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JOURNAL

OF

THE LINNEAN SOCIETY,

ZOOLOGY.

VOL. XXXV.

2 T3206

LEFORNADIOINT:
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON. HOUSE,
PICCADILLY, W. 1,
AND BY
LONGMANS, GREEN, AND CO.,
AND
WILLIAMS AND NORGATE,
1922-1924,

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

No. 231, ' pp. 1-79,
PAR NTL
» 143-216,
oR. Ol aiae),
293-892,
926, ,, 393-447,
237, 4, 449-579,
, 238, ,, 581-672,

(Index, Title, etc.),

published June 12, 1922.

m9.

September 30, 1922.
March 3, 1923.
June 5, 1923.
August 15, 1925.
December 31, 1928.

February 14, 1924,

April 16, 1924.

PRINTDD BY TAYLOR AND FRANCIS,

RED LION COURT, FLEET STREET,

ISH IP Jew ede esis)

Page
Aurson, A. M. On the Method of Oviposition and the Egg of Lyctus
brunneus Steph. (Communicated by Dr. A. D. Inms, F-.L.S8.)
(Blatia 12), arnal B WersiciNeAMires,)) oococcesaconcancascansonens cesosogbsoesbbcossqbon 217

—— On the Genital System of Lyctus branneus Steph., with a Note on
Lyctus linearis Goeze (Coleoptera). (Communicated by Dr. A. D.
Tus, F.L.8.) (Plates 31-34, and Text-figures 1-4.) ...................5. 581

Barsson, Dr. WittiAM, F.R.S. See GArsranc, WALTER.
Borrapaius, Lancenor Anmxanper, Se.D., etc. On the Mouth-parts of the
Shore Crab. (Communicated by Prof. E. 8. Goopricu, F.R.S., Sec..8.)

(Plewsas WO, Wo) ocoosooceoseconsacasdonnenspnesc0n50an9n9nseoace09sn0000r ree Ararat 115

Bourne, Ginpert Cuarues, M.A., D.Sc., F.R.S., F.LS., late Fellow of Merton
College, Oxford, and Professor of Zoology and Comparative Anatomy.

The Raninide: a Study in Carcinology. (Plates 4-7.) .................. 25
Caiman, Winuiam Tuomas, D.Sc., F.R.S., Sec.L.8. See Hauperr, J. W.
CuarK, Huserr Lyman, Museum of Comparative Zoology, Cambridge, Mass.,
U.S.A. Some Echinoderms from West Australia. (Communicated by
IPO, We Vo IDATEIEN, TNILYS),)) (IPleuie) 18},) oucceasadososadvaopascanadsucespaasoune 229
Connincr, Watrer E., D.Se., F.L.8., Keeper of the Yorkshire Museum, York.
On the Terrestrial Isopod Hlwma ccelatum (Miers) = purpurascens,
Bucléle-Iwroel, (IPMN) 5) naosododo4os0co0baocncdnos0coonddcaxcaacaddnapaapeoseedoe 103
—— On Two new Terrestrial Isopods from Madagascar. (Plate 9.) ......... 107

Currier, Donatp Warp, M.A. See Sanvon, H.

Dakin, Wituiam Joun, D.Sc., F.L.S. See Cuark, H. L.; Hickson, 8. J.;
O’Donocuus, C. H.; and Tarrersarn, W.

Denpy, Arraur, D.Sc., F.R.S., F.L.8., Professor of Zoology in the University
of London (King’s College), and Lustre M. Frepericx, M.Sc., Harold
Row Student in the Zoological Department, King’s College. On a
Collection of Sponges from the Abrolhos Islands, Western Australia.
(Blates 25-26!) oo eee cece eee eee cee ete mentees serena 477

iv
Earuanp, Artuur. See Heron-Auuen, H., F.R.S., F.LS.
' Freperick, Miss Lestiz M., M.Se. See Dunpy, A.

Frew, J. G. H., M.Se., Ministry of Agriculture Research Scholar. On the
Morphology of the Head Capsule and Mouth-parts of Chlorops taniopus
Meig. (Diptera). (Communicated by Dr. A. D. Inms, F.L.8.) (With
CC ANGy anne inHe}sh) eo nanadoronduacecabnodudasecauas dobnbonsnmnpRosdndedtosooadongsonbeS
Garsranc, Waurer, M.A., D.Sc. (Oxon), Professor of Zoology in the Uni-
versity of Leeds. The Theory of Recapitulation: a Critical Re-
statement of the Biogenetic Law. (Communicated by Dr. W. Bateson,
TDGIRsShos Holts) (Uiatiols @ Were.) o0003-codoeoaocnpoansocvoogcuad0704000
Gincertist, Joun Dow Fisumr, M.A., D.Sc., F.L.8., Professor of Zoology in the
University of Cape Town. A Form of Dimorphism and Asexual
Reproduction in Péychodera capensis (Hemichordata). (With 7 Text-
TOUTES VI ole ee ABOU RRR AGA MUR ae ea ee ab Ee ee cr cam Oe ae

Git, E. L., M.Sc. See Warson, D. M.S.

Goopricn, Epwarp SrepHen, M.A., F.R.S., F.L.8. See Borrapairn, L. A.,
and Huxtry, J. 8.

Gurney, Roperr, M.A., F.L.8. The Crustacean Plankton of the English
Lake District. (Plate 23, and 3 Text-figures.) .............:..ceesceenencees

Havpert, J.N.,M.RJ.A. Notes on Acari, with Descriptions of New Species.
(Communicated by Dr. W. T. Canmay, F.R.S., F.L.S.) (Plates 20-22.)...

Harmer, Sir Sipyey F., K.B.E., Se.D., V.P.R.S., F.L.S., Director of the
Natural History Departments of the British Museum. On Cellularine
eval @ulaeie lohan, (CEES IG=1G))) | foossseaseqenoscns cneossdanoboasadaenaone

Heron-ALLen, Epwarp, F.R.S., F.L.S., and A. Eartanp, F.R.M.S. The
Foraminitera of Lord Howe Island, South Pacific. (Plates 35-37). ...

Hickson, SypNry Joun, Professor of Zoology, University of Manchester. On
Two Sea-pens from West Australia. (Communicated by Prof. W. J.
LD) sehg Ul Ore) Mme nanunaHeBe Ads satan dubnansooandceausadecaueeraneBnacnddaatocced aac

Huxney, Junttan Sorent, M.A. Courtship Activities in the Red-throated
Diver (Colymbus stellatus Pontopp.); together with a discussion of the
Evolution of Courtship in Birds. (Communicated by Prof. B.S. Goop-
rich, M.A., F.R.S., Sec.L.S.) (Plates 14, 15, and 4 Text-figures.)

Page

399

81

393

All

363

293

599

. 253

Imus, Aucusrus Danret, M.A., D.Sc. See Aurson, A. M.; Frew, J. G. H.

O’Donocuun, Cuarues H., D.Sc., F.R.S.C., Professor of Zoology, University of
Manitoba, Canada. Report on Opisthobranchiata from the Abrolhos
Islands, Western Australia, with Description of a new parasitic
Copepod. (Communicated by Prof. W. J. Dakin, D.Sc., F.L.S.)
(Blates:2 7230S) ype opemtr cc reer mee eri tactics os slisaiereiie Ganatsta s'erouarclactecie st

Sanpon, H. Some Protozoa from the Soils and Mosses of Spitsbergen.
Results of the Oxford University Expedition to Spitsbergen, No. 27.
(Communicated by D. Warp Curer, F.L.8.) (Plate 24, and 6 Text-
HT SMALE) eeetoess era eistSeteclefe ciorae atte oatelsye siciya tere eeaclae lors aisles lalostonclaioetstialldiarn geeistres

Tarrersati, WiitiAM, D.Se., Keeper of the Manchester Museum. The Perey
Sladen Trust Expeditions to the Abrolhos Islands (Indian Ocean),
under the Leadership of Prof. W. J. Daxry, F.L.S., &.Z.5.—Amphipoda
and [sopoda. (Communicated by Prof. W. J. Daxty, F.L.8.) (Plates
l= }a) el gubaadeetiognes¢e dace ad aa aud con GneeCeaa mm anth cere srACnC Ren np panera car Anecoannn

TintyarD, Rosin Joun, M.A. (Cantab.), D.Sc. (Sydney), F.L.S., C.M.Z.S.,
F.E.S., Entomologist and Chief of the Biological Department, Caw-
thron Institute, Nelson, N.Z. The Wing-Venation of the Order
Plectoptera or Mayflies. (With 10 Text-figures.)........... .2.cceceeee eee

Watson, Davip Merrepira Seargs, F.R.S., and E. L. Girt, M.Se. The
Structure of Certain Paleozoic Dipnoi. (Communicated by Prof. E. 8.
GoopricH, F.R.S., Sec.L.8.) (With 34 Text-figures.) .......... Sa0boso00n

Page

521

449

143

16-19.
20-22.

25, 26.
27-80.
31-34.
35-37.

vi

EXPLANATION OF THE

Abrolhos Amphipoda and Isopoda.

Raninide.

Eluma celatwm (Miers).

Isopoda from Madagascar.

Mouth-parts of Shore Crab.

Lyctus brunnews Steph.

Echinoderms from West Australia.
Louisiana Heron, Hydranassa tricolor.

Little White Egret, Hgretta candidissima.
Two female Whitethroats, Sylvia cinerea.
Willow-Warblers, Phylloscopus trochilus.
Polyzoa.

Species of Acari.

Crustacean Plankton from the English Lakes.
Protozoa from Spitsbergen.

Sponges from the Albrolhos Islands.
Opisthobranchiata from the Abrolhos Islands.
Genital System of Lyctus brunneus.
Foraminifera from Lord Howe Island.

PayAw, Eis

vil

ERRATA.

Page
47, line 21, for Thiers, read Miers.
219, ,, 6, read Robinia Pseudo-Acacia.
245, ,, 7 from bottom, for Pentanogaster read Pentagonaster.
288, ,, 30, read extremely.

i, 4
577, ,, 177 read DaKInt.
578, ,, 40

Addenda to “ The Foraminifera of Lord Howe Island.”
Page 632, after line 12, insert
156 a. DiscorBinA Dimrp1ata Parker & Jones.
Discorbina dimidiata Carpenter, Parker & Jones, 1862, ISF. p. 201, fig. 328.

A 5 Parker & Jones, 1865, NAAF. pp. 385 & 422, pl. 19.
fig. 9.
py “a Heron-Allen & Harland, 1914-15, FKA. p. 698. 3

Frequent. The specimens are rather smaller than those characteristic of
Australian shore-sands, but are quite typical.

Page 636, before line 13 from bottom, insert

175a@. PULVINULINA MENARDII (d’Orbigny).
Rotalia menardii dOrbigny, 1826, TMO. p. 273, no. 26, Modéle no. 10.
Pulvinulina ,, Brady, 1884, FC. p. 690, pl. 108. figs. 1, 2.
ip » Heron-Allen & Harland, 1914-15, FIA. p. 715.
A single small pelagic specimen.

Plates 27, 28, in legend read OPISTHOBRANCHIATA.

JUNE 12, 1922. Price 12s.

THE JOURNAL

OF,

THE LINNEAN SOCIETY.

Vou. XXXYV. ZOOLOGY. No. 231.

CONTENTS.
Page
I. The Percy Sladen Trust Expeditions to the Albrolhos Islands
(Indian Ocean), under the Leadership of Prof. W. J. Daxiy,
F.L.S., F.Z,.8.—Amphipoda and Isopoda. By Wm. Tarrzr-
SALL, D.Sc., Keeper of the Manchester Museum. (Communi-

eated by Prof. W. J. Dakin, F.L,.8.) (Plates 1-3.) ............ i
Il. —— On Two Sea-pens from West Australia. By Sypnery J.

Hickson, F.R.S., Professor of Zoology, University of Man-

chester. (Communicated by Prof. W. J. Daxiy, F.I.S8.) ...... 21

III. The Raninide : a Study in Carcinology. By Giupert C. Bourne,
M.A., D.Sc., F.R.S., F.L.S., late Fellow of Merton College,
Oxford, and Professor of Zoology and Comparative Anatomy.
Cielito state) mateo vaniti tes disiciodesetnannvacstevetsausenetaicnams aces 25

LONDON:

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LINNEAN SOCIETY OF LONDON.

LIST OF THE OFFICERS AND COUNCIL.
Elected 24th May, 1922.

PRESIDENT.
Dr. Arthur Smith Woodward, F.R.S.

VICE-PRESIDENTS.

EK, T. Browne, M.A. Horace W. Monckton, F.G.S
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COUNCIL.

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Prof. Dame Helen’ Gwynne-Vaughan, Rothschild, F.R.S.

D.B.E. Dr. E. J. Salisbury.
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LIBRARY COMMITTEE.
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James Britten, Esq. L. V. Lester-Garland, M.A.
Dr. R. R. Gates. Dr, E. J. Salisbury.
Dr, A. D. Imms. B. B. Woodward, Esq.

THE JOURNAL

OF

THE LINNEAN SOCIETY,

(ZOOLOGY.)

THE PERCY SLADEN TRUST EXPEDITIONS* TO THE ABROLHOS
ISLANDS (Inpiaw Ocray).

Under the Leadership of Prof. W. J. Dakty, F.L.S., F.Z.S.

Amphipoda and Isopoda. By W. M. Tarrmrsart, D.Sc., Keeper of the
Manchester Museum. (Communicated by Prof. W. J. Daxiy, F.L.S.)

(Prats 1-3.)
{Read 5th May, 1921.]

I am indebted to Professor Dakin for the opportunity of examining the small
collection of Amphipoda and Isopoda which he made during his expedition
to the Abrolhos Islands. Nine species of Amphipoda and three of Isopoda
were found in the collection. The number of specimens was very small, and
considerable difficulty has been encountered in the endeavour to identify
them satisfactorily. I have not been able to satisfy myself that any of the
Amphipoda represent undescribed species, but in almost all cases the
specimens showed some departure from existing descriptions. The recent
work of Chilton has, however, shown that in the Amphipoda we are dealing
with a deminant and yirile group of Crustacea exhibiting relatively endless
variations, and ihe time has now arrived when it is better to determine the
extent of specific variation than to regard each departure from the normal as

of specific value. Future work will undoubtedly reduce the number of valid

species of Amphipoda

* Acknowledgement must be made here also of a grant from the Government Grant
Committee of the Royal Society, and a grant from the British Association.

