287. |
cursor, Gervais, 288, 295, 296.
domesticus, de Geer, 285, 287, 288,
289, 290, 291, 293, 294, 295, |
296, 298.

spinipes, Koch, 285, 287, 288, 289,
292, 2938, 294, 295, 296, 297,
298.

Glyphidodon saxatilis, Lénn., 485.

Gnathostomes, 58.

Gobiesox cephalus, Lacép., 483.

Gobius Jeffreysii, Giinther, 446.
minutus, Gi., 446.
soporator, C. § V., 483.

Gonatonotus pentagonus,

White, 109.

Goniastrea, 570.

Goniomerus, Mich., 401, 405.
musculinus, Mich., 405, 406 ; para-

sitic on field-vole (Arvicola
agrestis), 406. _

Goniosoma crucifera, Fabr., 110.
ineequale, Walk., 110, 116, 117.
natator, Herbst, 110.

Gonodactylus, Latr., 526.
chiragra, Fabr., 1138, 506, 526.

Gonolobus micranthus, 478,

Gorgonia axis, 504.

Gorgoniz, 397.

Grantia ciliata, /Vlem., 460.

Grapside, 110, 512.

Grapsoidea, 512.

Grapsus, Lamh., 512, 518.
maculatus, Cateshy, 506, 512.
strigosus, Latr., 110.

Grey trout, 71 ftnote, 72.

Gryllidz, 550.

Gryllodes, 554.

Gryllus assimilis, /abr., 5380, 583.
forticeps, Sauss., 580, 533.

Gulick, Rev. J. Thos., Divergent Evo-

lution through Cumulative Segrega- |
tion, 189. |

Adams &§

INDEX,

Gumminia, 568,
Gygis candida, Gm., 480.

Hemulon chrysargyreum, Giinther, 483.
Halecium Beanii, /ohnst., 460.
halecinum, Linn., 460.
muricatum, ZU. § Sol., 460.
Halictus, 474, 543
alternipes, Kird., 531, 542.
atripyga, Kirb., 581, 548.
levipyga, Kirb., 531, 542, 543.
Halticella liberator, Walk., 35.
Haploops setosa, Boeck, 456.
tubicola, Lill7., 456.
Haplosonyx, 156.
sumatree, 161.

Harpacticide, 147, 151.

Harpacticus chelifer, Wiiller, 152.

Harpinia plumosa, Kréy., 456.

Harporhynchus rufus, 315.

Hedgehog (Hrinaceus europeus), Sym
biotes tripilis, Mich., parasitic on,
404.

Heliothis armiger, Hiidu., 531, 544.

Helix entodonta, Pfeiffer, 500.

(Ophiogyra ?) quinquelirata, Smith,
500, 503.

Hemidactylus mabouia, Mor., 475, 481.

Hemidianeura Cameroni, W. F. Kirb.,
34, 37.

Hemiprogne, 390.

zonaris, 301.
zonatus, 392.

Hemiptera, 552.

Hemirhamphus
483.

Hereeus percultus, Dist., 547.

variegatus, Kirb., 582, 547.

Heredity (Gulick), 192.

Hero, self-fertilized plant of Zpomea
purpurea (Darwin & Gulick), 264.

Hero, 264, 265, 266.

Hesperocichla, 309-314,

neevia, 312, 314, 392.

Heteerius ferrugineus, 120.

Heterocera, 531.

Heterogyna, 531.

Heteromera, 550.

Heteroplax dentatus, S¢imps., 110.

Heteropora, 278.

cervicornis, D’Orb., 281.

Heteroptera, 552.

Hexactinellida, 568.

Hippa scutellata, Fabr., 506, 515.

Hippasteria plana, 45.

Hippidee, 515.

Hippidea, 515.

Hippoglossoides limandoides, B/., 446.

platessoides, Fabr., 446,

unifasciatus, Ranz.,

INDEX.

Hippolyte, 468. |

Se nncdi, M.-Edw., 454.
pusiola, Kroy., 454. /
securifrons, Norman, 454.
spinus, Sow., 45, 454, |

Hippomedon Holbolli, Kriy., 456.

Hipponyx antiquatus, Linn., 492.
Grayanus, var., Menke, 493.
radiatus, Quoy § Gaimard, 498. |

Hippothoa carinata, Norman, 452.

Hircinia, 564, 565, 566, 568. |

Hirundinide, 306, 318, 352-864, 378, |
384, 387-391.

Hirundinide, Anatomy of the North
American (Dr. R. W. Shufeldt), 352-
365. |

, Osteology of the (Dr. R. W. Shu- |
feldt), 358.

Hirundo, 352, 353, 361.

rupestris, 353.
rustica, 353.
urbica, 352, 353, 366, 368.

Histeride, 120.

Holectypus, 59.

Holocentrum longipinne, C. f V., 485.

Holorhaphidota, 568.

Holothuria intestinalis, Asc. § Lath., |
458, 470.

Hornera, 275, 276, 278, 288.
coneatennata, Feuss, 275, 278.
fissurata, Bush, 275, 276, 278, 280.
foliacea, Macg., 276.
frondiculata, Lamk., 276, 278.
galeata, Smitt, 276.

hippolyta, Defr., 276.
lichenoides, Linn., 276, 278.
ramosa, Macq., 276.
robusta, Macq., 276. .
serrata, euss, 278.
subdubia, Goldst., 276.
violacea, Sars, 276.

Hoyle, W. E., On the Deep-water |

Fauna of the Clyde Sea-area, 442.
Human races, segregate vigour and
segregate fecundity between (Gulick),
267-269.
Hyalinecia tubicola, O. ', Miill., 458,
469.

Hyas araneus, Linn., 454, 468.
coarctatus, Leach, 454, 468.

Hyastenus diacanthus, De Haan, 109.
oryx, 4. M.-Hdw., 109.
planasius, ddams § White, 109.

Hydra, ftnote, 101.

Hydractinia echinata, //em., 460,

Hydrallmania falcata, Linn., 460.

Hydrocorallina, 276.

Hydroid Zoophyte, 565.

Hydroida, 465.

Hydrophilus, 432.

581

Hydrophylide, 432, 433, 438.

Hydrozoa, 462.

Hylotomine, 34.

Hymenia perspectalis, Hiibn., 531, 545.

Hymeniacidon sanguinea, Bh., 566,
568.

Hymenoptera, 474, 531.

Hypera alba, 46.

Hypopus, 286, 287, 292-298.

Hyporrhagus marginatus, Fabr., 550.

Jache latirostris, 304.

Ianthina fragilis, Zamk., 492.

Japyx Saussurii, Hwind., 556, 557.
solifuga, Halliday, 557.

, var. Wollastoni, 557.

Ichneumon cynipiformis, Rossi, 31.

Icteridx, 510.

_ Idmonea, 276, 278.

atlantica, Forbes, 276.
concava, Reuss, 276.
eboracensis, Bush, 276.
fissurata, Busk, 275.
gracillima, Busk, 276.
irregularis, Meneghini, 279.
lichenoides, Goldf., 276.
Meneghini, Heller, 378, 279, 280.
Milneana, D’Orb., 279, 280.
notomala, Bush, 279.
radians, Lamk:., 276, 278.
radicata, Kirkp., 276.
serpens, Linn., 276, 454.