LINN. JOURN.—ZOOLOGY, VOL. XXXY. 1

2 SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

The main interest of the collections lies in the bearing which it has on the
general question of the origin of the fauna of the Abrolhos Islands. As
Professor Dakin has already pointed out, these islands are remarkable as
being the most southerly islands in the world with coral reefs fringing their
shores, and this fact would suggest that the fauna of the islands should, in
the main, bear a strong tropical facies. ‘The Amphipoda in this collection
hear out that suggestion. I have in the following table indicated the known
geographical distribution of all the species in the collection. It will be seen
at once that six of them have a wide distribution in the tropical and sub-
tropical waters of the Indian and Western Pacific Oceans, ranging from
South and Hast Africa to the Oceanic Islands of the Pacific. Two other
species, Parharpinia villosa and Grubia setosa, have so far only been met with
in the waters round South and Hast Australia and New Zealand. Only one
species, Parawaldeckia kiddert, seems to be a migrant from the Southern
waters of the Antarctic sub-Polar seas.

Of the Isopoda, two species are described as new, but all three species in
this collection belong to the Spheeromidee. a family of Isopoda characteristic
of the warmer waters of the globe.

|
|
|
|
|

g | ea ; yd |
n-* oo 2) | | ||
EN xa) Ee \imeree=ta
no) Yd tal oS S | cs é
3 <s foe | | oN &
el ‘aS ‘3 3} ; Bet
Ss : & = 2) iS = os & 3 oe | 3S E | ao}
2 CH (eer e+ GH 0s (ey fl | p=]
a i) 5 an Go} |] Bpos Sr ol 2 ||
I a : ‘S S
a om Oo 28 a ow = | xF, Oo
a | & (OR (SS les See] & ce | iS
o | ae. S | = cond
a < Be | 28 (ER Sagi N | awe Pa
<=) » = 9 | BO oe) ew e 1m | o
2 2 |i)! Sa | Bei joes! 5 | ee ee)
oA) S |4n,e <q ZA | ne | iS)
| | | |
| — a pee =
| |
|\Parawaldechia kiddert ....| Gu ie Ox x x xX Kerguelen.
= d pe! ene | LA oa aa
|Parharpinia villosa ...... | x Ieex 5
Leucothoé spinicarpa...... | Xx x Xx x x || |
ahs | —<————|___—_ SS —
Ceradocus rubromaculatus | x x x x x x 1
|
= —— IL
| | |
Mera tenella............ x | |) os =] *
|
a | | \—— | | ==
Mera mastersti.......... | 28 x Se Se x x oo
:
Elasmopus subcarinatus....| do |i} 2X x Xx :
Eurystheus atlanticus Xx x . x ‘Cape Verde.
= — gated!
\|Gaubta setosa .......-.-..| : [femal 3 2
| = = cee — a.
| ESV
: z - . 10 | |
Go) >R Ord ~ is) 5 . : 2
Joe |aS] ae SaRalgs Re) es
tes Sa | os = 1 (SS | Ss
4 Cores O psi r= Oo AQ) D lo
| Lim 1 mal Se eo i a | aoe =
b0_> or S Teale -~5
acl at los | S leadiaes| ae) a
) oer tS) oO 8 @ 10 i ta OD 5/06 5 5
Qa2s|/H5 | 4O 2 (2ES|Fe se] og B=)
ad ree =O a Qatlions|ms rs
Se tSe PS | eo Ieee ee | S|
; mm | FR | & a jARO|IOCH| OH | & |

DR. W. M. TATTERSALL : AMPHIPODA AND ISOPODA. 3

AMPHIPODA.
Family LYSIANASSID &.
Genus ParaAwALDEcgta, Stebbing.

1. PARAWALDECKIA KIDDERI (S. J. Smith). (Pl. 1. figs. 1-6.)
Nannonyx kiddert, Chilton, 1909, p. 615.
Parawaldeckia Thomsoni, Stebbing, 1910 (1), p. 571.

Locality. Dredge off Wallaby Group, one male and two females, 6-7 mm.

Distribution. Kerguelen (Smith), Torres Straits (Miers), Tasmania (Thom-
son), New Zealand (Thomson), Campbell Islands and Auckland Islands
(Chilton), Kermadee Islands (Chilton).

Remarks. The difficulties which present themselves to the student of
Amphipoda in attempting to identify isolated specimens from remote
localities are nowhere more abundantly illustrated than by a reference to the
history of this species, as detailed by Chilton in the paper quoted above. It
had been referred to three different genera and been described under at least
five and probably as many as eight different specific names. Its adventurous
career was even then not at an end, for in the next year Stebbing, who had
not been able to consult Chilton’s paper before his own went to press, pro-
posed a new genus, Parawaldechia, for the reception of Vannonyx thomsoni,
one of the many synonyms of this species. It seems that at last the species
has a permanent abiding place of its own.

I believe my specimens to helong to the species as described by Chilton.
They agree closely with his description except in one point. The adult male
presents the usual sexual differences in the third uropods exhibited by so
many Lysianassidze, in having the rami of these appendages enlarged and
fringed with plumose sete. Chilton says that in the male the third uropods
are the same as in the female. I give a figure of the telson of one
of my specimens, showing it to be slightly excavated but not distinctly
cleft. Hach lobe of the apex is armed with two short spines, and there are
two delicate plumose sete on each lateral margin. In this respect my
specimens are in substantial agreement with Chilton’s descriptions. The
outer plate of the maxilliped is broadly rounded and unarmed, the inner
plate truncate and armed with three blunt teeth.

The genus Parawaldeckia is characterised by the possession of accessory
lobes to the branchial vesicles and by having the last joint of the peduncle of
the second antenna of the male dilated and moderately long, the remaining
joints being small. It is very closely allied to Waldeckia, and differs only in
having the telson slightly excavated, whereas in JValdechia it is deeply cleft.
- My specimens agree with the definition of the genus Parawaldeckia in these

points. There can be no question as to its relationship to Waldeckia, for a
1*

4 SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

comparison of the figures of some of the appendages given herewith with
those published by Chevreux and Walker for Waldeckia will show the closest
agreement in general form, especially in the first and second gnathopods.

Family PHOXOCEPHALIDA.
Genus PaRrHARPINIA, Stebbing.

2. PARHARPINIA VILLOSA (Haswell). (Pl. 1. figs. 7-14.)

Phoxus villosus, Haswell, 1880 (1), p. 258, pl. ix. fig. 2.
Parharpinia villosa, Sic] gaa 1906, p. 147.

Locality. Sandy Island, Wallaby Group, one female with incubatory
lamellee, 6 mm.

Distribution. Coasts of Australia (Haswell) and New Zealand (Thomson).

Remarks. In comparing this specimen with Stebbing’s description, it is
found to differ in the following points :—

|
| Stebbing. Present specimen.
1. Antenna 1.
Flagellum. 13 joints. | 10 joints.
AGUSEEDY App. | | WOgeaast | 7 joints.
2, Antenna 2 |
Flagellum. [17 joints. 11 joints.
3. Outer “plate of Extend to more than half- | Barely extend bevond the
Maxillipeds. way along the long first joint of the palp
second joint of the palp,| and have only eight
and are fringed with a serrate spines.

dozen graduated spines |

serrate on both edges. |

4, Gnathopod 2. | Fifth joint about as long | Fifth joint much shorter
| as the sixth. than the sixth.

5. Pereopods 1 & 2. | Apical spine of the fifth | Apical spine of the fifth
joint nearly as long as joint half as long as the
the sixth joint. | sixth joint.

6. Pereopod 3, | Fifth joint scarcely longer Sixth joint longer than
than the sixth. | the fifth,

ie Perzeopod 5. Second joint not produced | Second joint produced to

| to end of fourth. | end of fourth. .
8. Uropod 2. Pedunele fringed with ten | Peduncle with only eight
| outstanding spines. | spines.

The most important of these differences are numbers 3 and 4. The length
of the outer plate of the maxillipeds is one of the characters used by
Stebbing to separate the genera Parharpinia and Pontharpinia. In the
former it is described as elongate and fringed with serrate spines on the
inner margins. In the latter it is simply described as short. In my
specimen, the maxillipeds do not appear to differ markedly from those of
the genus Pontharpinia, except that the spines on the inner margin of the

DR. W. M. TATTERSALL: AMPHIPODA AND ISOPODA. 9)

outer plate are serrate on both margins, a feature not, so far as I aware,
deseribed in Pontharpinia.

{ am quite unable to understand Stebbing’s description of the second
gnathopod unless, indeed, by some chance the descriptions of enathopods
1 and 2 have become inadvertently transposed in his diagnosis. As will be
seen from the figures of these appendages given here, the fifth joint is
decidedly larger in gnathopod 1 than in gnathopod 2, and with this sup-
position, Stebbing’s description of the gnathopods would apply to my
specimen.

Stebbing’s description is based on specimens from New Zealand recorded
by Thomson (1882) as Phowus batei, Haswell. Stebbing identifies Thomson’s
specimens with Phowus villosus, Haswell, and refers Haswell’s P. batei to
Pontharpinia rostrata (Dana). On comparing my specimen with Thomson’s
description and figures, I find the most complete agreement except that the
carpus of the second gnathopods is distinetly shorter than Thomson shows.
Thomson does not describe or figure the fourth perzeopods.

My specimen differs from Haswell’s original description in the larger size

of the eyes, in the fewer joints in the accessory appendage of antenna 1, and
in the shape of the second joint of the fifth pereeopods. In the characters of
the eyes and the fifth pereeopods it agrees with Haswell’s description of
Phoxus batei, and as the latter is said by Haswell to differ from P. villosus
in the larger size of the eyes and the form of the fifth perzopods, it looks as
if my specimen really belonged to Phowus batei, Haswell= Pontharpinia
rostrata (Dana). But the long and slender form of the fourth perzeopods
agrees much more with those figured by Haswell for P. villosus.

The genera Pontharpinia and Parharpinia are distinguished, according to
the definitions of these genera given by Stebbing in ‘ Das Tierreich,’ by the
following characters :—

(1) In Pontharpinia the third joint of the mandibular palp is shorter
than the second. In Parharpinia the third joint is longer than
the second.

(2) In Pontharpinia the outer plates of the maxillipeds are short. In
Parharpinia they are elongate and fringed with serrate spines on
the inner margins.

(3) In Pontharpiuia the fourth and fifth joints of the third and fourth
pereeopods are expanded. In Parharpinia they are not expanded.

(4) In Pontharpinia the expanded second joint of the fifth pereeopods is
greatly produced, while in Parharpinia it is only moderately
produced.

These differences appear to me of doubtful value. My specimen is a
Parharpima in the characters of the mandibular palp and the form of the
third and fourth perseopods, and a Pontharpinia in the characters of the
maxillipeds and of the fifth perseopod.

(<r)

SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

On the whole, my specimen agrees most closely with those described by
Thomson as Phovus batei. As these specimens formed the basis of Stebbing’s
description of P. villosa, Haswell, and were those upon which Stebbing
founded the genus Parharpinia, I here record the present specimen as
Parharpinia villosa (Haswell), with an expression of doubt as to the separate
identity of the genera Parharpinia and Pontharpinia.

My specimen differs from Pontharpinia uncinata, Chevreux, in the evenly
rounded epimeral plate of the third segment of the pleon, from P. stimpsonii, —
Stebbing, in the shorter and evenly rounded rostrum, and from P. pinguis
(Haswell) in the less robust pereeopods. I should have been quite content
to record my specimen as Pontharpinia rostrata (Dana) except for the longer
and more slender fourth perzeopods. In Dana’s figure the eyes are larger
and more rounded in form and the rostrum shorter than in my specimen,
but these differences disappear in the light of Haswell’s description of Phowus
bate?, which Stebbing regards as synonymous with P. rostrata.

A redeseription of Australian and New Zealand species is a necessity
before they can be placed on a sound basis, and it is to be hoped that this will
be attempted by some one on the spot who has access to material.

Family LEUCOTHOID A.
Genus LrucorHo®, Leach.

3. LeucorHo® spintcarpa (A bildgaard).
LL. spinicarpa, Chilton, 1912 (2), p. 478.
5 » Barnard, 1916, p. 148.

Localities. Wooded Island, Easter Group, two specimens. Sandy Island,
Wallaby Group, five specimens.

Dnstribution. Cosmopolitan.

Remarks. The two papers quoted above give the full synonymy of this
species and full notes on its known distribution.

Family GAMMARID AN.
Genus Cerapocus, A. Costa.
4, CERADOCUS RUBROMACULATUS (Stimpson). (PI. 1. figs. 15, 16.)

C. rubromaculatus, Stebbing, 1906, p. 430 (synonymy).
5 ss Walker, 1904, p. 272; 1905, p. 927 ; and 1909, p. 334.
Chevreux, 1907, p. 479.

” ”
9 0 Stebbing, 1908 (1), p. 8L; 1908 (2), p. 456; 1910 (1), p. 598.
90 % Chilton, 1916, p. 369.

Locality, Sandy Island, Wallaby Group, one female, 10 mm., and one male
and two females, 6 mm.

Re

DR. W. M. TATTERSALL: AMPHIPODA AND ISOPODA. 7

Distribution. Caylon, the Maldives and Laccadives, and the coast of Africa
(Walker) ; South Africa (Stebbing) ; Australia and Tasmania (Haswell &
Stebbing) ; New Zealand (Chilton) ; French Oceania (Chevreux).

Remarks. From the published descriptions and figures, this species would
appear to be subject to great variation in the antennze, antennules, first and
second gnathopods, fifth perseopods, and in the size and extent of the
serrations on the margins of the pleon and its epimeral plates.

These facts have led me to ascribe all the four specimens in this collection
to the same species, notwithstanding the differences and the extraordinary
dimorphism in the second gnathopods of the only male specimen.

In the large female specimen, 10 mm. in length, the dorsal surface of the
pleon is microscopically shagreened. The teeth on the posterior margins of
the first three segments are longer and coarser than in the smaller specimens,
and there is a seta alternating with every tooth. ‘The epimeral plate of the
first pleon segment has two teeth on the lower border and eight on the hinder
border, or ten teeth in all. The second pleon segment has five teeth on the
margins of the epimeral plate, and the third segment fourteen teeth. It is
not easy to decide where the lower and hinder margins meet. The antennal
peduncle is equal in length to the antennular. The accessory flagellum of
the antennule has seven joints and the main flagellum nvomlget ma. The
flagellum of the antenna has fourteen joints. The hinder lower corner of
the second joint of the fourth and fifth perseopods is prolonged into a spine,
as figured by Chevreux (1907) for specimens from French Oceania.

In the smaller specimens the teeth on the dorsal border of the tirst three
pleon segments are shorter and finer than in the larger specimen. ‘The
epimeral plates of the first two segments of the pleon have one or two teeth
on the lower border but none on the hinder. The third segment of the pleon
has six teeth on the hinder border and two on the lower border of the
epimeral plate. The flagellum of the first antenna has seven joints and the
accessory appendage four joints. The flagellum of the second antenna has
from 15-21 joints. The hinder lower corner of the second joint of the
fourth and fifth perseopods is much less produced than in the large
specimen, and is more like the typical form.

The male specimen presents a strking dimorphism in the second gnathio-
pods. The right one is of the normal type, the palm rather oblique and
furnished with two flat-topped teeth. It is not so robust as in the specimens
figured by Cheyreux from French Oceania, and the palm is more oblique and
not transverse. But Chilton describes specimens from New Zealand which
appear to agree with mine. ‘The second gnathopod on the left side is quite
different. The propodus is larger and more robust, the palm very oblique
with no prominent tooth marking its edge, but with a very strong obtuse

8 SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

tooth near the base of the nail. The whole limb recalls to some extent that
found among the males of the genus Hlasmopus, though it is not setose at
all. Stebbing, in the ‘Challenger’ Report and in recording this species
from South Africa, notes the dissimilarity between the right and left second
gnathopods in some of his specimens; but the dissimilarity usually takes the
form of one of the gnathopods being of the female type and the other of
the male. No such striking instance as the present one has been noticed
before.

With regard to the serrations on the epimeral plates of the first three
segments of the pleon, the typical form has one or two serrations on the
lower margin in all three segments, bat only the third segment is serrated on
the hind margin. Chevreux describes serrations on both lower and hinder
margins of the epimeral plates of the second and third pleon segments in
specimens from French Oceania. Haswell says of Australian specimens
“lateral plates (epimera) of the three anterior segments of the pleon
serrated posteriorly,” while in J/era spinosa, now regarded as a synonym of
C. rubromaculata, he gives only the third segment with serrations on the
posterior margin of the lateral plate. There is thus abundant evidence of
variation in this character. I may note that, in all my specimens, the first
joint of the mandibular palp has a spiniform process on its inner corner as
described by Walker (1904) and Chevreux (1907).

Genus Mara, Leach.

5. Mana TENELLA (Dana).
Gammarus tenellus, Dana, 1852, p. 952, pl. 65. fig. 7.
Mera tenella, Walker, 1904, p. 272, pl. 5. fig. 31.
M., tenella, Stebbing, 1206, p. 438.

Locality. Wooded Island, Haster Group, one male, 5 mm.

Remarks. The single specimen agrees closely with the description and
figures of a specimen from Ceylon referred to Dana’s species by Walker.
The only noteworthy point of difference is that the dactylus of the second
gnathopods in the male has a well-marked tooth about the centre of its inner
margin, which fits into a corresponding excavation on the palm. The serrate
hind margin of the third segment of the pleon and the slightly different
form of the second gnathopods cf the male seem to distinguish this species
from M. viridis, Haswell, to which it is otherwise very closed allied ; in fact,
this species seems to me to afford an additional argument in support of
Chilton’s contention that Haswell’s species belongs to the genus Mera, and
not to Elasmopus to which Stebbing would refer it.

Distribution. Fiji (Dana) ; Ceylon (Walker).

DR. W. M. TATTERSALL: AMPHIPODA AND ISOPODA. 9

6. Mara masrersit (fasvwell).

Megamera mastersit, Haswell, 1880 (1), p. 265, pl. 11. fig. 1.
M. thompsoni, Miers, 1884, p. 318, pl. 34. fig. B.
Mera mastersti, Stebbing, 1906, p. 459.

0 a Chevreux, 1907, p. 481.

“9 5 Stebbing, 1910 (1), p. 642.
5 9 Chilton, 1911, p. 564.

” 5 Barnard, 1916, p. 195.

6 i Chilton, 1916, p. 867.

Locality. Sandy Island, Wallaby Group, one male, 5 mm.

Distribution. Port Jackson, Australia (Haswell) ; several localities in the
Torres Strait (Miers); Gambier Archipelago (Chevrenx); Kermadee
Islands (Chilton) ; Mozambique (Barnard); New Zealand (Thomson &
Chilton).

Genus ELasmopus, A. Costa.

7. HLASMOPUS SUBCARINATUS (Haswell).

Megaumera subcarinata, Wuswell, 1880 (2), p. 385, pl. 21. fig. 4.
Elasmopus subcarinatus, Chilton, 1915, p. 321, figs. 1-6.

Locality. Dredged off Wallaby Group, one male, 6 mm.

Distribution. Shores of New Zealand, Australia, Ceylon, South Africa,
and Indian Ocean.

Remarks. Chilton, in the paper quoted above, has given a full account and
synonymy of this species and of its geographical distribution. The species
is distinguished from its congeners by the two carinze of the fourth segment
of the pleon each ending in a prominent tooth. In this character my
specimen agrees with the type, but it differs from typical specimens as
described by Chilton in the following characters :—

(1) In the first antennee, which are much shorter than usual and have
only fifteen joints in the flagellum. The accessory appendage is
quite short and only has two joints. Walker recorded specimens
from Ceylon in which the accessory appendage of the first antenna
was only two-jointed.

(2) In the hinder margin of the second joint of the last pair of
pereopods, which is closely and coarsely pectinate as described
for the species 7. pectenicrus, Sp. Bate. Chilton says of all the
pereeopods that the “ basal joint ...... posterior margin simply
serrate,” while Stebbing says that the margins are ‘“ strongly
serrate.” But the condition in the present specimen agrees with
Walker’s description in HH. serrula (=. pectenicrus) : “the
greater part of the hind margin elegantly cut into flat-topped

10 SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

teeth of a peculiar form” (Walker, 1904). Barnard (1916), in
describing HE. pectenicrus, Bate, from 8. Africa, to which species
he refers Walker’s /. serrula as a synonym, figures the second
joint of the fourth perseopod, showing a condition exactly
similar to the second joint of the fifth pereeopod in my specimen.
I may note that on one side the hinder margin of the second
joint of the fifth perzeopod is concave and on the other
convex or evenly curved. Barnard suggests that the concave
margin is a condition of maturity. The fifth perzopods of
my specimen are not so setose as is described by Chilton for
New Zealand specimens. Having only one specimen, it is
impossible to say whether these differences are constant and
of specific value. The two dorsal carinz of the fourth pleon
sevment exclude this specimen from all other described species,
and in the character of the second gnathopods the specimen is a
typical adult male of /. subcarinata as figured by Chilton. I
preter, therefore, to regard the specimen as at most an individual
variation of this widely distributed species.

Family PHOTIDZ.
Genus Etrysrnenus, Bate.

8. HurYsTHEUS ATLANTICUS (Stebbing). (Pl. 1. figs. 17-20.)

posers atlantica, Stebbing, 1888, p. 1101, pl. 114.

G. zeylanicus, Walker, 1904, p. 282, pl. 6. fig. 41.

G4 gardineri, Walker, 1905, p. 929, pl. 88. figs. 11-14, 16-17.

Lurystheus Cipla Stebbing, 1906, p. 611.

E. atlanticus, Stebbing, 1908, p. 86 (1), pl. 40 B.

LE. zeylanicus, Walker, 1909, p. 339

LE. atlanticus, Walker, 1909, p. 339.

we 5, stebbing, 1910 (1), p. 614.

Locality. Dredged off Wallaby Group, one female with ova, 6 mm. ; one
adult male, 6 mm. ; two young males, 4 and 4°5 mm.

Distribution. Cape Verde Islands (Stebbing), Ceylon, Maldive and
Laccadive Archipelagoes, Seychelles (Walker), S. Africa (Stebbing),
Australian coasts (Stebbing).

Remarks. These four specimens are ali imperfect; orly one, the larger of
the immature males, has any of the antenne still attached, and that only the
peduncle with the accessory appendage on one side only. The accessory
appendage is composed of four joints, three large ones and a small terminal
one.

Only the two larger specimens have the lageniform eyes characteristic of
the species. In the two smaller specimens the neck of the flask, so to speak,
is wanting.

DR. W. M. TATTERSALL : AMPHIPODA AND ISOPODA, iat

I have figured the distal joints of the second gnathopods of all four speci-
mens. The second gnathopods of the female agree very fairly well with
those figured for /. afer by Stebbing (1888), for /. zeylanicus by Walker
(1904), and for L. gardinert by Walker (1905). Those of the smaller
immature male resemble the figures given by Stebbing for the type-specimen
(a female) from Cape Verde (1888, pl. 114, gn. 2) of FE. atlanticus, by
Walker for the young male of LH. zeylanicus (1904, pl. 6. fig. 41, gn. 2,
3 jr), and by Walker for the young male of . gardineri (1905, pl. 88.
fig. 14).

The second enathopods of the larger of the immature males agree closely
with those figured for the adult male of . zeylanicus by Walker (1904, pl. 6.
fig. 41, gn. 2, d), and is not unlike the figures given by Stebbing for the
adult male of /. atlanticus (1908 (a), pl. 40 B). Stebbing’s specimen had,
however, lageniform eyes.

The second gnathopods of the adult male are not quite like those figured
for any of the species of Hurystheus, but seem to be the adult condition of
the immature males in this collection. I feel convinced that the four
specimens here dealt with belong to one species. They were all collected
at the same time in the same place, and appear to me to represent
the adult female and three stages of growth in the male. From this
it follows that the lageniform shape of the eyes is an adult character
only, or, alternatively, a character which varies greatly. I feel sure, too,
that my specimens belong to the same species as 2. zeylanicus, Walker, and
LN. gardineri, Walker. Neither of these species has lagenitorm eyes, but
judging from my specimens, most of Walker’s specimens were immature.
Walker himself in 1909 united these two species, and in a footnote to the
same paper expresses the opinion that both are synonymous with /. atlanticus,
Stebbing. In accepting that verdict, it is necessary to point out that the type
of HE. atlanticus is a female which has the lageniform eyes of the adult and
second gnathopods of the form ascribed here to an immature male with oval
eyes. Both Chilton 1912 (2) and Barnard (1916) have suggested the possi-
bility of . atlanticus being the same as L. afer (Stebbing), and support is
given to that idea by the close resemblance of the second gnathopods of the
adult female in the present collection to those figured by Stebbing for E. afer,
female. But the latter species has long oval eyes, not laveniform in shape.
Chilton’s specimens, identified with doubt as L. afer, seem to have gnathopods
altogether longer and more slender than in the present specimens.

It will be seen that there has been considerable difficulty in arriving at a
conclusion with regard to the identity of these specimens. I can best sum
up the evidence yielded by them, by saying that they are certainly the
sume as 1, zeylanicus and LE. gardineri of Walker, and that I accept his
opinion that these species are synonymous with LH. atlanticus, Stebbing, a
widely distributed and apparently very variable species.

12 SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

Family AMPHITHOIDA,
Genus Grupa, Czerniavshy.
9, Grupta seTosa (Haswell). (Pl. 2. figs. 21-24.)

Amphithoé setosa, Haswell, 1880 (1), p. 270.
Grubia setosa, Stebbing, 1906, p. 644.

Localities. Four males and four females, up to 15 mm. in length, from
Sandy Island, Wooded Island, and Pigeon Island. (Shore collecting in each
case.)

Distribution. Sidney Harbour and Botany Bay, New South Wales
(Haswell & Chilton).

Remarks. Haswell’s original description is short and unaccompanied by
figures, but the present specimens are in agreement with it. Chilton,
who examined specimens from Sydney Harbour which he referred to
this species, stated that the first antenna possessed a small accessory
appendage. The present specimens, or at least such of them as still
retain the first antenna, possess a small two-jointed accessory flagellum on
that appendage, barely longer than the first joint of the main flagellum.
The species is therefore a true Grubia, in which genus Stebbing has already
placed it.

The species is mainly distinguished by the form of the first and second
gnathopods in both sexes, and especially by those of the male. I figure
these appendages herewith. They are similar in the male and female,
but in the former they are more massive and are densely fringed with long,
delicate, plumose sete, especially on the second, carpal, and propodal joints.
The palmar margin is undefined in the first gnathopod, but in the second it
is bounded by a rather prominent tooth, on the inside margin of which is a
strong spine. In the second gnathopod of the male the limiting tooth is
stronger, and immediately inside it on the palm is a small excavation into
which the nail fits. ‘The nail in both gnathopods is provided with a closely-
set row of blunt spines on the inside margin. The figures will serve to show
the general form of these limbs and the proportions of the joints.

For the rest, the species has no very marked characters. The first antenne
are almost as long as the whole animal, the third joint of the peduncle about
1-1 as long as the second, and the flagellum composed of about 50 joints.
The second antenne are 2 as long as the first. The fourth joint of the
peduncle reaches to the distal end of the second joint of the peduncle of the
first antenne. The fifth joint equals the fourth in length, and the flagellum
has 35 joints. The lower margin of the peduncle is beset with long
plumose hairs, particularly in the male. The inner lobe of the first maxilla
bears six or seven fine plumose sete. ‘The third uropods have the peduncle
much larger than the rami, with a row of seven short, blunt spines on the
dorsal apical margin. The inner ramus bears one spine and a number of sete

DR. W. M. TATLERSALL: AMPHIPODA AND ISOPODA. Res

at its apex. The outer ramus bears two recurved hooks of equal size and a
short, blunt spine near the proximal end of the dorsal surface. In the. light
of Chevreux’s description of G. brevidactyla and of the present species,
Stebbing’s diagnosis of the genus Grubia in ‘ Das Tierreich’ will require
emendation in the following points:—antenna 1 has a short accessory
flagellum of one or two joints, and maxilla 1 the inner plate with a few
small lateral setee.

The present species is distinguished by the form of the first and second
legs and, possibly also, by the structure of the third uropods.

ISOPODA.

Tribe FLABELLIFERA.
Family SPHAROMID A.
Group HEMIBRANCHIATA.
Genus Cinrcmopsis, Hansen.

CILICAHOPSIS DAKINI, sp.n. (PI. 2. figs. 25-29; Pl. 3. fig. 34.)

Locality. Woody Island, Haster Group, one adult male, 13 mm. long,
9 mm. broad.

Description. The single specimen of this new species, a male, measures
13 mm. in length and 9 mm. in breadth, and is thus one and a half times as
long as broad. The body is strongly convex in dorsal view and without
granulations or tubercles. The posterior margins of all the thoracic somites,
the lateral margins of the body, the uropods, and the abdomen are fringed
with short, thick hairs, the general appearance of which can be seen from
Pl. 3. fig. 34. Similar hairs are scattered over the dorsal surface of the
abdomen, especially on the two large bosses. The distinguishing features of
the species are to be found in the abdomen and uropods. The abdomen
shows three proximal segments separated off from the large terminal telsonic
somite: the sutures separating the first two abdominal somites not complete
in the mid-dorsal line. The posterior margin of the third abdominal somite
is considerably and broadly produced in the mid-dorsal line, but does not
form a process of any kind. he terminal abdominal somite bears two
relatively enormous mammiform bosses. The posterior margin is evenly
rounded, and shows the merest trace of a median process separated by shallow
notches from the rest of the margin. The uropods consist of a massive basal
joint and a long, narrow, terminal joint. The basal joint, I take it, represents
the fused basipodite and endopodite, and the long terminal joint the exopod.