Idotea, 47.

parallela, Bate § Westw., 47, 456,
469

Idya fureata, Baird, 152.
Inachus dorsettensis, Penn., 454, 468.
dorynchus, Leach, 454.
Insecta, excepting Coleoptera, of Fer-
nando Noronha (W. F. Kirby), 530,
Tophon Pattersoni, Bowerh., 460.
Ipomeea purpurea, 264; Darwin on
Self-fertilized Generations of
(Gulick), 264.
tuba, 467.
Ischnoptera lucida, Walk., 530, 582.
Isias clavipes, Boeck, 150.
Isocardia cor, Linn., 450, 467.
Isopoda, 46, 1138.

Jacquinia, 546.
Janira maculosa, Leach, 456, 469.
Julis lucasana, 483.

noronhana, Bov/., 483.

Kapala, Camer., 32, 33.
furcata, Camer., 32, 33.
Kirby, W. F., A Synopsis of the
Genera of Chalcidide, Subfamily
Eucharine, with descriptions of

582

several new Genera and Species of
Chalcididz and Tenthredinidex, 28. |
Kirby, W. F., Insecta, excepting Coleo- |
ptera, of Fernando Noronha, 5380. |
Kirkpatrick, R., Polyzoa of Fernando
Noronha, 504.

Labidophorus talpx, Kramer, ftnote,
287.

Labidura riparia, Pall., 530, 5382.

Lafoéa dumosa, Flem., 460.
fruticosa, Sars, 460.

Lambrus laciniatus, De Haan, 109.
jlongimanus, Leach, 109.
longispinus, Miers, 109,
prensor, Herbst, 109.

Lamellibranchiata, 503.

Lamellicornia, 549.

Lantus, 312; 316,317.
ludovicianus excubitorides, ftnote,

313.

Laophonte curticaudata, Boeck, 152.
serrata, Claus, 152.

Laridx, 479.

Larridz, 531.

Lasius flavus, 118.
niger, 118, 121, 127, 181, 182, 133,

134, 136.

Latirus brevicauda, [eeve, 488.
concentricus, Peeve, 488.
gracilis, Reeve, 488.
spadiceus, [eeve, 488.

Leda minuta, Mii//., 450, 467.

, f. brevirostris, Jeffr., 450.

Leguminose, 474, 479.

Leiolophus, Miers, 513.
planissimus, Herbst, 506, 513.

Lepidoptera of Fernando Noronha

(Kirby), 531.

Lepisma, 557, 558.
corticola, Pidley, 558.

Leai, Ridley, 557.
saccharina, 558.

Lepralia cleidostoma, Simitt, 504.
depressa, Bush, 504.

Leptodius, 4. Milne-Edw., 511.
americanus, Saussure, 511.

Leptoplana tremellaris, O. 7. Mill,

458, 469.

Leptoxena, 186, 188.
eximea, Baly,-186, 188.

Lepus americanus, 141.
glacialis, 144.

Leuckartia flavicornis, Claus, 148.

Leuconia sacculata, Carter, 566, 568,

569;

Leucophzea surinamensis, Linn., 580,

539.

Leucosia craniolaris, Linn., 111.

hematosticta, Adams § White, 111.

INDEX,

Leucosia marmorea, Bell, 111.
Whitei, Bel/, 111.
Leucosiide, 111.
Leucothoé spinicarpa, Abi/d., 456.
Leucozonia cingulifera, Lamk., 487.
leucozonalis, Lamk., 487.
ocellata, Gmelin, 488.
rudis, Reeve, 487.
Libellula basalis, Burm., 537.
Jlavescens, Fabr., 537.
Libellulide, 531.
Lichenopora, 278, 280, 281, 282, 284.
bullata, Macq., 281.
californica, Busk, 282, 285.
ciliata, Busk, 283.
echinata, Macg., 281, 282, 283,
285.
fimbriata, 282.
grignonensis, Bush, 282, 283, 284.
hispida, Flem., 454. ~
, var. meandrina, Peach, 454.
Holdsworthii, Busk, 281, 282, 285.
novee-zelandix, Bush, 282.
pristis, Macq., 281, 283.
radiata, Aud., 281, 282.
reticulata, Macq., 281, 284.
verrucaria, O, Fabr., 454.
victoriensis, Waters, 284, 285.
wanganuiensis, Waters, 282.
Lichenopore, On the Ovicells of some,
by A. W. Waters, 280, 282, 283.
Ligyrocoris balteatus, S¢a@, 532, 547.
bipunctatus, Ktrb., 532, 547.
Lima elliptica, Jeffr., 450, 467.
multicostata, 499.
squamosa, var., Lamk., 499.
Limnza, 74, 83.
ovata, var. peregra, 83.
Lineus, 437.
Liparis liparis, Linn., 446.
Lirata, Camer., 33.
luteogaster, Camer., 38.
Liripora lineata, Macg., 284.
Lithodes maia, Linn., 45, 454, 468.
Littorina angulifera, Zamk., 491.
nodulosa, D’Orb., 491, 508.
trochiformis, var., Smith, 491, 503.
Loch-Leven Trout (Salmo levenensis),
On the, by F. Day, 71.
Locustidee, 530.
Lomechusa, 120.
strumosa, 120.
Longevity of Ants, Bees, and Wasps
(Lubbock), 133,
Longicornia, 555.
Longipedia coronata, Claus, 151.
Lophactea, A. M.-Hdw., 511.
lobata, Milne-Edw., 511.
Lophius piscatorius, Zinn., 446.
Lophopus, 61-64.

INDEX,

Lophopus erystallinus, 63,
Lendenfeldi, ley, 62, 64.

Lophozozymus epheliticus, Zinn., 109.

Lophyrocera, Camer., 32, 33.
nigromaculata, Camer., 32.
stramineipes, Camer., 32.

Lottia onychina, Gould, 495.

Lowne, B. T., On the Structure of the
Retina of the Blowfly (Calliphora
erythrocephala), 406.

, On the Structure and Develop-
ment of the Ovaries and their
Appendages in the Blowfly (Calli-
phora erythrocephala), 418.

Lubbock, Sir John, Observations on
Ants, Bees, and Wasps, Part XI.,118.

Lubbockia squillimana, Claus, 153.

Lucilia cesar, 125.

Lucina pecten, Lamk., 503.

Luidia Savignyi, 45.

Lumbricus terrestris, 561.

Lumpenus lampetriformis on the north
coast of Scotland, occurrence of, by
Geo. Sim, 38.

Lumpenus, 38-48.

lampetriformis, 38, 39, 41, 48.

Luperodes, ftnote, 163.

alboplagiatus, 163.
discrepens, ftnote, 163.
preeustus, ftnote, 163.

Lycenide, 531.

Lygeeidee, 532.

Lygeus anticus, Walk, 546.

rufoculis, Kird., 532, 546.

Lysianax tumida, K7rdy., 456.

Mabuia punctata, Gray, 481.

Machairopus, Zhomps., 152.
sancte-crucis, Thomps., 152, 15d.

Machilis, sp., 558, 559.

Macrochires, Studies of the, by Dr. R.

W. Shufeldt, 299-394.

Macrophthalmide, 110.