The figures which I give of the epistome, the antennule, antenna, second
and eighth thoracic limbs will suffice to show the general features of these

14 SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

appendages. The epistome is quite truncate anteriorly; the front margin
fringed with short, thick hairs similar to those on the rest of the body. The
second to the eighth thoracie limbs are bi-unguiculate and characterised by
the development of strong spines on the inner margins of the merus, carpus,
and propodus.

The pleopods are of the typical hemibranchiate type. The exopod of the
third, fourth, and fifth pairs is two-jointed. In the third pair the margins
of the endopod and the distal joint of the exopod are fringed with plumose
setee. In the fourth and fifth pairs there are no plumose setee, but the outer
margins of both branches of the fifth pleopods possess a fringe of simple
short hairs. ‘The appendix masculina on the second pleopods is longer than
the branches, with the terminal portion narrower than the rest and slightly
hooked. j

This species differs from the definition of the genus Cilicwopsis given by
Hansen (1905) in possessing a vestige of a mesial lobe on the posterior
margin of the last abdominal somite, and from the type-species C. granulata
(Whitelegge) and its ally, C. whitelegge: (Stebbing), in being without a long
process from the median portion of the anterior part of the abdomen. In this
latter character it agrees, however, with Ciliewa stylifera, Whitelegge, and
C. ornata, Whitelegee, which Hansen refers tentatively to the genus Cili-
cwopsis. It may be noted that both the last-named species show a further
resemblance to C. dakini in having the posterior margin of the third abdo-
minal somite preduced in the mid-dorsal line.

I know of no species of the group Cymodocini with which the present
species can be confused in the form and structure of the abdomen, and I have
pleasure in associating it with the name of its discoverer.

Genus Cymopocn, Leach.

CymopocrE MAMMIFERA, J/aswell (?). (Pl. 3. fig. 35.)
C. mammifera, Haswell, 1880 (4), p. 474, pl. 18. fig. 1, le.

Locality. Off Wallaby Islands, dredging, two specimens, 12 mm.

Remarks. It is with some doubt that I refer these two specimens to
Haswell’s species, but they agree completely with his short description and
figures. Of the two specimens, one is apparently an immature male in so
far as it has a pair of penial filaments on the sternum of the last thoracie
somite, but no appendix masculina on the second pleopods. he other is a
female. ‘The species is a typical hemibranchiate sphzeromid in the structure
of the pleopods, and would seem to fall into the genus Cymodoce by the well-
developed mesial lobe in the abdominal notch, by the absence of a mesial
process on the abdomen of the male, and by the well-developed endopod of

DR. W. M. TATTERSALL : AMPHIPODA AND ISOPODA. 15

the uropods. The body in both sexes is smooth, except for a sparse coating
of short fine hairs on the sides of the thorax and on the abdomen and the
uropods. There are no granulations or small tubercles. Jn the male, the
abdomen of which I have figured, the posterior margin of the fourth
abdominal somite, which is completely separated from the terminal part of
the abdomen, is produced into two small tubercles, one on each side of the
middle line. The terminal part of the abdomen has two large mammiform
bosses on the dorsal surface separated by a shallow depression, each with a
small tubercle on the highest part of the boss. The posterior margin of the
abdomen is tridentate, a large mesial lobe separated by notches from a small
lateral lobe on each side. ‘The large median lobe is very much bigger than
the lateral lobes and almost masks them in dorsal view. The uropods are
shorter than the median lobe of the abdominal notch, the endopod well-
developed and as long as the exopod, with the apex truncate ; the apex of the
exopod is rather acute. The female differs from the male in being without
the tubercles on the posterior margin of the fourth abdominal somite, and in
having the bosses on the terminal portion of the abdomen less well-marked
than in the male and without the tubercle on their highest parts.

Haswell’s types were collected at Port Denison in Queensland, and the
species does not appear to have been met with since. The species was
omittel from Haswell’s ‘Catalogue of Australian Crustacea,’ and has been
lost sight of since the original description appeared in 1880.

CyMODOCE PELSARTI, sp. n. (PI. 2. figs. 30-33 ; Pl. 3. fig. 36.)

Localities. Sandy Island, two males, 9-12 mm. Pigeon Island, one male,
5mm. ; one female, 6 mm. ; and six juv.

Description of the Male. The sides of the thoracie somites and the whole
of the abdomen and uropods are fringed with rather long hairs. Similar but
much shorter hairs are sparsely scattered over the dorsal surface of the
thorax. The thoracic somites have a double row of small round tubercles
extending as a band right across the posterior portion. These tubercles are
rather obscure on the anterior somites, but become successively more clearly
marked on the posterior ones. They are more clearly defined in the larger
specimens. The abdomen has an ornamentation of granules and tubercles,
as shown in PI. 3. fig. 36, which is taken from a male specimen, 9 mm. in
length. The whole of the abdomen and dorsal surface of the uropods is
finely granulose, and scattered among the small granulations are larger
rounded tubercles and a few still larger and pointed tubercles. The posterior
margin of the fourth abdominal somite, which is complete right across the
dorsal surface, is produced into two prominent pointed tubercles, one on each
side of the middle line. Hach of these tubercles bears a clump of long hairs
immediately in front. The remainder of the anterior portion of the abdomen

16 SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

is covered with larger and smaller tubercles. The posterior part of the abdo-
men has, in the main, three large bosses—an anterior pair, one on each side
of the middle line, and behind them a single median posterior boss. The
anterior pair of bosses are somewhat elliptical in shape, and each bears a
large, prominent, almost erect, conical spine with additional spines at its
apex. Behind this prominent spine are four or five sharply-pointed tubercles,
more prominent than the rest of the tubercles and granulations covering the
surface. The posterior median boss is circular in outline, quite smooth and
polished, and bearing a single almost erect stout conical spine. The posterior
margin of the abdomen is tridentate; the median lobe rather longer than the
lateral lobes, with the apex truncate and bearing two spine-like tubercles.
The lateral lobes have their apex pointed and bearing a single spine-like
tubercle. The uropods extend some way behind the posterior margin of the
abdomen ; their dosal surface is covered with fine granulations, with here and
there a larger and more prominent tubercle. he inner branch is longer and
stouter than the outer; its apex obtusely rounded, with a spine-like tubercle
at its tip. The outer branch is more pointed than the inner, and has a similar
spiniform tubercle at the apex. The mouth-parts and thoracie appendages
present no special features, and the figures which I give of the peduncles of
the antennules and attennze, second and eighth thoracic limbs will serve to
show the general structure of these appendages. The second to the eighth
thoracic limbs are stout and bi-unguiculate, with an armature of strong
spines on the inner margin of the merus, carpus, and propodus.

The pleopods are typically hemibranchiate, with the exopods of the third,
fourth, and fifth pairs two-jointed. The appendix masculina on the second
pair of the male is nearly twice as long as the branches, narrowing to a
finely-pointed apex, the distal portion curved slightly inwards.

The female differs from the male in having the body smooth, without
tubercles and granules, and with the hairs on the thorax and abdomen very
short and very much less numerous. The anterior part of the abdomen,
2.e. the combined first four segments, has the general surtace raised into.
two obscure bosses. The terminal portion of the abdomen has likewise two
bosses, more clearly defined and separated medianly by a shallow groove.
The posterior margin is tridentate ; the median lobe somewhat larger than
the lateral lobes, with its apex bluntly rounded.

Among all the deseribed species of the genus Cymodoce, this species comes
nearest to C. pilosa, M. Wd., and its allies C. longistylis, Miers, C. bicarinata,,
Stebbing, and C. sanzibarensis, Stebbing, but differs from all of these species
by the large, conical, erect spine which crowns each of the three main bosses
of the abdomen. I have associated the name of the discoverer of the
Abrolhos group of isiands with this pretty little species found there by
Professor Dakin.

DR. W. M. TATTERSALL : AMPHIPODA AND ISOPODA. 17

List or Lrrerature.

Barnarp, K, H., 1916.—Contributions to the Crustacean Fauna of South Africa. _Azn.
S. African Mus, yol. xv. pp. 105-803, pls. 26-28.

CuEvreEvx, E., 1907.—Amphipodes recueillis dans les possessions Francaises de l’Océanie
par M. le Dr. Seurat . . . . 1902-1904. Mém. Soc. Zool. France,
t. xx, pp. 470-527, 55 text-figs.

Cur iton, C., 1909.—The Crustacea of the Subantarctic Islands of New Zealand. The
Subantarctic Islands of New Zealand, vol. ii. pp. 601-671, 19 text-
figs.

1911.—The Crustacea of the Kermadec Islands. Trans. New Zeal. Instit.
vol. xliii. pp. 544-573, 4 text-figs.

1912 (1).—Miscellaneous notes on some New Zealand Crustacea. Trans.
New Zeal. Instit. vol. xliv. pp. 128-135.

1912 (2). The Amphipoda of the Scottish National Antarctic Expedition.
Trans. Royal Soc, Edinburgh, vol, xlyiii. pt. 2, no. 23, pp. 455-520,
2 pls.

1915.—The New Zealand species of the Amphipodan genus Elasmopus.
Trans. New Zeal. Instit. vol. xlyii. pp. 820-830, 12 text-figs.

1916.—Some Australian and New Zealand Gammaride. Trans. New Zeal.
Insti. vol. xlyiii. pp. 859-370, 6 text-tigs.

Dana, J. D., 1852.—United States Exploring Nxpedition. Vol. xiii. Crustacea.

Hansen, H. J., 1905.—On the propagation, structure, and classification of the family

Spheromide. Q. J. M.S. vol. xlix. N.S, pp. 69-135, pl. 7.

Haswett, W. A., 1880 (1).—On Australian Amphipoda. Proc. Linn. Soc. N.S. Wales,
vol. iv. pp. 245-279, pls. 7-12.

1880 (2).—On some additional new genera and species of Amphipodous

Crustaceans. Proc. Linn. Soc. N.S. Wales, vol. iv. pp. 319-850,
pls. 18-24.
1880 (3).—On some new Amphipods from Australia and Tasmania.
Proc. Linn. Soc. N.S. Wales, vol. v. pp. 97-105, pls. 5-7.
1880 (4).—On some new Australian marine Isopoda. Part I. Proc.
Linn. Soc. N.S. Wales, vol. v. pp. 470-481, pls. 16-19.
Miers, E. J., 1884.—Crustacea in Report on the Zoological Collections made in the Indo-
Pacific Ocean during the voyage of H.M.S. ‘Alert,’ 1881-2,
pp- 178-322, pls. 18-84.
Srepsine, T. R. R., 1888.—‘ Challenger’ Amphipoda. ‘ Challenger’ Reports, vol. xxix.
1904.—Marine Crustaceans. XII. Isopoda, with descriptions of a new
genus. Fuuna Geog. Mald. Lace. Arch. vol. ii. pt. 3, pp. 699-721,
pls. 49-53.
1905.—Report on the Isopoda collected by Professor Herdman, at
Ceylon, in 1902. Ceylon Pearl Oyster Fisheries, 1905—Supple-
mentary Reports, No. xxiii. pp. 64, 10 pls.

1906.—Amphipoda. “I, Gammaridea. Das Tierreich, Lief. 21.

1908 (1).—South African Crustacea. Pt. 1V. Ann. S. African Mus.

vol. vi. pp. 1-96, 14 pls.

1908 (2).—General Catalogue of South African Crustacea, dann. S.

African Mus. vol. vi. pp. 281-598, 8 pls.
1910 (1).—Crustacea. Part V. Amphipoda, (‘ Thetis’ Scientific
Results.) Mem. Austr. Mus. vol. iv. pp. 567-658, pls. 47 *-60 *.
LINN, JOURN.— ZOOLOGY, VOL. XXXV. 2

”

”

”

”

18 SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

SreppinG, T. R. R., 1910 (2).—Reports on the Marine Biology of the Sudanese Red Sea.
XIV. On the Crustacea Isopoda and Tanaidacea. Jowrn. Linn.
Soe. London, Zool., vol. xxxi. pp. 215-230, pls. 21-23.

1910 (3).—Isopoda from the Indian Ocean and British East Africa.
Trans. Linn. Soc. London, Ser. 2, Zool., vol. xiv. pp. 83-122,
pls. 5-11.

Tuomson, G. M., 1882.—Additions to the Crustacean Fauna of New Zealand. Trans. New

Zeal. Instit. vol. xiv. pp. 230-238, pls. 17, 18.

Watker, A. O., 1904.—Report on the Amphipoda collected by Professor Herdman, at
Ceylon, in 1902. Ceylon Pearl Oyster Fisheries, 1904—Supple-
mentary Reports, No. xvii. pp. 229-300, 8 pls.

1905.—Marine Crustaceans. XVI. Amphipoda. Fauna Geog. Mald.
Lace. Archip. vol. ii. Suppl. i. pp. 9238-932, pl. 88 & text-figs.
140-142,

1909.—Amphipoda Gammaridea from the Indian Ocean, British East
Africa, and the Red Sea. Trans. Linn. Soc. London, Ser. 2, Zool.,
vol, xii. pp. 825-544, pls. 42 & 43.

”

”

7

EXPLANATION OF THE PLATES.
PuatTe 1,
Parawaldeckia kidderi (S. I. Smith).

. Telson, x 680.

. Pedunele of antenna of male, x 33.
. Second thoracic leg, x 33.

Third 5 » X33.

. Fifth oF » =X 33.

. Highth 99 » X33.

Down se con pe

Parharpinia villosa (Haswell).

Fig. 7. Mandible, x 57.

8. Maxilliped, x 57.

9. Second thoracic leg, x 33.
10. Third m9 » X33.
11. Fifth * » X33.
12. Sixth oF Fos A SEREY
13. Seventh _,, op os ILL
14. Eighth __,, » X33.

Ceradocus rubromaculatus (Stimpson).

Fig. 15. Left third thoracic leg of male, 6mm., x 33.
16. Right ,, " » same specimen, X 39.

Eurystheus atlanticus (Stebbing).

Fig. 17. Hand of third thoracic limb of female with lageniform eye, x 460.
18. Hand of third thoracic limb of young male with oval eye, x 460.
19. Hand of third thoracic limb of young male with oval eye, x 460.
20, Hand of third thoracic limb of adult male with lageniform eye, x 460.

Tattersall.

ABROLHOS AMP

Journ. Liww. Soc. Zoon. Vol XXXV PI, 1.

JT Rennie Reid Lith Edint
AND ISOPODA.

Salar

atlas

Tattersall.

Aa etuin

Tattersall.

ee

SS
Sarai

SS
r /

pi)

OST. del.
ABROLHOS AMI

Journ. Linn. Soc. Zoon. Vol XXXV PL. 2.

J. TRennie Reid Lith Edin?

AND ISOPODA.

Journ Linn Soc ZOOL. Vol XXXV Pl. 2.
Tattersall.

Tattersall. Journ. Linn. Soc. Zoon. Vol XXXV Pl. 3.

OST. del. JTRenmeReid, Lith Edint

ABROLHOS AMPHIPODA AND ISOPODA.

DR. W. M. TATTERSALL : AMPHIPODA AND ISOPODA. 19

PLATE 2.

Grubia setosa (Haswell).

Fig. 21. Second thoracic limb of female, x 22.

22, Third i. Py 9D 5 BBY,

23. Second 5 0 male, x 22.

24, Third 0 1 mp 6 oe

Cilice@opsis dakini, sp. n.

Fig. 25, Epistome.

26, Antennule of male, x 22.

Nilo ANWR, pp SK BR

28. Second thoracic limb, x 22.

29. Highth 55 PD eee

Cymodoce pelsarti, sp. n.

Fic. 30, Antennule of male, x 33.
3l. Antenna ,, 4, X00,
32. Third thoracic limb, x 33.
82. Highth ,, os:

PLATE 3.

Fig. 34. Cilicceopsis dakini, sp. n., abdomen of male, x 10.
35. Cymodoce mammifera, Haswell, abdomen of young male, x 11.
36. Cymodoce pelsarti, sp. n., abdomen of male, x 16.

[I am greatly indebted to my wife for the drawings illustrating

this report. |

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SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS. 21

THE PERCY SLADEN TRUST EXPEDITIONS * TO THE ABROLHOS
ISLANDS (Iypran Ocray).

Under the Leadership of Prof. W. J. Daxry, F.L.S, F.Z.S.

On Two Sea-pens from West Australia. By Sypney J. Hickson, F.R.S.,
Professor of Zoology, University of Manchester. (Communicated by
Prof. W. J. Dakin, F.L.8.)

(Read 5th May, 1921.]

Ar the present time only three species of Pennatulacea have been recorded
from the waters of the Western coast of Australia, and these were all found
in Shark’s Bay, over 200 miles north of the locality from which the two
species here recorded were obtained. The species previously recorded are
Policella australis by Gray (1862), Virgularia elegans by Gray (1870), and
Pteroides hymenocaulon by Broch (1910).

The two specimens obtained by Professor Dakin from the neighbourhood
of the Abrolhos Archipelago belong to, or are closely related to, species that
are found in the Malay Archipelago; but as other species of the same genera,
namely Veretillum cynomorium and Pteroeides griseum, occur in the Mediter-
ranean Sea, and the genera are widely distributed in shallow to deep water,
the presence of the two sea-pens off Abrolhos is a fact of no value, at present,
for the determination of the affinities of the local fauna. It is quite possible,
or even probable, that they have travelled down from the North, but it would
not be surprising to find them a great deal further South, as they are not
tropical forms.

VERETILLUM MALAYENSE (Hickson) +. 20 fathoms. Off Long Island.

Length of rachis 80 mm., length of stalk 60 mm., diameter of rachis and
of stalk 12 mm., diameter of terminal bulb 14mm. Autozooids 14 mm. in
length by 2°5 in diameter.

The rachis is club-shaped, but differs from the rachis of the type-specimen
in coming gradually to a blunt extremity. The stalk has a well-marked

* Acknowledgement must be made here also of a grant from the Government Grant
Committee of the Royal Society, and a grant from the British Association.
+ ‘Siboga’ Pennatulacea, 1916, p. 48.

22 SLADEN TRUST EXPEDITIONS TO ABROLHOS ISLANDS.

basal bulbous swelling. The autozooids are large, and most of them are fully
expanded. The siphonozooids are numerous and irregularly scattered on
the rachis; they extend a considerable distance down the stalk beyond the
last autozooid. The stomodwa have a dorsal ventral diameter of 0-14 mm.
The axis is well developed, quadrangular in section, the diagonal measurement
being 15 mm. in its thickest part. I have not dissected out the axis from
this beautifully preserved specimen, but it can be seen through the transparent
tissues to be at least two-thirds of the total length of the colony.

The spicules of the rachis are long, thin plates with jagged edges, large
ones measuring 0°25x 0:07 mm., but they are very variable in shape and
size, and scattered or clustered. In the expanded part of the autozooids,
spicules are very scarce. I have seen one or two at the base of the tentacles,
but none in the tentacles themselves.

The type-specimens of this species were found in the Bay of Bima on
Sumbawa Island, Malay Archipelago, at a depth of 55 metres, and I have
been able to compare the characters of the specimen from W. Australia with
the type-specimens which are at present under my care. The specimens
from Shark’s Bay that were named by Gray Policella australis were trans-
ferred to the genus Veretillwm by Kiikenthal and Broch, and, in my opinion,
this transference was fully justified.

The principal difference between the species Veretillum malayense and
V. australe is that the autozooids of the former are about twice the size of the
autozooids of the latter. This character —the size of the autozooids—may be
more variable than we are justified in assuming on our knowledge of the
half-dozen specimens that have been described, and the two species may be
merged into one in the near future. But it is clear that the specimen
described above agrees more closely with the specimens hitherto called
V. malayense than it does with those deseribed as Policella (or Veretillum
australis).

PrEROIDES sp.?, juv. 20 fathoms. Outside Wallaby Group.

Length of rachis 25 mm., length of stalk 31 mm. Leaves 11-10.
Number of rays on largest leaves 5. Siphonozooid plate basal.

This specimen is evidently a juvenile form, as is shown by its small size, by
the small number of leaves, by the rudimentary character of the lower
leaves, and by the relative length of the stalk. It may also be regarded
as a sign of juvenility that there is only a single row of autozooids on the
greater part of the margin of the largest leaves, an additional autozooid of
what may be a second row appearing only in one or two isolated places.

As it is impossible, in the present state of our knowledge, to determine
accurately the relations of the juvenile forms of this genus, the principal
point of interest is to consider whether it is a juvenile of the only species of

PROF. S. J. HICKSON : TWO SEA-PENS FROM WEST AUSTRALIA. 23

the genus that has hitherto been identified from the West coast of Australia,
namely Pt. hymenocaulon. Broch (1910) examined and carefully described
two specimens of this species from Shark’s Bay. One was 94 mm. in length
and one 140 mm. in length, and therefore it is probable that they possessed
most of the adult characters. My reasons for believing that the young
specimen from the Wallaby Group is not P. hymenocaulon are: (1) that the
siphonozooid plate is marginal in P. hymenocaulon and basal in our specimen,
and (2) that the spicules in the stalk of P. hymenocaulon are smaller and
more numerous than in our specimen. I do not wish to maintain that these
reasons are conclusive, but I am inclined to think that the species to which
the specimen belongs is one of my Group II. 3 (see Monograph on ‘ Siboga’”
Pennatulacea, p. 231), represented by such species as Pt. grisewm or Pt.
malayense.

DR. G. C. BOURNE ON THE RANINIDA. 2

Or

The Raninidee: a Study in Carcinology. By Ginpert C. Bourne, M.A.,
D.Se., F.R.S., F.L.S8., late Fellow of Merton College, Oxford, and Linacre

Professor of Zoology and Comparative Anatomy.
gy | y
(PLaTEKs 4-7.)

[Read 15th December, 1921.]
MM. A. Minne Evwarps and BE. L. Bouvier conelude the Introduction to

their Report on the Deomiacea and Oxystomata dredged by the U.S. Coast
Survey Steamer ‘Blake’ with the following sentence :

“Nous avons
distrait du groupe des Oxystomes la famille des Raninidés qui nous parait
aberrante & plus d’un titre. Ces Oxystomes anormaux sont représentés par
de curieuses formes nouvelles dans les matériaux recueillis par le Blake ;
nous les étudions avec la plus grand détail, et ils font prochainement l’objet
dune étude trés complete.” So far as I have been able to ascertain by
search in the records of zoological literature, somewhat incomplete as they
are through disturbances arising out of the late war, this promised memoir
on the Raninide has not yet been published, and the following pages give
an account of my own studies on this aberrant and, as I shall strive to show,
primitive family of crabs. My attention was directed to them some eight
years ago, and I spent some time in the attempt to procure a well-preserved
specimen of Ranina, of which I had several ill-preserved examples collected
by Messrs. C. & E. Hose in Celebes. Hventually, after much fruitless corre-
spondence with other naturalists, I obtained a fine specimen of Ranina
dentata from Professor Kishinoye, of Tokio. It was admirably preserved in
spirit, and I have to express my best thanks to him for his trouble in pro-
curing me so excellent an example for study and dissection.
In addition to Ranina, I had examples of the following members of the
family in the collections of the Oxford University Museum :—
Notopus dorsipes, Fabr.; two examples collected by H.M.S. ‘ Nassau’ in
the Sulu Sea, 1871-72.
Raninoides personatus, White MS., Henderson; one example from the
Bay of Bengal, presented to the University by the Caleutta Museum.
Lyreidus tridentatus, de Haan ; two examples from Port Jackson, presented
to the University by the Trustees of the British Museum of Natural
History.
‘Two examples of a species labelled “‘ Notopus,” collected in the Sulu Sea
by H.M.S. ‘Nassau.’ These I am describing as a new genus and
species under the name of Notosceles chimmonis.

bo
lor)

DR. G. C. BOURNE ON THE RANINIDA:

I have also been able to examine the following species in the British
Museum of Natural History, and take this opportunity of thanking Dr. W.T.
Calman for his courtesy and assistance in bringing the material together for
examination :—

Notopoides latus, Henderson.
Notopus ovalis, Henderson.

Lyreidus Channeri, Wood-Mason.
Zanclifer caribensis, de reminville.
Cosmonotus Grrayt, Adams & White.

The above are preserved in spirit. I also had the opportunity of examining
a dried specimen labelled Ranilia muricata, M. Edw., and am inclined to the
opinion that, while it is a distinct species, it is closely allied to and should be
placed in the genus Votopus of de Haan.

Thus I have had opportunities of studying more or less closely nearly all
the known genera and most of the known species of the Raninidee, which
must be accounted one of the most ancient crab families, for it occurs in the
Cretaceous along with other forerunners of existing crabs. The proper
place of the Raninidi in classification las always been a subject of dispute.
Latreille first placed them among the Brachyura, but subsequently removed
them to the Macrura. Milne Hdwards in his ‘ Histoire Naturelle des Crus-
tacés’ (25, 1837) includes them along with the Dromididee, Homolide, and
Pactolidee as a tribe of his section Anomura Apterura. De Haan (34, 1850)
showed that the resemblances between the Raninide and Hippide were
superficial, and placed the- former among the Oxystomata, laying special
stress on the athinities of Lyrecdus to the Leucosiidee, ‘“ Uti in ceteris
Oxystomatibus, os Raninoideorum parte anteriore est canaliculatum. Rani-
noidea et Leucosidea unice sunt forme inter Decapoda, quorum regiones
pterygostomianze cum sterno coherent, ita ut maxillarum quintarum articuli
primi infra palpos sint dispositi et radices ipsis regionibus pterygostomianis
tegantur ; unicee, quorum cayitates branchiales a parte anteriore simplicem
tantum aperturam offerunt, uti jam Cl. Ndwards observavit. Lyreidus tam
distinctum vineulum constituit inter Raninas et Leucosias, ut vix dubium
remanere possit, quin justus hie sit illius familie locus.”

Boas (4, 1885) retained the Raninidee among the Brachyura, but laid
stress on their abnormal characters, and concluded by saying, “il m’est
impossible de comprendre ..... pourquoi les Ranines sont rangés avec les
autres Oxystomes.” In the ‘Challenger’ Reports, Henderson (85, 1888),
following Milne Edwards’ arrangement, describes the Raninidee among the
Anomura, and the same author, in a subsequent memoir (36, 1893), makes
them into a group, Ranidea, equivalent to his groups Dromidea and Hippidea
of the Anomura. Ortmann (41, 1892) follows de Haan in placing the
Raninidee in the Oxystomata, and subsequently (42, 1901) erected the

A STUDY IN CARCINOLOGY.

ne
1)

Oxystomata into a division (Abtheilung) equal in value to the Dromiidea
(Boas’s Dromiacea) and the Brachyura, the last-named division including all
other true crabs. This arrangement, with some minor modifications, has been
adopted by Alcock (1, 1896), Borrodaile (12, 1904), and Calman (23, 1909).
Against these authorities must be set the opinion of A. Milne Edwards and
Bouvier, quoted above, and the similar judgment of Boas that the Raninidze
cannot be included in the Oxystomata. But if carcinologists differ as to the
exact position that the Raninidee should occupy among the crabs, it is now
generally agreed (except Ortmann [42], who keeps the Oxystomata apart
from the Brachyura) that they are true crabs belonging to the section
Brachyura and not to the section Anomura of the Decapoda reptantia.
Indeed, this was clearly established by de Haan seventy years ago, and it
will be convenient to enumerate here the Brachyuran characters of the
Raninide, most of which were recognised by the clear-sighted author of the
volume ‘Crustacea’ of the ‘ Hauna Japonica.’ They are :—(1) The reduced
abdomen and the absence of uropods. (2) The absence of pleopods on the
first abdominal somite of the female. (3) The absence of posterior abdominal
appendages in the male. (4) The union of the pterygostomial region of the
carapace with the epistome. (5) The adherence of the branchiostegite to
the thoracic epimera. (6) The reduction of the branchiz. (7) The absence
of an exopoditic scale on the second antenna. (8) The presence of a sella
turcica posterior. (9) The expansion of the ischium and merus of the third
maxillipeds to form a plate-like covering to the other mouth-parts. (10) The
presence of orbits in which the eye-stalks can be concealed, and correlated
with this the reduction in size of the first and second antenne. Other
characters might be mentioned, such as the specialisation of the thoracic
limbs as organs of locomotion, involving a great development of the apo-
demes forming the walls of the cavities in which the muscles of the limbs are
contained. In other words, it is part of the distinctive facies of a crab that
the musculature is highly developed in connection with the locomotory
thoracic limbs, and is feeble almost to the point of disappearance in the
abdomen. ‘This feature is strongly marked in the Raninide.