Macrophya corallipes, Hversm., 34.
femoralis, Hversm., 34.
Saundersi, W. #. Kirb., 34, 37.

Macrura, 112.

Macrurous Crustacea, 4C5.

Mactra elliptica, Brown, 450, 467.
solida, 46,
stultorum, 46.

Madreporaria of Fernando Noronha,

by Prof. P. M. Duncan, 569.

Madreporaria aporosa, 569.
fungida, 570.

Meandrina, 570.

Meera longimana, Leach, 456.
Loveni, Bruz., 456.

Maia Miersii, Walk, 109, 118, 117.

Maiide, 109,

583

Maioidea, 507.

Malacodermata, 550.

Mammalia of Fernando Noronha, 476.

Mammals and Birds of Hudson’s Bay
Company's Territory, and of Arctic
Coast of America, by Dr. J. Rae,$136.

Margaronia jairusalis, Walk., 531, 545.

Marginella fluctuata, C. B. Adams, 403.

sagittata, Hinds, 489,

Marimboudo, 541.

Mastigophora Dutertrei, Audowin, 504.

Matuta Banksii, Leach, 111.

lunaris, Herbst, 111.
victrix, Mabr., 111.

Matutida, 111.

Mbebu, name of Sula leucogastra, in
Fernando Noronha, 480.

Mecynocera, Thomps., 150.

Clausi, Zhomps., 150, 155.

Melampyrum pratense, Ants and Seeds
of (Lubbock), 133.

Melitodes, 397.

ochracea, 397.

Mella zinckenella, 7reitschke, 531, 546.

Melobesia, 565, 566.

Melophagus ovinus, 454.

Membranipora catenularia,-/J@meson,452.

Flemingi, Bush, 452.
pilosa, Linn., 452.

Menippus, 157.

Merluccius merluccius, Zinn., 446.

Meroncidius indistinctus, Walk., 536.

viridinervis, Kirb,, 530, 536.

Merula, 309, 310.

Mesomyodian birds, 387, 388.

Metagea, W. F. Kirb., 30, 33.

zalates, Walk., 30.

Metopograpsus messor, 115.

Michael, A. D., Researches into the
Life-histories of Glyciphagus domes-
ticus and G. spinipes, 285.

, On some Unrecorded Parasitic
Acari found in Great Britain, 400.

Micippa curtispina, Hasw., 109,

mascarenica, Kossm., 109.

Microphrys, M.-Edw., 507.

bicornutus, Lafr., 507.
Microporella ciliata, Pal/., 452.
impressa, Azd., 452.
violacea, Johnst., 504.
Micropus, 362-384,
melanoleucus, 303, 365, 377, 380-
383, 390, 393, 394.

Microvelia vagans, White, 548.

Migration of Organism (Gulick), 195.

Mimiecry among Ants (Lubbock), 121.

Mithraculus coronatus, A. Milne-Edw.,
510,

Mithrax, Leach, 507.

verrucosus, M.-Edw., 507.

584

Mithrax (Mithraculus)

Herbst, 510.

(Teleophrys) cristulipes, Stimps.,
508.

Mitra, 489.

Adamsi, Dohrn, 489.

ansulata, Sowerb., 488, 489.

barbadensis, Ginelin, 488.

caffra, 396.

cavea, Reeve, 489.

consanguinea, Reeve, 489.

dermestina, Lamk., 489.

formosensis, G. B. Sowerb., 395,
400.

histrio, Reeve, 489.

leucodesma, Reeve, 488, 489.

microzonias, Reeve, 488.

pardalis, Kiister, 489.

picta, Reeve, 488.

pisolina, Lamk., 489.

pulchella, Reeve, 489.

recurva, G. B. Sowerb., 396, 400.

Smithi, G. B. Sowerb., 396, 400.

tessellata, Kiener, 488.

(Pusia) ansulata, Sowerb., 488.

Mitrularia alveolata, 4. Adams, 492.

uncinata, Reeve, 492, 503.

Modiola barbata, Linn., 450, 467.

modiolus, Linn., 450.

Modiolaria marmorata, Forbes, 450.

Mollia antiqua, Simitt, 504,

Mollusca, 46, 448, 462-476.

Mollusca of Fernando Noronha, by
E. A. Smith, 488.

Mollusks, terrestrial, of
Islands (Gulick), 189.

, diversity of species not caused by
differences in environment, nor proba-
bly to sexual selection (Gulick), 189.

Molva molva, Linn., 446.

Momordica charantia, 474.

Momotus Lessoni, 547.

Monedula signata, Linn., 474, 531, 540.

Monoculodes longimanus, Spence Bate &

Westw., 456.
Stimpsoni, Spence Bate, 456.

Monolepta, Hrichs., 156, 163, 164, 178.

Cumingii, Zaly, 165, 167, 187.

foveicollis, Ba/y, 165, 167, 188.

hemorrhoidalis, Fabr., 164, 167,
187.

pauperata, 163.

terminata, Guér., 166, 167, 188.

verticalis, Baly, 167, 188.

Wallacei, Baly, 166, 167, 188.

Morphology of Aimpelis cedrorwm (Dr.
R, W. Shufeldt), 306-318.

—— of certain Cypseli and Trochili
(Dr. R. W. Shufeldt), 865-387.

Mucronella Peachii, Johnst., 452.

coronatus,

Sandwich

INDEX.

Mucronella ventricosa, Hass., 452, 454.
Multitubigera, 280.
Munida rugosa, Fabr., 454, 469.
Munna whiteaua, Spence Bate § Westw.,
456.
Murena catenata, Bl., 483.
pavonina, Pich., 483.
vicina, Cast., 483.
Murex erosus, Broderip, 486.
obeliscus, 4. Adams, 486.
pauperculus, C. B. Adams, 486.
(Ocinebra) alveatus, Kiener, 486.
(-—) erosus, Broderip, 486.
(—-—) obeliscus, A. Adams, 486.
(——) pauperculus, C. B. Adams,
486

Mus musculus, 476.
rattus, 474, 476, 477.
Musca, 408, 437, 438.
domestica, 439.
vomitoria, 408, 420.
Muscide, 532.
Muscles, pectoral, of Antrostomus and
Chordeiles (Dr. R. W. Shufeldt), 345.
Musk-rat, 142.
Mutilla, 35.
Mya truncata, Linn., 450.
Myocoptes, 400.
musculinus, 400, 401, 402, 403.
tenax, Mich., 400, 401, 406.
parasitic on field-vole (Arvicola
agrestis), 403.
Myodes hudsonicus, 143.
Myra affinis, Bell, 111.
australis, Hasw., 111.
carinata, Bell, 111.
Myriopoda of Fernando Noronha, by
R. I. Pocock, 526.
Myrmedoniz, 121.
Myrmica, 119.
leevinodis, 119.
rubra, 120.
Mysidee, 45.
Mysidopsis didelphys, Vorman, +56.
Mysis neglecta, G. O. Sars, 456.
Mytilus exustus (Lamk.), Reeve, 499,
503.
Myxilla incrustans, Johnst., 460.

Naidide, 94.

Nais, 92, 93, 97, 98.
auricularis, 94, 101.
barbata, 94.
decapoda, 93.
digitata, Miller, 92, 93, 94, 103.
quadricuspida, 94.