On the other hand, it is generally recognised that the Raninidee exhibit a
number of Macruran characters which, on the assumption that the Brachyura
are descended from the Macrura, are primitive. Among the more important
of these are :—The comparatively well-developed rostrum, which, however, is
absent in Cosmonotus. The proximal segments of the second antenna are
not fused with the epistome, a primitive feature also shared by the Dromiacea.
The presence of a sternal canal and the relatively well-developed anterior
thoracic apodemes. The narrow and keel-like posterior thoracic sternites.
The oviducal apertures opening on the coxz of the third pereiopods, another
primitive character shared with the Dromiacea. The external lobes of the

28 DR. G. C. BOURNE ON THE RANINIDH:

second maxille are narrow, the inner lobes normal (Ortmann). Finally, as
I shall show, the nervous system is more Astacuran in type than in any
other Brachyura. In addition, the Raninide exhibit a number of features
peculiar to themselves, and these I shall discuss in detail in the later part of
this paper.

For the present it is only necessary to refer to the theory originally pro-
pounded by Huxley and brilliantly sustained by Bouvier in his essay, ‘ Sur
Porigine homarienne des Crabes’ (22), that the crabs are not descended from
Paguridee nor from Galatheidee nor Thalassinide, but directly from the
Astacura, and in particular from the Homaride through the Dromiacea.
“Les Dromiacés par Vintermédiare des Homolodromies ou de formes trés
voisines, dérivent directement des Homaridés jurassiques ou de Homaridés
qui leur ressemblent.” Exception being made of such crab-like forms as
Porcellana, Lithodes, and the Porcellanopagurus recently described by
Borrodaile, Bouvier’s theory of the origin of crabs is now generally accepted,
and it follows that, if the Raninidee be true crabs, they must be descended
from a Homarid or Nephropsid ancestor by way of the Dromiacea. ‘Thus
Bouvier (22) has given convincing arguments for the derivation of the
Dorippide from the Dromiacea, and Alcock (2) sees in his genus Cymonomops,
which differs but little from Cyclodorippe, a number of Raninoid characters
which, he holds, warrant the conclusion that ‘‘we have, in fact, in some of
these deep-sea forms the clearest evidence of the close relation of the Ranina
to the Dorippe type and quite suflicient justification for accepting de Haan’s
scheme of the Oxystomata, almost without modification.” Cymonomops,
however, is clearly a highly-specialised Cyclodorippid ; it has been placed in
this family by Bouvier, and its supposed resemblances to the Raninide are
of the most superficial character. If the last-named family were descended
from the Dromiacea it would hardly be through a Dorippid line, but it is my
object to show in the following pages that the Raninidee, though by defini-
tion they must still be included among the crabs, are not derived from a
Dromiid ancestor, but have been evolved as an independent group from the
Astacura. In sustaining this thesis I shall place reliance first of all on the
nervous system, and secondly on the character and degree of development
ot the endophragmal skeleton.

Our knowledge of the nervous system of the Decapoda is not very exten-
sive, and what we do know is due largely to the researches of Bouvier (21).
He has demonstrated that the degree of concentration of the nervous system
varies so considerably in the several groups of Decapoda reptantia that it
cannot of itself be taken as a sure guide of affinity, but I shall be content
to found my argument upon a sentence taken from his essay, ‘ Sur lorigine
homarienne des Crabes.’ ‘* Le systtme nerveux des Crustacés décapodes, en
effet, subit une concentration longitudinale réguliére & mesure qu’on d’éloigne

A STUDY IN CARCINOLOGY. Pag)

des Macroures pour se rapprocher des Brachyures et la réduction qu’il présente
dans le sens de la longueur, est un indice, non pas des aftinités précises de
Vanimal, mais du degré d’évolution cancerienne auquel il est arrivé.” If we
accept this proposition, and for my own part I accept it without reserve,
it must follow that a crab with a nervous system of which the ganglia are
more concentrated in the longitudinal direction is farther removed from its
Macruran ancestry than one in which the ganglia are far less concentrated,
and that the former cannot be a progenitor of the latter.

Of the Raninidee I have been able, thanks to the excellent state of preser-
vation of the example sent me by Prof. Kishinoye, to make a thorough study
of the nervous system of Ranina dentata and a sufficient study of that of
Lyreidus tridentatus, of which I possessed two well-preserved examples.
I have also dissected the nervous systems of Notopus dorsipes and Notosceles
chimmonis, and was able to ascertain that they are of the same character and
the ganglia fully as much extended in the longitudinal line as in Ranina
and Lyreidus, but their state of preservation did not admit of great accuracy
in making out details.

Pl. 4. fig. 8 is a drawing of the nerve-ganglion chain of Ranina dentata,
as seen from above, the whole series being represented as flattened out in one
plane. Pl. 4. fig. 9 shows the actual position of the nerve chain as seen
from the side, and its relation to the endophragmal skeleton. Owing to the
deep infolding of the sternal apodemes of the posterior thoracic somites,
the posterior thoracic ganglia and the abdominal ganglia closely applied to
them are directed nearly vertically upwards.

The cerebral ganglion is relatively large and quadrangular in outline, and
the ocular, antennulary, and antennary nerves are of large size ; otherwise it
does not present any special features. A small pair of nerves passes forward
to the rostrum. The circum-cesophageal connectives are long and stout.
The small ganglia on either side of the cesophagus give off a fairly stout
nerve to dilator muscles of the cesophagus as well as the more slender
connectives of the stomatogastrie system. The post-cesophageal commissure
is well marked. The subcesophayeal ganglion mass is large and goblet-
shaped in outline, as seen from above, and gives off five stout nerves on either
side. These nerves are somewhat swollen at their origins, and the last two
come off rather apart from the rest and give the appearance of originating
from a separate ganglionic centre. Otherwise the six ganglia supplying the
mandibles, first and second maxille, and three pairs of maxillipeds are
indistinguishably fused in the subcesophageal mass. It will be noticed that
the mandibular nerve does not spring from the subcesophageal ganglion but
from the cireum-cesophayeal connectives, some way in advance of the ganglion.
The ganglion pair of the chelipeds is distinet, but so closely fused to the
subcesophageal mass that the connectives nniting the two are indistinguishable.

30 DR. G. C. BOURNE ON THE RANINID®:

The succeeding ganglion pair, supplying the first pereiopods, is situated well
back in the thorax, and is connected with the pair supplying the chelipeds by
two long connectives separated from one another by a well-marked interval.
The ganglion-pair of the second pereiopods is unitéd with the preceding pair
by similarly distinct connectives, but only one-third as lone as those con-
necting the chelipeds and first pair. The connectives between the second
and third ganglion-pairs of the pereiopods are shorter, but set widely apart,
leaving an oval space for the passage of the large sternal artery. he two
last thoracic ganglion-pairs, supplying the third and fourth pereiopods, are
fused together, and the connections between them cannot be distinguished,
but the separate ganglia can be clearly recognized. Closely applied to the
last thoracic ganglion-pair is the chain of abdominal ganglia, completely
withdrawn into the thorax and represented by a short and thick nervous mass
divided by distinct transverse constrictions into five segments. The terminal
segment apparently represents the fifth and sixth abdominal ganglia fused
together. Paired nerves issue from each abdominal ganglionic segment, and
pass backwards closely bound up in the same connective-tissue sheath as the
fused mass of abdominal ganglia, but they are represented as spread out by
dissection in fig. 8. The nerves issuing from the first abdominal ganglion
segment can be traced to the first pair of modified copulatory abdominal
limbs of the male. In this condensed and abbreviated representative of the
abdominal nerve chain no trace of connectives nor of ihe paired stucture of
the ganglia can be detected, The abdominal ganglia and the nerves passing
from them to the abdomen lie in the concavity of the narrow sella turcica
posterior formed by the deeply infolded sternal apodemes of the last two
thoracic segments. Pl. 4. fig. 10 is a representation of the ventral nerve
chain of Lyreidus tridentatus. The general plan is similar to that of Ranina,
but the details are different. The subcesophagal ganglion is an ovoid mass,
formed by the fusion of the ganglia of the mandibles, maxille, and maxilli-
peds. As I could only count five nerves given off from this ganglion on
either side, I conclude that the mandibular nerve issues, as in Ranina, from
the cireum-cesophageal connectives, but I was not able to verify this point.
The ganglion-pair of the chelipeds instead of being approximated to the
cesophageal as in Ranina, is placed further back in the thorax and is united
with the subcesophageal by a thick cord, in which I could not trace any
separation of the connectives in the middle line. My specimen was none
too well preserved, but I can say for certain that these connectives are bound
together in the same connective-tissue sheath. The ganglion-pairs of the
chelipeds and first pereiopods are distant and, as in /anina, their connectives
are separated by a distinct slit-like space. The posterior thoracic ganglia
are more closely approximated than in Ranina, and the short connectives
between the ganglion-pairs of the first and second pereiopods are bound

A STUDY IN CARCINOLOGY. 31

together in the same connective-tissue sheath posteriorly, leaving only a
small circular passage between them anteriorly. The connectives between
the next succeeding ganglion-pairs are pretty widely divaricated to admit
the passage of the sternal artery, and the last two thoracic ganglion-pairs are
so intimately fused that the separate ganglia cannot be distinguished. The
abdominal ganglia are precisely like those of Ranina, but somewhat larger in
proportion to the rest of the ventral chain. In Notopus dorsipes and
Notosceles chimmonis the ventral nervous chain presents much the same
characters, but my examples of these species had lain for many years in spirit
and were too rotten to admit of exact study. In Notopus the ganglion-pair
of the chelipeds is separated from the subcesophageal ganglion mass by an
interval somewhat shorter than in Lyreidus, but the connectives joining them
are distinctly paired and are not included in a common connective-tissue
sheath.

The above descriptions, with the figures illustrating them, show that the
thoraco-abdominal nerve system of the Raninide approximates more nearly
to the Macruran type than does that of any other Brachyuran—indeed, more
nearly than any Anomuran except Hippa, in which genus, as I have
ascertained by dissection, the thoracic ganglia are spaced fairly evenly along
the longitudinal line and are united by distinctly paired objectives, except
the last two which are fused together. The abdominal chain also in Hippa
consists of at least five fairly distant ganglion-pairs united by distinctly
paired connectives, but I have not yet finished my observations on this genus,
and must postpone the description of its nervous system to another paper.

As compared with the Decapoda, of which the nervous systems have been
accurately described and figured, the abdominal ganglia of the Raninidee
most nearly resemble the condition found in Dromia vulgaris, as described
and figured by Bouvier (22). I have myself verified the accuracy of this
deseription. But in Dromia the thoracic ganglia, though distinct, are
closely approximated in longitudinal series, and form a sort of oval beaded
ring round the perforation for the passage of the sternal artery, as is the
case in Calappa among the Oxystomes, in Corystes, and generally in the
Catametopan section of the Brachygnatha. In respect of the longitudinal
concentration of the thoracic ganglia, the Raninide exhibit a much more
primitive condition than Dromia. Similarly, the extended thoracic chain of
the Raninidee is ina much more primitive condition than are the approxi-
mated though distinct thoracic ganglia of Galathea and Porcellana among
the Anomura, though these are more primitive as regards the abdominal
chain. ‘The same may be said in respect of Hupagurus, in which the thoracic
ganglia are in some respects more completely fused together than in the
Galatheidee. If we carry on the comparison to the Macruran types, the
Raninidee cannot be brought into close relation with the Lobsters, and still

32 DR. G. C. BOURNE ON THE RANINIDA:

less with the Crayfishes, because in the first-named the thoracic connectives
are separate and show but little tendency to lateral fusion. But of all the
Macrura they most closely resemble Nephrops in this and other respects.
In both the subeesophageal ganglion mass is in a very similar state of con-
centration, the differences being that in Ranina the mandibular nerves are
given off from the circum-cesophageal connectives (this may be correlated
with the great length of these connectives) and in Nephrops the ganglion
pair of the third maxillipeds retain a certain distinctness. The connectives
between the subesophageal and the ganglion-pair of the chelipeds are very
nearly of the same length and in the same condition of incipient lateral
fusion in Vephrops as in Notopus, though the concentration is carried further
in the lateral sense in Lyreidus and in the longitudinal sense in Ranina.
The connectives between the ganglion-pairs of the chelipeds and _ first
pereiopods are much longer in the Raninidee than in Nephrops, but are
equally distinctly paired in both and show no tendency to lateral fusion.
The same may be said of the connectives next following. In the Raninidee.
the last two thoracic ganglion-pairs are more (Lyreidus) or less (Ranina)
closely fused together. These differences apart, if the long chain of six
abdominal ganglion-pairs of Nephrops were shortened up to the condition
found in the Raninidee and withdrawn into the thorax, there would be very
little difference between the two nervous systems. So far, then, as this part.
of their anatomy may serve as a guide, the Dromiacea have progressed
further from the Macrurous in the direction of the Brachyurous type than
have the Raninidee, but the latter retain many distinct Macrurous features.
and exhibit specially close affinities with Nephrops.

It will be noticed that there is a marked tendency for the thoracic ganglia
to form two groups, one at the anterior the other at the posterior end of the
thorax, in the Raninidee. This must be correlated with the powerful develop-
ment of the first, second, and third pereiopods as digging or swimming
organs. These limbs are exceedingly strong, and are actuated by largely
developed muscles enclosed in remarkably deep cavities formed by the endo—
phragmal skeleton. The size and extent of these muscle cavities as compared
with those of other Brachyura is one of the most striking features when one.
undertakes a dissection of any member of the Raninidze, and the abundant
musculature of the limbs makes the dissection of the nervous system difficult.
It is also worthy of remark that in Lyreidus the last pair of pereiopods are
very weak and attenuated and the corresponding ganglion-pair is practically
unrecognizable, whereas in Fanina, in which the last pair of pereiopods
differs little in size from the rest, its ganglion-pair is distinct and emits.
large nerves to the limbs.