Nassa capillaris, Watson, 503.

Natica Alderi, Forbes, 448, 466.
Montagui, Yorbes, 448, 467.
sordida, Phil., 448, 467.

INDEX.

Natural History of the Genus Dero, by
E. C. Bousfield, 91.

Selection (Gulick), 1990.

, divergent Evolution not ex-
plained by (Gulick), 202-212.

Necrophaga, 549.

Nemoria denticularia, Walk., 531, 544.

Nephrops norvegicus, Linn., 45, 454,
469.

Neptunus pelagicus, Linn., 110.

Nerita, 493.

ascensionis, Gmelin, 493, 503.

Nesticus, 128.

speluncarum, 125.

Neuroptera, 531.

Nicotiana tabacum, 264.

Nika edulis, Risso, 454, 469.

Nitella, 474.

Noctua armigera, Hibn., 544.

Noctuz, 531.

Notes on some of the Birds and Mam-
mals of the Hudson’s Bay Company’s
Territory, and of the Arctic Coast of
America, by Dr. J. Rae, 136.

Notes on a Collection of Crustacea
from Singapore, by A. O. Walker,
107.

Notes on the Zoology of Fernando No-
ronha, by H. N. Ridley, 473. -

Nucula nitida, Sow., 46, 450, 467.

nucleus, Linn., 450, 467.
suleata, Bronn, 450, 467.
tenuis, Mont., 450, 467.

Nursia plicata, Herbst, 111.

Nyctibius, 299, 339, 351, 388.

Nyctidromus albicollis, var. Merrilli,
and Cypseloides niger, Notes on (Dr.
R. W. Shufeldt), 383.

Nyctidromus, 351, 383, 384, 388.

albicollis, var. Merrilli, 351 ftnote,
383, 384.

Nyctiphanes norvegica, M. Sars, 456,
469.

Nyctornis sthereus, 340,

grandis, 340, 343.

Observations on the Anatomy of <An-
trostomus apart from. the Skeleton
comparisons with Chordeiles) (Dr.
R. W. Shufeldt), 843-352.
Observations on Ants, Bees, and Wasps,
Part XI., by Sir John Lubbock,
118.
Ochralea, 163.
Octopus rugosus, Bosc, 484.
Ocypoda, Fadr., 512.
arenaria, Catesby, 512.
arenarius, Kingsl., 512.
Ocypodide, 512.
Odonata, 531.

585

Cicanthide, 534.
Cicanthus pallidocinctus, Kird., 530,
Dao.

Cicella, Kirb., 530, 535.
furcifera, Kirb., 530, 535.

inidea, Baly, 156, 157, 159.
ceruleipennis, Baly, 159, 187.
Jacobyi, Baly, 160, 187.
pallipes, Fabr., 161, 187.

Oithona challengerii, Brady, 151.

plumifera, Dana, 151.
setiger, Dana, 151.
spinifrons, Boeck, 151, 456.
Oligochzeta of Fernando Noronha, by W.
B. Benham, 560.
Oligocheeta, 99, 100, 103, 560.
Oliva Bulowi, G. B. Sowerb., 395,
400,
emicator, Meusch., 395.
fulgida, Reeve, 487, 503.
literata, Lamk., 487.
nivea, 487.
pulchella, Duclos, 487, 503.
(Olivella) nivea, Gmelin, 487.
Onexa obtusa, Dana, 153.
cr te neptunus, ddams §& White,
109.

Onus cimbrius, Linn., 446.
maculatus, Rzsso, 446.

Onychomorpha __lamelligera,

bid, 117.

Ophiacantha, sp., 560.

Ophidonais, ftnote, 98.

Ophiocnida brachiata, 44.

Ophiocoma echinata, Ag., 560.
nigra, O. F. Miill., 458, 470.
pumila, Liith., 560.

Ophioglypha affinis, Liith., 458, 470.
albida, Forbes, 44, 458, 470.
lacertosa, Penn., 44, 458, 470.

Ophiopholis aculeata, O. F. Miill., 458,

470.
Ophiothrix pentaphyllum, Penn., 45,
458, 470.

Ophiura cinerea, M. § Tr., 560.

Ophiuroids, 44.

Ophrys apifera, 237.

Opsibotys flavidissimalis, Grote, 5381,

546.

Orasema, Cameron, 29, 33.
flavipes, Cam., 29.
stramineipes, Cam., 29.

Oreophorus reticulatus, ddwms § White,

Lae.

Organism, migration of (Gulick), 193.

Oribatidse, 296.

Orthalicus MacAndrewi, G. B. Sowerb.,

398, 400.
Bensoni, 399.
Orthoptera, 530.

Stimps.,

LINN. JOURN.—ZOOLOGY, VOL. XX. 4b

586 INDEX,

Osteology of Chordeiles, Phalzenoptilus,
and Antrostomus (Dr. R. W. Shu-
feldt), 348-352.

of the Hirundinidz (Dr. R. W.

Shufeldt), 358.

of ‘Trogon mexicanus and T.

puella (Dr. R. W. Shufeldt), 318-

338.
Otocoris, 309, 310, 316, 317.
alpestris arenicola, ftnote 309, 312,
314.
Ovicells of Cyclostomatous Bryozoa, on
some, by A. W. Waters, 275.
of some Lichenopore, on the, by
A. W. Waters, 280.
Ovulum, 397.
Philippinarum, 397.
(Birostra) Haynesi, G. B. Sowerb.,
397, 400.
Oxalis Noronhe, 474.
Oxyrhyncha, 109.
Oxystomata, 111.

Pachygrapcus transversus, Gibbes, 110,
des

Pachyzancla detritalis, Guén., 531, 546.

Paguride, 112.

Pagurus Bernhardus, 45.

maculatus, Catesby, 512.

Prideauxi, 45.

Thompsoni, 45. saa
Paleinon earcinus, Fadr., 112.
Palemonide, 112, 518.

Palinuride, 112, 516.

Palinurus ornatus, Fabr., 112.

Palmipes membranaceus, 45.

Palpicornia, 549.

Pandalus, 465.

annulicornis, Leach, 45, 456.

brevirostris, Rathke, 456.

Pantala flavescens, Fabr., 531, 537.

Panulirus, Gray, 506, 516, 517.

echinatus, S. LZ. Smith, 516.

inermis, Kirkp., 516.

ornatus, fadbr., 516.

Papilio Hanno, Stoll, 548.

Paracalanus parvus, Claus, 148.

Paradesmus gracilis, C. Koch, 526,
528.

Parasitic Acari found in Great Britain,
on some unrecorded, by A. D. Michael,
400.

Paraulaca, 168, 188.

angulicollis, Motsch, 188.

(Aulacophora) angulicollis, Motsch,

168.

Paridea, 26.

thoracia, Baly, 27.

Parthenopide, 109.

Partula, 501.

Pasiphwa sivado, Risso, 456.

Patagial Muscles of the Pectoral Limb
of Antrostomus and Chordeiles, on
the mode of Insertion of the (Dr. R.
W. Shufeldt), 344.

Patagona, 370.

gigas, 369.

Paussidex, 121.