Two good figures are given in Cuvier’s ‘Régne Animal,’ but otherwise
de Haan is the only author who has dealt with the endophragmal skeleton of

A STUDY IN CARCINOLOGY. 33

the Raninide. He says (34, p. 136), “thoracis interior structura offert
sellam turcicam uti tantum in Brachyuris, atque canalem sternalem uti in
Macrouris ceterim tantum invenitur. Apodemata singula tam sterni quam
epimerorum ab utraque parte medio thorace junguntur in quoque articulo
sternali, canalem sternalem constituentia vertebrarumque formam plane
simulantia. Apodemata cephalica pariter adsunt quinque transyersa, ita ut
Ranine hoe respectu opposites sunt Dromiis, in quibus apodemata thoracica
et cephalica separatim versus centrum commune petunt.” We have here
a combination of Macruran and Brachyuran characters which, when the
ancestry of the Raninidz is in question, deserves close study. As a prelim-
inary to this part of the subject, | must go in some little detail into the
structure and morphology of the endophragmal skeleton. It was described
both for Macrura and Brachyura by Milne Hdwards (25), and since his
day does not seem to have been studied in further detail. We have, sub-
sequently, the brief but very accurate description of these structures in the
Crayfish by Huxley (38). His figures, however, are not drawn from a point
of view that clearly illustrates the structures described, and I shall endeavour,
with the aid of new figures, to make clear some points that are necessary for
a detailed comparison with the homologous structures in crabs. Tig. 11 is a
ventro-lateral view of the sterna and the endophragmal skeleton of the second
and third pereiopodal segments of the common Crayfish, seen from the left
side ; the thorax has been tilted over to the right; the outlines of the
articular cavities of the limbs have been thickened to make them more con-
spicuous. Ventrally we see the keel-shaped sternites, and at the outer side
of the posterior end of each is a small articular socket, into which fits a
corresponding knob on the coxa of the appropriate limb. Above are the
corresponding epimera, separated from one another by a groove which is not
a suture but a deep infolding of the cuticle, technically known as an apodeme.
If we follow this groove down below the upper limits of the articular cavities
of the limbs, we see that it bends forwards, then turns rather abruptly back-
wards, and seems to end just below a little knob which fits into a corre-
sponding cavity on the coxa of the appropriate limb. Thus the articular
knobs, which I shall eall strophidia, are borne on prolongations of the anterior
ends of the pleura, the articular cavities, which I shall call strophingia,
on the posterior ends of the sterna™. The line indicating the epimeral

* These small articular knobs and cavities have keen described in some detail, and their
mechanical action has been described by Herrick (37), but though often figured, they seem
to have been neglected by other authors. As I shall have occasion to refer to them
frequently and to use them as landmarks, I have found it necessary to give them names, and
call the socket the strophidiwm, the pee which fits into it the strophingium. The names are
derived from the peg (arpodiyy:ov) and socket (otpodevs) hinge of a Greek door. For the
correct formation of the diminutive orpodevs, otpodeidSiov, strophidium, I am indebted to
Mr. Eric Barber of Exeter College.

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 3

34 DR. G. C. BOURNE ON THE RANINIDZ:

apodeme seems to end just below the articular knob : in reality it is prolonged
beyond it, and can be traced along the inner and hinder edge of the twisted
Jamella or arthrophragm that separates each articular cavity from the one
next behind it. The arthrophragm, in fact, is a double fold of the cuticle or
apodeme, continuous with the pleural apodeme; the two edges of the fold
run down rather to the inside of the hinder edge of the arthrophragm, and in
the entire animal pass respectively into the arthrodial or joint membrane of
the lmb in front and the limb behind. At the lower end of the arthrophragm
the groove forms the apparent boundary between the contiguous sterna,
but here again there is no suture; the sterna are really continuous, and
are only separated by apodemes. All this is familiar, but as separate names,
epimera, sterna, arthrophragms, etc., are given to the various parts, I have
thought it desirable, in view of the comparisons that are to follow, to emphasize
the fact that these names only apply to local areas and indurations separated
from one another by more or less deep infoldings of a continuous cuticle.
In the upper and lower parts of the arthrophragm its apodeme is shallow, and
consequently the arthrophragmal partition extends only a little way into the
cavity of the body; but at about the middle of its course the apodeme
becomes very deep and gives rise to a lamina running upwards, forwards,
and inwards towards the middle line, short of which it expands into
horizontally flattened plate with jagged edges, which makes more or less
intimate union with its fellow of the opposite side. The lamine in question
are the endosternites : they and their flattened summits form the walls and
roof of the so-called sternal canal within which the thoracic nerve-ganglion
chain lies. A sternal canal is found in the Macrura reptantia and in some
Anomura, but, with the exception of the Raninide, it does not occur in the
Brachyura. PI. 4. fig. 11 also shows the epimeral apodemes or endopleurites.
As the articular cavities slant backwards, the endopleurites alternate in
position with the endosternites, and, as is familiarly known, each endopleurite
divides internally, sending a posterior branch to unite with the endosternite
of the segment behind, an anterior branch to unite with the endosternite in
front. The result of all these complicated arrangements is that the cavity of
each limb communicates internally with two imperfectly delimited chambers,
which may properly be called muscle-cavities as the abductor and adductor
muscles of the limbs are attached to the several apodemic ingrowths that
constitute their walls. In the Macrura, as the articular sockets of the limbs
look nearly ventrally and the epimera are not much inclined inwards from
the vertical, the muscle-cavities lie in nearly the same horizontal plane and
alternate with one another. A consideration of the relative positions of the
dorsal strophingia and ventral strophidia by which the coxa of the limb is
hinged to the articular frame will show that the outermost and slightly
dorsal of the two muscle-cavities encloses the abductor muscles whose action

A STUDY IN CARCINOLOGY. 35

moves the coxa outwards, backwards, and slightly upwards ; the innermost
and slightly more ventral muscle-cavity encloses the adductor muscles whose
action is the reverse of the abductors.

On comparing the thoracic skeleton of a crab such as Carcinus menas
with that of a crayfish or a lobster, the differences are considerable. The
carapace being removed, the epimera are seen to slope inwards towards the
median dorsal line like the roof of a house. he sterna instead of being
narrow and keel-like are widely extended laterally, and form by their union
the broad ventral plastron across which run a series of transverse grooves
indicating the sternal apodemes. As a consequence the sockets of the limbs
are placed not ventrally but on the sides of the body, forming a row which is
not straight but curved upwards at both ends. In some cases the hinder end
of the row curves upwards, the fourth pereiopods being dorsal in position.
Although Dromia is in many respects a primitive crab retaining many
Maeruran features, its endophragmal skeleton is so characteristically crab-
like that it will serve for comparison and contrast with that of the Crayfish,
Pl. 4. fig. 12 is a drawing of the left side of the thorax of Dromia vulgaris
after removal of the carapace. The thorax has been slightly tilted over to
the right to bring it as nearly as possible into comparison with PI. 4. fig. 11,
and for simplicity’s sake the perspective of the ventral surface of the sternum
has been omitted. Fig. 15 (PI. 5) is a posterior view of the same preparation
which illustrates some features not clearly shown in fig. 12.

As in Astacus, the strophingia and strophidia, the former situated at the
upper ends the latter at the lower ends of the arthrophragms, serve as
convenient landmarks. ‘The most conspicuous feature in the crab is the
great depth of the arthrophragms, especially those between the muscle-
cavities of the segments of the second and third and third and fourth
pereiopods (intersegments xii/xiii and xiii/xiv). It is also characteristic
that- the three posterior arthrophragms slope forward, the two anterior
arthrophragms backward, in such wise that the flexor muscle-cavities of
which they form the walls converge towards a point situated in the middle
of the thorax. A third characteristic feature is that, owing to the low pitch
of the gabled roof formed by the epimera (see Pl. 5. fig. 15) the abductor
muscle-eayities lie above and not, as in the Macrura, in the same horizontal
plane as the adductor cavities. All these features, which ge a long way to
make up the characteristic cancroid facies, are correlated with the habits of
erabs. They run much more actively than any Macrura, and the attach-
ments for the muscles of their pereiopods are proportionally increased. ‘To
provide these attachments the intersegmental apodemes are very deep,
particularly on the sternal surface and on the sides of the body, and the
arthropbragms, instead of being mere flattened bars as in the Macrura, are

conspicuous partitions extending deep into the body. But their internal

36 DR. G. C. BOURNE ON THE RANINIDZ:

edges stop some way short of the middle line and are not prolonged into
conspicuous endosternites. Towards their lower ends they do, indeed, give
off triangular or thorn-shaped offsets (such an offset is shown in the inter-
segmental arthrophragm x/xi in PI. 4. fig. 12), but these are not expanded
into flattened summits and do not come into contact with their fellows of the
opposite side of the body. Therefore there is no sternal canal. The
epimeral apodemes or endopleurites are well-developed in crabs and, as in
Macrura, they bifurcate at their inner ends, the anterior bifurcation uniting
with the intersegmental apodeme in front, the posterior bifurcation with the
intersegmental apodeme next behind, as is shown in fig. 12. It should be
noted that in all these respects Droméa has departed so far from its Macruran
ancestry and has taken on so completely the characters of crabs, that it has
been possible to make use of it as a typical example of this aspect of cancroid
structure.

If we now bring the Raninide into comparison, taking as our example
Ranina dentata, we find a curious admixture of crab-like and Macruran
characters, but the latter predominate and the former must be called
deceptively crab-like, for a detailed examination shows that they must have
originated independently, in relation to the habits of the animal, and not by
way of inheritance fromany other recognized crabs.

In the first place, as may be seen by a comparison of PI. 5. figs. 13, 14, and
15, the contour of the thorax of Ranina as seen from behind is more like
that of a lobster than like that of a crab. In the crab the width of the
thorax largely exceeds its height ; in the lobster the height clearly exceeds
the width ; in Ranina the height is rather greater than the width. Looking
at the sternal surface, Ranina has a broad ventral plastron and in so far
resembles a crab, but the most eursory inspection shows that it is made up
almost entirely of the greatly enlarged and flattened sternum of the tenth
segment. The more posterior thoracic sterna, particularly those of the
twelfth and thirteenth segments, are narrow ‘and keel-like ; indeed, they are
proportionately narrower than in the lobster. The broad sternum of the
tenth segment is flat and has no vertical depth; that of the cleventh
seoment has the form of a pair of aliform expansions extending along the
posterior edges of the twelfth sternum and making up with it the ventral
plastron. Posteriorly these aliform sclerites meet in the middle line to form,
the short and somewhat thickened body of the eleventh sternum, tilted
somewhat upward. The twelfth sternum is very narrow, of considerable
vertical depth, and is curved upwards. The thirteenth and fourteenth sterna
are still deeper vertically, and are so much tilted upwards that the cavities of
the third pair of pereiopods look nearly directly backwards and the eavities
of the fourth pair look backwards and upwards (Pl. 5. tig. 14). The large
articular cavities of the clelipeds are placed laterally, but those of the first,

A STUDY IN CARCTNOLOGY. oN

and second pereiopods are almost ventral. Thus, in the middle of the thorax,
the articular sockets of the limbs have the same position as in the Astacura,
but at the two ends their position is similar to that seen in crabs (PI. 5.
fig. 17). The epimera of Lanina are of great depth and exhibit peculiar
features that will be described later ; for the moment it is sufficient to note
that, owing to the size and depth of the epimera, the arthrophragms are
relatively short and as in the Astacura are somewhat narrow curved Jaminz
separating the articular cavities. But, as also in the Astacura, the arthro-
phragmal apodemes are deeply extended into the body for a limited space on
either side of the mid-ventral line. These extensions are the so-called endo-
sternites, and in all the Raninidz they are very largely developed, the largest
of them, namely the intersegmental pair xi/xii, extending upwards as much)
as two-thirds of the entire depth of the thorax. As may be seen in Pl. 4.
fic. 9 and Pl. 5. fig. 16, the four posterior pairs of endosternites slope
sharply /orwards from their points of origin from the arthrophragms, but the
intersegmental pair ix/x stands nearly vertically and the pair next in front
slopes somewhat backwards. The upper ends of the endosternites are widely
expanded, and their innermost expansions, the mesophragms of Huxley,
unite in the middle line to form the roof of a very well-marked and very
deep sternal canal, at the bottom of which lies the ventral nerve-ganglion
chain. This is a wholly Macruran character and, as we have seen, it is
unrepresented in Mromia. In this respect, therefore, as in the nervous
system, the Dromiacea have departed further from the Astacuran and have
advanced further towards the cancroid type of structure than have the
Raninidz. The former cannot be in the direct ancestral line of the latter.
To return to a consideration of the epimeral elements of the thorax. The
epimera (fig. 17), with the exception of the last thoracic, have a great
extent dorso-ventrally and present features absolutely distinctive of the
Raninide. That of segment ix stands nearly vertical and the broad
epimeron of segment x nearly so. These two do not present any specially
abnormal features, but the three succeeding epimera slope sharply forwards,
so that their upper extremities converge towards the upper posterior angle
of the tenth epimeron, and their extensive lateral surfaces are divided into
dorsal and ventral moieties by a ridge running obliquely upwards from the
hind margin of the articular cavity of the cheliped to the insertion of the
abdomen. Against this ridge the lower edge of the branchiostegite fits
closely and is held in position. by two projections, one near the anterior edge
of the eleventh, the other at the posterior edge of the thirteenth epimeron,
just in front of the articular cavity of the last pereiopods. The lower moieties of
the epimera of the eleventh, twelfth, and thirteenth segments are therefore
exposed on the surface, and form a somewhat excavated and roughly quadri-
lateral area between the coxee of the limbs and the branchiostegite. The

38 DR. G. C. BOURNE ON THE RANINIDZE:

epimeral walls in this area, being exposed, are strongly calcified. This
exposure of a large portion of the epimeral surface is a remarkable and
unique feature in the Raninide. Hven in the Astacura, where the lower
edge of the branchiostegite is free, it reaches down to the bases of the limbs,
and in all crabs the lower edge of the branchiostegite is so closely adapted
to the coxze of the thoracic limbs as to leave no passage for water into the
branchial chamber behind the chele. This peculiarity did not escape the
attention of Milne Edwards, but subsequent authors seem to have paid very
little attention to it. If we now examine the interior of the thorax, we find
that the apodemes between the ventral and exposed parts of the three
thoracic epimera in question are shallow and form nothing more than two
inconspicuous thickenings internally, strengthening the framework of this
part of the flanks but not projecting into the cavity of the body. But the
apodemes above the line of the adherence of the branchiostegites are yery
deep and form well-marked endopleurites which, as usual, bifureate and their
branches unite with the expanded summits of the endosternites next in front
and next behind them. It appears that the excessive depth of the sternal
canal is correlated with the restriction of the endopleurites to the upper
moieties of the epimera or, to put it in another way, with the downgrowth of
the ventral edges of the epimera beyond the line of the attachment of the
branchiostegite. The ventral extension of the epimera is an adaptive
feature, and receives its explanation when the habits of the animals are taken
into account. The Raninide are digging crabs, with large and powerful”
chelipeds and first, second, and third pereiopods. In many the fourth pair
of pereiopods is weak and slender, though in Ranina this last pair is not
much weaker than the others. The powerful digging limbs imply powerful
abductor muscles and large adductor muscle-cavities. Reference to fig. 9
shows that these cavities are in fact unusually deep, and their extent is due
to the great length of the endosternites within the body and to the large
ventral extensions of the epimera forming their outer walls. The same
figure shows that, as in crabs, the abductor muscle-cavities lie above and not
alongside of the adductor muscle-cavities. But cancroid as this character
is, it has clearly a very different origin from the similar arrangement
in crabs.