Pecten maximus, Linn., 450.
noronhensis, Smith, 503.
opercularis, Linn., 450, 467.
pusio, Linn., 450, 467.
septemradiatus, Miil/., 450, 467.
striatus, Mill., 450, 467.
tigrinus, Miull., 450.

Pectinaria belgica, Padl., 458, 469.

Pectinatella, 62.

Pectoral Limb of <Axtrostomus and
Chordeiles, on the Mode of Insertion
of the Patagial Muscles of the (Dr.
R. W. Shufeldt), 344.

Pectoral Limbs of certain Cypseli and
Trochili, A Critical Comparison of
the (Dr. R. W. Shufeldt), 369.

Pectoralis major, 357.

Pectunculus angulatus, 399.
crebreliratus, G. B. Sowerb., 399,

400.
pectinatus, Gmelin, 508.

Pelecanidz, 480.

Pelvic Limb in Antrostomus and Chor-
deiles, Anatomy of the (Dr. R. W.
Shufeldt), 346.

Penzide, 112, 523.

Penzeus affinis, 1.-Edw., 112.

monodon, fabr,, var. carinatus,
Dana, 112.
velutinus, Dana, 112.

Pennatula phosphorea, Linn., 44, 460.

Pentatoma testacea, Dall, 532, 546.

Pentatomide, 532.

Periceride, 507.

Perigonimus repens, Wright, 460.

. Perilampine, 28.

Perionyx, 563.
Periplaneta americana, Linn., 530,
532.
Petrochelidon, 354, 361.
lunifrons, 304, 354, 361.
Petrolisthes, Stimps., 513.
asiaticus, Leach, 515.
corallicola, var., Hasw., 112, 117.
dentata, M.-Edw., 112.
marginatus, Stimps., 513, 515.
Phaethon ethereus, Linn., 480.
Phakellura hyalinata, Linn., 531, 545.
Phalena-Geometra hyalinata, Liun.,
545.
Phalenoptilus, 322, 334, 349, 350, 351,
384, .

’
oo, oe

INDEX,

Phalenoptilus Nuttalli, 302, 389, 348,
349, 351, 384, 393.

Phanopteride, 530.

Pharomacrus mocinno, 321.

Phasianella, sp., 503.

Pheidole omnivora, Kird., 531, 589.

pusilla, Heer, 559.

Philhydrus marginellus, Fadr., 549.

Philine scabra, 46.

Philyra pisum, De Haan, 111.

Pholas crispata, 46.

Phycididee, 551.

Phycis zinckenella, Treitschke, 546.

Phylactoleamatous Polyzoa, 61.

Phyllodromia poststriga, Walk., 530,
532

Physiological Selection and Segregate
Feeundity (Gulick), 197-202.

Phytophaga, 156.

Picarizx, 388.

Picidz, 330.

Pilumnus de Haanii, Miers, 110.
labyrinthicus, Miers, 110.
vespertilio, Fadr., 110.

Pimpinella saxifrage, 82.

Pinnotheres obesus, Dana, 111.

Pinnotheride, 111.

Pisa bicornuta, Latr., 507.

Pisania pusio, Linn., 486.

Pisces, 446, 462, 464.

Pisces of Fernando Noronha, by G. A.

Boulenger, 483.

Pisum sativum, 237, 265.

Plagusia, Latr., 513.
depressa, Yabr., 513.

Planorbis, 475.
fragilis, Brazier, 502.

Gilberti, Dunker, 502.
noronhensis, Smith, 502, 503.
Platalea ajaja, 580.
Platyarthrus, Experiments with (Sir J.
Lubbock) 124.

Platyarthrus Hoffmanseggii, 136.

Platypus parallelus, /’., 554.

Platyxantha, Baly, 158.
clypeata, Baly, 158-187.

Plecotrema, 501.

Plesiofungide, Dwnc., 570.

Pleuromma abdominale, Claus. 148.

Pleuronectes cynoglossus, Linn., 44,

446

flesus, Linn., 448.

limanda, Linn., 448.

microcephalus, Donov., 448.

platessa, Linn., 448.
Pleurotoma cuprea, Reeve, 485.

fuscescens, Gray, 485.

nebula, 395.

nigrescens, Gray, 485.

' paxillus, Reeve, 4805.

587

Pleurotoma solida, C. B. Adams, 485.
turricula, Mont., 46.
(Bela) turricula, Mont., 448, 466.
(Crassispira) fuscescens, Gray,
485

(Mangelia) Brione, G. B. Sowerd.,
395, 400.
Plumatella, 61, 62.
Aplini, Macgill., 64.
repens, var. @, Adlm., 64.
Plumularia Catharina, Johnst., 460.
pinnata, Linn., 460.
Pocock, R. I., Crustacea of Fernando
Noronha, 506.
oo of Fernando Noronha,
526.
Podargus, 299, 339, 351, 388.
gigas, 340.
Podoceropsis Sophia, Boeck, 456.
undata, Spence Bate, 456.
Podophthalmia, 109, 113.
Podophthalmus vigil, Fabr., 110.
Polistes, 474.
instabilis, Sawss., 541.
Ridleyi, Kirb., 531, 541.
called Marimboudo in Fernando
Noronha, 541.
Pollen, Darwin on the influence of Po-
tency in (Gulick), 242.
Polycarpa pomaria, Sav., 448, 466.
Polycremuus ochtodes, Herbst, 110.
Polyergus rufescens, 118, 120.
Polyonyx cometes, Walk., 112, 116,
117.
obesulus, White, 112.
Polyplacophora, 503. , .
Polytherses, Duchass. e¢ Mich., 564, 566,
568, 569.
Polytrema, 564, 560.
miniaceum, 564.
Polytrichum, 44.
Polyzoa of Fernando Noronha, by R.
Kirkpatrick, 504.
Polyzoa, 61, 452, 462, 464, 465, 504.
Pomacentrus leucostictus, MW. 4 T.,
483.
Pompilide, 531.
Pompilus nesophilus, Kirb., 474, 531,
540,
sobrinus, Blanch., 540.
Pontobdella muricata, Linn., 458
Porania pulvillus, O. F. Miill., 45,

458.
Porcellanella picta, Stimps., 112, 117.
(Petrolisthes) corallicola, var.,

Hasw., 118.
Porcellanidx, 112, 513.
Porcellanidea, 513.
Porcellidium viride, Philippi, 152.
Porella compressa, Sow,, 452.

588 INDEX.

Porifera of Fernando Noroxha, by H.
J. Carter, 564.
Porifera, 460, 462.
Portunide, 110.
Portunus, 45.
depurator, Linn., 45, 454, 468.
holsatus, Mabr., 45, 454.
marmoreus, Leach, 454.
pusillus, Leach, 45, 454, 468.
Prairie-hen, 139.
Priapulus caudatus, 45, 47.
Pristiurus melanostomus, Bonap., 446.
Progne, 352-366.
subis, 301, 304, 352, 355, 358, 365,
393.
Proto, Dutrochet, 94, 103.
Psalurus, 299, 351, 388.
Psammobia Ferroensis, 46.
Psammonemata, 568.
Pseudocalanus elongatus, Boeck, 148,
456.
Pseudocophora, Jacoby, 29, 168.
ambusta, Hrichs., 169, 170, 172,
174, 175, 188.
bicolor, Jacoby, 169.
brunnea, Baly, 26, 169, 170, 173,
174, 188.
Buquetii, Guér., 169, 170, 171, 172,
188.
distineta, Baly, 169, 171, 188.
Erichsoni, Baly, 169, 171, 188.
flaveola, Baly, 169, 173, 188.
pectoralis, Baly, 169, 174, 188.
perplexa, Baly, 170, 175, 188.
uniplagiata, Jacoby, 169, 173, 174,
188.
Wallacei, Baly, 169, . 173, 174,
188.

Pseudotrimera, 556.
Psilogaster, Blanch., 80, 33.
cupreus, Blanch., 80.
pallipes, Brudlé, 30.
Psilopus metallifer, Watk., 474, 532,
548.
Psclus phantapus, Strussenf., 458.
Pterylographical tracts of Antrostomus
and Chordeiles, on the (Dr. R. W.
Shufeldt), 859-343.
Puncturella Noachina,
467.
Pupa pellucida, Pfeiffer, 501.
solitaria, Smith, 501, 5038.
Purpura accincta, @’ Orb., 486.
hemastoma, Linn., 486.
Pyecnarthrum gracile, Hichh., 554.
setulosum, Waterh., 558.
Pycnogonum littorale, 46.
Pygidicrana notigera, S¢d/, 530, 532.
Pyrales, 531.
Pyralis manihotalis, Guén., 531, 545.

LTinn., 450,

Pyralis perspectalis, Hiibn., 545.
Pythia inflata, Pfeiffer, 501.

Rae, Dr. J., Notes on some of the Birds
and Mammals of the Hudson’s Bay
Company’s Territory, and of the
Arctic Coast of America, 136.

Raja clavata, Linn., 446.

fullonica, Linn., 446.
maculata, Montagq., 446.

Ranunculus aquatalis, 82.

Recognition of Friends in Ants, Bees,
ancl Wasps (Lubbock), 128.

Relation between Formica sanguinea
and its slaves, 118.

Remips, Latr., 515.

cubensis, de Sauss., 515.
scutellatus, Mabr., 515.

Reptilia of Fernando Noronha, by G. A.
Boulenger, 481.

Researches into the Life-histories of
Glyciphagus domesticus and G. spi-
nipes, by A. D. Michael, 285.

Rhagovelia incerta, Kirb., 532, 548.

Rhaphidonemata, 568.

Rhincalanus cornutus, Dana, 148.
gigas, Brady, 148.

Rhipipallus, W. F. Kirb., 31, 33, 37.
Cameroni, W. F. Kirb., 37.
volusus, Walk., 31, 37.

Rhizopide, 110.

Rhomboidichthys lunatus, Linn., 483.

Rhombus megastoma, Donov., 446.
norvegicus, Giimther, 446.
punctatus, B/., 446.

Rhopalocera, 531.

Rbhynchophora, 553.

Rhynchopora bispinosa, Johnst., 504.

Ridley, H. N., Notes on the Zoology of
Fernando Noronha, 473.

, Thysanura and Collembola of
Fernando Noronha, 556.

Ridley, S.O., On the Characters of the
Genus Lophopus, with description of
a new species from Australia, 61.

Rissoa, sp., 503.
abyssicola, Yorbes, 448, 466.

Rock-grouse, 140.

Rossia Oweni, Ball, 448.

Rotifera, 97.

Rusticus adolescens Hanno, Hiibn., 548.

Sabella pavonia, Sav., 458, 469.

Saccharissa, W. F. Kirb., 87.
contingens, Walk., 37.

Salarias atlanticus, C. g& V., 483.
vomerinus, C, f& V., 483.

Salmacis, 58.

INDEX. 5S9

Salmo albus, 90.
ceecifer, Parnell, 71.
cumberland, Lacép., 74.
fario, 71, 75, 76, '79, 86, 90.
ferox, ftnote 84, 86.
iridens, ftnot», 87.
lemanus, 75, 90.
levenensis, Walk., 71, 72, 76.
salar, ftnote, 71.
taurinus, Walk., 71, 72.

Salmon-trout, ftnote, 71.

Samea castellalis, Guér., 581, 545.

Sand-wasps, 474. }

Sapphiriva inzqualis, Dana, 153.
metallina, 153.
serrata, Brady, 153.

Sarcophaga calida, Wiedem., 532, 548.
carnaria, 420.

Sarcoptide,. 401, 405.

Saxicava rugosa, Linn., 452, 467.

Sayornis, 315.

Sealaria hellenica, Forbes, 503.

Scalpellum vulgare, Leach, 456.

Scaphander lignarius, Linn., 448.

Scapteriscus abbreviatus, Scudd., 530,

533.

Schizaspidia, Westw., 31, 35.
furcifera, Westw., 31.
maculata, Westw., 29.
Murrayi, Kirb., 37.
plagiata, Walk., 31.

Schizophrys aspera, A. M.-Hdw., 109,

Lis.

, var. spinifrons, A, M.-Edw.,
113.
Schizoporella unicornis, Johnst., 452,

Scolecithrix Danze, Lubbock, 148.
minor, Brady, 148.
Scolopendra morsitans, Linn., 526.
Scoparia dulcis, 474.
Scrobicularia prismatica, 46,
Scrupocellaria, 504.
frondis, Kirkp., 504, woodcut,
505.
reptans, Linn., 452.
Seutellidum tisboides, Claus, 158.
Scyllarid, 112.
Seymus, sp., 556,
xeranipelinus, Muls., 556.
Seythrops, 338. ©
Sea-pen, 44.
Segregate breeding (Gulick), 192-195.
Segregation, Causes of Cumulative
(Gulick), 222-274.
_——, Cumulative divergence through
Cumulative (Gulick), 212-222.
, Spencer’s Law of (Gulick), 195.
Seira Buskii, Ludd., 559.
musarum, idley, 559.

Selandria meridionalis, W. F', Kirb., 34,
37

Selandriinx, 34.

Selection, effects of, and independent
generation contrasted (Gulick), 202-
212.

Semele chinensis, Sowerd., 498.
cordiformis, Chemnitz, 498.
lenticularis, Sowerd., 498.
luteola, Reeve, 498.
luteola, A. Adams, 498.
modesta, Reeve, 498.
modesta, A. Adams, 498.
nitida, Miill., 452, 468.
reticulata, Sowerb., 498.
sinensis, A. Adams, 498.
subtruneata, Reeve, 498.
tenuis, Mont., 452, 468.

(Abra) alba, Wood, 452, 468.

Separation and Variation of species
(Gulick), 191.

Sepiola Rondeleti, Leach, 448.

Serpula vermicolaris, Zinn., 458, 469.

Serricornia, 550.

Sertularella fusiformis, Hincks, 460.
Gayi, Lamzx., 460.
rugosa, Linn., 460.

Sertularia abietina, Linn., 44, 460.
argentea, Lill. § Sol., 460.

Sesarma Bocourti, 4. M.-EHdw., 110,

LUZ.

Sessile-eyed Crustacea, 46, 74, 83.

Setella gracilis, Dana, 151.

Sharpe, R. Bowdler, Aves of Fernando
Noronha, 477.

Shells, descriptions of fourteen new
species of, by G. B. Sowerby, 395.
Shufeldt, Dr. R. W., Studies of the
Macrochires, Morphological and
otherwise, with the view of indi-
cating their Relationships and de-
fining their several Positions in the
Systein, 299.

Siderastraea siderea, Ellis g Soland.,

570.
stellata, Verrill, 570.

Silpha, 126.
thoracica, 126.

Silver-grey trout, 72.

Sim, -Geo., Occurrence of Zumpenus
lampetriformis on the North Coast
of Scotland; with Notes on its
Habits, Food, and the Ground it
frequents, 38.

Siphonaria brunnea, Hanley, 497.

hispida, Gould, 497.
lineolata, d’ Orbigny, 497.
picta, var., Smith, 497, 503.

Siphonodentalium tetraschistum, Wat-

son, 503.

590 INDEX.

Sitophilus oryzx, Linn., 553.

Sitoplidus oryze, 475.

Skink, 475, 476, 481.

Sladen, W. Percy, and Prof. P. M.
Dunean, On the Anatomy of the
Perignathic Girdle and of other
Parts of the Test of Déscoidea cylin-
drica, Lamk., sp., 48.

Slavina, ftnote, 98.

Smell in Ants, Bees, and Wasps (Lub-
boek), 125.

Smith, E. A., Mollusca of Fernando
Noronha, 483.

Smittia reticulata, MacGill., 452.

Smittipora antiqua, Busk, 504,

Snow-goose, 138,

Solarium, sp., 503.

Solaster endeca, Gm., 458.

Solea variegata, Donov., 448.

Solen pellucidus, Penn., 46, 450.

Sowerby, G. B., Descriptions of four-
teen new species of Shells, 395.

Spatangus purpureus, 4. Ff. Miill., 45,
458, 470.

Spencer, H., Law of Segregation (Gu-
lick), 195.

Speotyto, 323.

Spermacoce, 479.

Sphagnum, 44.

Spirastrella cunctatrix, Sd7., 565.

Spirobolus, 526.

noronhensis, Pocock, 528.
paraensis, Humb. § Sauss., 529.

Spondylus, sp., 500.

Sponges, 465.

Spongillide, 64.

Spongiophaga communis, 564.

Spotted Gunnel, 39.

Squilla chiragra, Fabr., 526.

nepa, Fabr., 113.

Stalk-eyed Crustacea, 45.

Steatornis, 299, 339, 351. 352, 388.

Steganoporella Smittii, Hincks, 504.

Stelgidopteryx serripennis, 305, 362.

Stelletta, 565.

Stenogyra (Opeas) Beckiana, var.,

Smith, 502.
(
502

(——) subula, Pfeiffer, 592.
Stenopola dorsalis, Thunb., 530, 536.
Stenopusculus, Jeichters, 506, 523.

crassimanus, Richters, 523, 525.

spinosus, Kirkp., 506, 523.
Stenorhynchus longirostris, M.-EHdw.,

454, 468.

Stenorhynchus rostratus, Linn., 454,

468.
Stenothoé monoculoides, Mont., 456.
Stichzeus lampetraeformis, 446.

) octonoides, C. B. Adams,

Stichaster roseus, O. F. Miill., 45
458.
Stilbula, Spin.; 31, 33. —
cynipiformis, Fossi, 31, 37.
Stomatella nigra, Quoy & G., 503.
Stomatoceras, 36.
diversicornis, VW. F. Kirb., 36, 37.
Magretti, W. F. Kirb., 35, 36,
3

the

Stomatopoda, 113, 526.

Stomatopora granulata, M.-Edw., 454.

Strategus anteeus, Fadr., 550.

Strongylocentrotus, 58.

Studies of the Macrochires, Morpholo-
gical and otherwise, with the view of
indicating their Relationships and
defining their several Positions im
the System, by Dr. R. W. Shufeldt,
299.

Sturnella, 310.

Styela erossularia, V. Ben., 448, 466.

rustica, Linn., 448, 466.
Stylaria, ftnote, 99.
Suberites ficus, Johnst., 460.

massa, Sd¢., 566, 568.

suberea, Mont., 460.

Sula leucogastra, 480.

Survival of the fittest (Gulick), 189.

Symbiotes, 401, 402.

setiferus, 404.

tripilis, Mich., 403, 406; para-

sitic on hedgehog (Hrinaceus
europeus); 404.

Synascidia, 565.

Synnotum aviculare, Pieper, 504.

Syrphide, 532.

Syrphus vesiculosus, Fabr., 548.

Tachycineta, 361, 362, 364.
bicolor, 805, 354, 361.
thalassina, 305, 354, 361, 362, 364,

394,
Tachypetes aquila, 480.
Tachytes inconspicuus, Kirb., 474, 531,
540.
iridipennis, Smith, 540.

Tapinoma, 129.
erraticum, 121.

Tarucus Hanno, Séod/, 531, 543.
monops, Zell., 543.

Tectura fulva, Mill., 450.

Teleophrys cristulipes, Stimps., 508,

509.

Tellina cordiformis, Chemnitz, 498.
decussata, Wood, 498.
fabula, 46.
reticulata, Linn., 498.
sordida, 452.

Temnocera vesiculosa, Fadr.,

548.

474, 582,

INDEX. 591

Temnopleurus, 58.
Temora dubia, Lubbock, 150.
longicornis, Miill., 456.

Tenthredinide and Chalcidide, On
Genera and Species of, by W. F.
Kirby, 28, 34.

Tenthredinine, 34.

Terebrantia, 531.

Terebratulina caput-serpentis, Linn.,
452, 468.

Tethya cranium, 566, 568.

Tetragonaspis flavicollis, Mayr, 538.

gracilicornis, Mayr, 538.

Tetramelia, W. F. Kirb., 31, 38.

plagiata, Walk., 31.
Tetrao mutus, ftnote, 140.

rupestris, 140.

saliceti, 139, 140.

Thalamita sima, M.-Edw., 110.

Thalassina anomala, Herbst, 112.

Thalassinide, 112.

Thalassinidea, 515.

Thalestris Mysis, Claus, 152.

rufocineta, Norm., 152.

Thalotia Wilkiz, G. B. Sowerb., 397,
400.

Thenus orient#lis, Herbst, 112.

Thermesia gemmatalis, Hiibn., 531,
544.

Thompson, J. C., Copepoda of Madeira
and the Canary Islands, with De-
scriptions of new Genera and Species,
145.

Thoracantha, Latr.. 32, 33.

atrata, Walk., 32.
coleopteroides, Waterh., 32.
furcata, Haliday, 32.
Latreillii, Guér., 82.
striata, Perty, 33.

Thuiaria thuia, 44.

Thyone fusus, O. FV’. Miill., 458, 470.
Thysanura and Collembola of Fernan-
do Noronha, by H. N. Ridley, 556.

Tlos muriger, Adams § White, 111.

Tomicus, 553. .

Tomigerus, 484, 501.

Torinia xthiops, Menke, 491.

cyclostoma, 492.
cylindracea, 492.
nubila, 492.

Tornatella fasciata, 46.

Trachea, Visceral Anatomy, and other
parts, On the (Dr. R. W. Shufeldt),
358.

Tragosia infundibuliformis, Johnst.,
460.

Tramea basalis, Burm., 531, 537.

Trapezia cymodoce, Herbst, 110.

Trematodes, 437, 488, 459.

‘Trichoxenia cineraria, Walh., 35, 37.

Tricoryna, W. F. Kirb., 29, 33.
Trigla gurnardus, Linn., 446.
Triglops Murrayi, Gunther, 446.
Tringa alpina pacifica, 140.
Tripneustes esculentus, Leske, 560.
Triton, 490.
Cantrainei, Récluz, 486.
gallinago, Peeve, 490.
obscurus, Reeve, 490.
pilearis, Lamk., 490.
Ridleyi, Smith, 489, 503.
testudinarius, Adams g¢& Reeve,
490.
(Epidromus) testaceus, Morch,
490

Trochili and Cypseli, A Critical Com-
parison of the Pectoral Limbs of cer-
tain (Dr. R. W. Shufeldt), 369.

, Notes on the Anatomy of

the Pelvic Limb in certain (Dr. R. W.

Shufeldt), 378.

, Morphology of certain (Dr.
R. W. Shufeldt), 365-387.

Trochilidx, 299, 301, 306.

Trochilus, Anatomical Notes upon the
Nestling of (Dr. R. W. Shufeldt),
285.

Trochilus, 365-392.

Alexandri, 303, 377.

Anna, 303.

Calliope, 304, 382, 394.

colubris, 303, 380, 381.

moschitus, 366, 368.

platycerus, 303, 365, 3868, 370, 374,
381, 394.

rufus, 304, 375, 377, 378, 394.

Trochus digitatus, Reeve, 493.

magus, Linn., 448.

nobilis, Philippi, 494.

Olfersi, Troschel, 493.

(Eutrochus) jujubinus, Gmelin,
493, 494.

) gemmosus, /?eeve, 494.

(Gibbula) cinerarius, Zinn., 448,
467.

(Infundibulum) baccatus, G. B.
Sowerb., 897, 400.

(Zityphinus) millegranus, Phil.,
448, 467.

) zizyphinus, Linn., 448, 467.

Trogon, 300-838.

Alexandri, 333.
ambiguus, 318.
mexicanus, 302, 306, 318-338, 387,
393.
puella, 302, 306, 318, 321-827,
333, 387, 388.
Trogon mexicanus and T. puella, Oste-

ology of (Dr. R. W. Shufeldt), 318-
338.

592 INDEX.

Trogonide, ftnote 300.
Trogophleeus pusillus, Grav., 549.
Trout, bill, ftnote 71, 72.
grey, ftnote 71, 72.
Loch-Leven, 71-90.
salmon-, ftnote 71.
silver-grey, 72.
Truxalis dorsalis, Thunb., 536.
Trypanidius dimidiatus, 7%., 555, 556.
isolatus, Waterh., 555.
melancholicus, 556.
Tryphosa longipes, Spence Bate, 456.
Tubifex, ftnote, 99. .
Tubularia indivisa, Linn., 44, 460.
Tunicata, 444, 448, 462, 464, 466, 471,
472.
Turbo digitatus, Deshayes, 493.
imbricatus, Sowerd., 493.
unguis, Wood, 493.
(Calear) Olfersi, Zroschel, 493.
Turricula, 396.
Turritella terebra, Linn., 448, 466.
Typhlocarcinus villosus, Szémps., 110,
Wg
Tyrannidex, 301, 309, 310, 311, 312,
313, 316, 478.
Tyrannus, 516.
tyrannus, 356.
verticalis, 809, 314, 315, 392.
Tyrant Flycatcher, 310.
Tyroglyphidex, 296.
Tyroglyphus, 286, 287, 295, 296, 297.
siro, Linn., 287.

Unicavea californica, D’Orb., 282.

Urobenus, 561.

Urocheeta, Perrier, 561, 562, 563.
corethrura, /. Miiller, 562, 563.
dubia, Horst, 562, 563.

Uromelia, W. F. Kirb., 33.
striata, 33.

Uronais, 94.

Utriculus canaliculatus, Say, 503.

Variation and Separation of Species
(Gulick), 191.

Velia collaris, Burm., 548.

Veliidae, 532.

Velutina levigata, Penn., 445, 466.

Venus fasciata, DaC., 450.

Venus lincta, 46.
ovata, Penn., 450, 467.
(Anaitis) paphia, Zinn., 497.
Vermes, 458, 462-472.
Vesicularia spinosa, Linn., 454.
Vespa signata, Linn., 540.
Vespide, 551.
Vireo gracilirostris, Sharpe, 478.
magister, 478.
Vireonide, 478.
Virgularia mirabilis, O. F. Mi/l., 460.
bianca, 480.
preta, 479.
Viuva preta, 479.
Viviparous Blenny, 41.

Wagner, Moritz, On the Law of the
Migration of Organisms (Gulick),193.

Walker, A. O., Notes on a Collection
of Crustacea from Singapore, 107.

Wasps, Ants, and Bees, Observations
on, by Sir John Lubbock, 118-136.

Waterhouse, Chas. O., Coleoptera of
Fernando Noronha, 548.

Waters, A. W., On some Ovicells of
Cyclostomatous Bryozoa, 275.

, On the Ovicells of some Licheno-
pores, 280.

Waxwings, 301.

Westwoodia nobilis, Baird, 152.

Westwoodilla caecula, Spence Bate, 456.

Willow-grouse, 159.

Witch-sole, 44, 45, 46.

Xantho, Dutroch., 93, 103.

decapoda, 93.

hexapoda, 93.

scaberrimus, Walk., 109, 118, 117.
Xenophora corrugata, ecve, 5038.
Xylopertha, sp., 550.
Xylophaga, 553.

Zenaida aurita, Gray, 479.
maculata (V.), Sel. & Salv., 479.
noronha, Gray, 479.

Zoarces (Viviparous Blenny), differen-

tiative characters in, 41, 42.

Zoarces, 41, 42.
viviparus, 39, 41.

Zoophyta, 44.

END OF THE TWENTIETH VOLUME.

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET.

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THE JOURNAL

THE LINNEAN SOCIETY.

Vou. XX. ZOOLOGY. : Nos. 124-125.

CONTENTS.

Notes on the Zoology of Fernando Noronha. By H. N.
iipier, oA. PGS. (Plate XXX.) ....00...0.0..0.05. 478

Titlepage, Contents, and Index to Volume.

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‘O68 T “FT Avy

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NOTICE.

—_— > —-

THE seeming irregularity in the issue of Numbers of the
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Journal of Zoology.

Vol. XX. is completed with the issue of the present number,
and contains Nos. 116 to 125 inclusive, with Title and Index to
the Volume.

Vol. XXI.— Devoted to the Fauna of the Mergui Archi-
pelago, comprises Nos.'126 to 135 inclusive, all of which have
been published, with Title and Index to the Volume.

Vol. XXII., devoted to Dr. De Man’s Report on the Crustacea
of the Mergui Archipelago, comprises Nos. 136 to 140 inclusive,
all of which have been published, with Title and Index to the
Volume.

Fellows are requested to compare these statements with their
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For issues of Botany, see Notice on back of the Botanical
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UNIVERSITY OF ILLINOIS-URBANA

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ae JOURNAL OF THE LINNEAN SOCIET

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