To turn to another feature noted by de Haan, “ thoracis interior offert
sellam turcicam uti tantum in Brachyuris.” What is this sella turcica ?
The name is due to Audouin and Hdwards, and the latter author has
described the elements entering into its composition with great minuteness
on p. 34 of the first volume of the ‘ Histoire Naturelle des Crustacés.’
Paraphrasing his description, the sella turcica posterior is a little arch or
vault, concave from side to side and convex from front to back, formed by
the forward extension and fusion together in the middle line of the nearly

A STUDY IN CARCINOLOGY. 39

horizontally disposed intersegmental apodemes xiv/xy ; in other words, the
apodemes separating the last thoracic from the first abdominal segment.
This sella turcica is not present in all crabs, but where present the arrange-
ments indicated above are due to the presence of a deep longitudinal mid-
ventral apodeme in the last one or two thoracie sterna. The effect of this
deep folding-in of the terminal thoracic sterna is to raise the hinder end of the
last thoracic sternum and to throw its arthrophragmal apodemes forward into
a horizontal position. The edges of the arthrophragms are thus brought into
contact, and fuse in the middle line. It will be understood from the above
description that the exhibition of a sella turcica is dependent on the
formation of a more or less deep apodeme in the mid-ventral line of the
penultimate thoracic sternum.

Figs. 13, 14, and 15 are drawings of posterior views of the thorax of the
lobster, of Ranina dentata, and of Dromia vulgaris respectively. In Homarus
—I should more correctly write Astacus !—the penultimate thoracic sternum
is large and escutcheon-shaped in posterior view: it has a distinct mid-
ventral apodeme and the central area or inescutcheon is membranous. The
terminal sternum consists of two distinct quadrangular pieces united at their
inner angles, and above these is a curved transverse bar which appears to be
a special calcification of the membrane connecting the last thoracic and first
abdominal segment. It is not present in Nephrops norvegicus nor in the
Crayfish (Potamobius). On either side of this bar the intersegmental arthro-
phragms xiv/xv diverge outwards to unite by slender ends with the large and
peculiarly shaped epimera of the fourteenth segment. From the front edge
of the sternum just where it passes into the arthrophragm of each side, the
two narrow and nearly horizontal episternites run forward diverging from
one another ; only their tips appear in the drawing. There is no sella turcica.
In Ranina the penultimate thoracic segment, though relatively smaller than
in the lobster, is similarly somewhat escutcheon-shaped. Both it and the
ante-penultimate sternum have a deep mid-ventral apodeme, indicated by the
median suture in fig. 14 and clearly shown in side view in fig. 9. The last
thoracic sternum, clearly recognizable by the position of the two strophidia,
is aliform with tapering wings diverging from one another and passing
forwards and upwards to unite with the last thoracic epimeron. The arthro-
phragms xiv/xv are triangular plates arising by their bases from the front
edges of the V-shaped arms of the last thoracic sternum. Their apices are,
in the natural position of the animal, directed forwards and downwards ;
their inner surfaces are concave and their admedian edges meet and unite in
the middle line, thus forming the sides and floor of the deep trough called
the sella turcica. Fig. 9 shows the relation of the abdominal nerve ganglia
and the nerves given off from them to this trough. The anterior ends of the
arthrophragms in question diverge from one another and, curving outwards

40 DR. G. C. BOURNE ON THE RANINIDA:

beyond the intersegmental endopleurites xili/xiv, unite with the inner edges
of the lower part of the endopleurites xii/xili. The sella turcica of Nanina
is therefore of large extent, larger than in so typical a crab as Carcinus
menas. Among the various Brachyura that I have studied, it most nearly
resembles in form and extent that of Calappa. But, as mentioned above, the
sella turciea is not found in all crabs. Fig. 15 represents a posterior view of
the thorax of Dromia vulgaris. The sterna of the twelfth, thirteenth, and
fourteenth segments can easily be recognised by the strophidia on their outer
extremities ; the large pair of ventro-lateral projections in front of them
belong to the eleventh sternum; their strophidia, being on their anterior
face, do not appear in the drawing. The fourteenth (last thoracic) sternum
does not differ much in shape from that of Fanina and offers resemblances
to that of tle lobster. The thirteenth and twelfth sterna are irregularly
shaped transverse bars, of which the outer portions are calcified and
distinctly separated by apodemes, but their admedian portions lose their
distinctness and pass into a median membranous pouch, the walls of which
are little if at all calcified. There is no trace of a median longitudinal
apodeme in this membranous representative of the penultimate and ante-
penultimate thoracic sterna, consequently there is no sella tureica. ‘The
arthrophragms of the intersegment xiv/xv are seen converging forward from
the anterior edges of the Y-shaped arms of the last thoracic sternum, and in
other crabs they unite above with the epimera of their own segment, in front
with the arthrophragms of the intersegment in front, but in the absence of
the ventral apodeme they fall far short of meeting in the middle Jine to form
a sella tureica. In this respect, then, Dromia has progressed less from the
Macruran towards the Brachyuran condition than Ranina, and this might be
fastened upon as an argument in favour of the derivation of the Raninidze as
of all other crabs from a Dromiacean ancestor, but the transversely elongated
twelfth and thirteenth sterna of Dromia are quite Brachyuran in character
and very different from the narrow and keel-shaped corresponding sterna
in the Raninidee.

On the balance the evidence afforded by the structure of the hinder -part
of the thorax is against the Dromiacean ancestry of the Raninide.

Another crab in which the sella tureica is absent is Philyra levis. In view
of de Haan’s emphatic opinion that the Leucosiidee are clearly united to the
Raninidee through Lyretdus, an opinion which has found favour with subse-
quent authors, the marked differences in the structure of the thorax demand
attention. A detailed comparison of the endophragmal skeletons of the
Raninidz and Leucosiide is best reserved for the discussion of the possible
derivation of the latter from the former family in another part of the paper,
but it may be conveniently pointed out in this place that in Philyra the
plastron is broadest at its posterior end; the intra-sternal apodemes are nearly

A STUDY IN CARCINOLOGY. — 4]

transyerse to the long axis of the body (with the exception of xiii/xv and
xiv/xv, which are, directly obliquely backwards from the middle line), are of
no great vertical height, and give off no offset to form a sternal canal. The
abductor muscle- cavities are of great length and slope backwards, converging
to a point at the postero-external angle of the thorax on either side. There
is no mid-ventral longitudinal apodeme and the arthrophragm xiv/xv is
rudimentary, so there is no trace of a sella turcica. The arrangements of the
several parts are as widely different as possible from those in Lyreidus or
any other of the Raninide.

Though de Haan in the quotation given above (p. 33) draws a contrast
between the Raninide and the Dromiacea in the matter of the “apodemata
cephaliea,” i.e. the pleural and sternal apodemes of intersegments v/vi-1x/x,
the Raninidze show more resemblance to the Dromiacea than to the Astacura
in this region, and the two first-named groups differ largely from the last.
Passing over the obyious differences due to the presence of a broad and flat
shield-shaped sternum in the Raninide, attention may be directed to the
apodemes entering into the composition of the anterior part of the sternal
canal. The reader will remember that in the Astacura, of which Nephrops
norvegicus may be taken as a convenient example, the pleural and sternal
apodemes of the three intersegments in front of ix/x, though some are missing
and others feebly developed, unite to carry forward the sternal canal towards
the region of the mouth. In front of them both the sternal and pleural
apodemes, but particularly the latter, of intersegment v/vi are very largely
developed, and their expanded internal extremities combine to form a
stout platform between the subcwsophageal ganglion mass and the stomach,
known to the older carcinologists as the sella turcica anterior. Huxley
named this platform the cephalic apodeme. In Notopus and Ranina, the
only two genera of the Raninide in which I have been able to study
these structures in detail, the enosternites and endopleurites of intersegment
ix/x are normal (figs. 9, 16, & 17), the former standing nearly vertically above
the plastron and ending above in slightly expanded summits which are
joined in the usual way by the anterior branches of the corresponding endo-
pleurites. The endosternites of intersegment viii/ix are small and directed
backwards to touch the summits of the endosternites behind them. The
endopleurite of intersegment viii/ix is minute, is pot,branched, and converges
towards the point where endosternites viii/ix and ix/x come into contact.
Intersegment vii/viii shows only rudimentary endosternites, but its endo-
pleurites are fairly well developed, exhibiting a very short anterior branch
and a posterior branch which makes connection with endosternite vili/ix.
In intersegment vi/vii there is no recognizable endosternite, but the endo-
pleurite is present in the form of a small incurved lamella. Intersegment
v/vi is largely membranous, and neither endosternites nor endopleurites can

42 DR. G. C. BOURNE ON THE RANINIDA:

be distinguished in it. Hence there is nothing to correspond with the selle
turcica anterior of the Astacura. In Dromia vulgaris much the same elements
can be identified as in the Raninidz, but the sternal and pleural apodomes
of intersegment ix/x do not meet to form a sternal canal. Similarly the
sternal and pleural apodemes of intersegment viii/ix unite on each side to
form a curved shelf running inwards from the thoracic wall, but leave a wide
space open in the middle line. The pleural apodemes vi/vii are fairly well
developed and project some way inwards and backwards, but are far from
approximating in the middle line. The intersegment v/vi is fairly stout, but
is simply an arthrophragm without any ingrowths that can be distinguished
as endosternites, and though the sixth pleura are “well developed, they have
no inwardly projecting apodemes separating them from the membrane that
corresponds with the fifth pleuron. In all these respects Dromia has
departed further from the Astacuran condition than have the Raninidze. In
another matter also, the Raninide stand closer to the Astacura. In the last
named, for example in Vephrops, the ninth, eighth, and seventh sterna do not
lie in the same straight line as the thoracic sterna behind them, but are
directed upwards, so that a line drawn through them forms an angle of about
140° with a line drawn through the hinder ends of the more posterior
thoracic sterna.

In the Raninidée, as shown in fig. 17 for Notopus, a line drawn through
the seventh, eighth, and ninth sterna forms an angle of 134° with a line
drawn through the tenth and eleventh sterna. In Dromia the corresponding
lines meet at an angle of 108°. This is another character in which the
Dromiacea have departed further from the Astacuran type than the Raninide,
but, as regards the six post-oral seoments, it would not be quite accurate to
say that they have advanced further towards the cancroid type. There are
certain points of resemblance between the Dromiacea and the Raninide
which may be taken as evidence of but slight modification from the common
Astacuran stock from which both have descended : not, however, as evidence
that one group has descended from the other. The evidence so far tendered
shows that the Raninidz in their nervous system and in the characters of the
endophragmal skeleton are much more nearly akin to the Astacura than are
Dromiacea ; they are, therefore, the more primitive group and cannot have
descended from the less primitive. In both groups the departures from the
Macruran type as exhibited in the first six post-oral segments are due to the
acquisition of a broad sternum in the tenth segment and the upward cant of
the sterna immediately in front of it, with which are associated the retreat
of the sub-cesophageal ganglion mass into the thorax (see fig. 9 for Ranina),
the decrease in length of the carapace and the increase in width of its
anterior margin. The mouth is also brought to a more anterior and less
ventral position than in the Macrura,

A STUDY IN CARCINOLOGY. 43

Correlated with these modifications is the extent to which the epimera of
the post-oral segments are involved in the branchiostegal fold. In most
of the Brachyura that I have examined they are more intimately involved
than is the case either in the Raninide or in the Dromiacea, and the corre-
sponding sterna and arthrophraems are modified and exhibit features
departing more widely from the Astacuran type. A preliminary survey
leads me to believe that a study of the structure and relations of the first six
post-oral segments in the Brachyura will yield results of considerable classi-
ficatory value, but any attempt to discuss the subject would involve a long
digression, and it cannot be pursued any further in this place.

But, before instituting a comparison between the pre-oral sternites and the
adjacent parts of the exoskeleton in the Dromiacea and Raninide, it will be
necessary to make a considerable digression with the purpose of clearing up
doubtful points in the morphology of this region in the Decapoda reptantia,
and defining more precisely certain terms which, to say the best of it, are
used in an ambiguous sense by carcinologists.

In the Astacura and in Braciyura in general there is in front of the mouth
a plate, usually of considerable breadth laterally, but of variable extent antero-
vosteriorly, to which is given the name of “epistome.” Behind this is the
triangular, or in Brachyura the more often quadrilateral area known as the
“mouth-frame,” The confusion arising from the loose application of these
names is really remarkable. They are due to Milne Edwards and Audouin,
and it will be well to quote the former author im ewtenso (25, vol. i. p. 251):
‘““Hn arri¢re des fossettes antennaires, on voit une surface plane, plus ou
moins étendue, qui représente le troisitme anneau céphalique et qui poste le
nom d’épistome. L’espace occupé par V’épistome, les fossettes antennaires et
la base des antennes externes constitue ce que nous appelons la région
antennaire..... les parties latérales et inférieures de la carapace, que
nous appellerons régions ptérygostomiennes, sont toujours dirigées plus ou
moins obliquement en dehors et en haut, et sur la ligne médiane elles laissent
entre elles un espace vide qui est occupé par l’appareil masticateur et que
nous désignerons sous le nom de cade buccal ; tant6t ce cadre buccal A la
forme d’un quadrilatére assez regulier, tantot il est triangulaire, et c’est
toujours & sa partie antérieure que viennent se terminer les conduits efférens
des cavités branchiales.” The definition of the “epistome ” is sufficiently
exact ; it represents the sternum of the third cephalic segment, that is, the
antennary segment. Huxley (8) calls the epistome “the sternal region
which appertains, in part, to the antennie and, in part, to the mandibles.”
He was correct in recognising the inclusion of the mandibular sternal
element in the “epistome,’ so we must take the term to connote the con-
Joined antennary and mandibular sterna. Both Milne Edwards and Huxley
further recognised the distinctness of the antennulary sternum lying in front

44 DR. G. C. BOURNE ON THE RANINIDAE:

of the ‘‘epistome.” But the French naturalist, usually so exact in the
definition and use of anatomical terms, goes sadly astray in the application
of the terms under discussion, and has thus introduced a confusion which
has persisted in carcinological literature down to the present day. Thus in
vol. ii. p. 170 of the ‘ Histoire Naturelle des Crustacés’ we find it stated of
Dromia, “V’épistome est triangulaire,’ but clearly the triangular sternal
element here referred to is the antennulary sternum (see infra, p. 50). In
the same place he says, “le cadre buceal est 4 peu pres carré,” and it is clear
from an examination in this region in Dromia, that the “cadre buccal”
is the combined antennary and mandibular sternum, that is, by h's own
definition, the epistome. Again in vol. ii. p. 97 we find it stated of the
Oxystomatous crabs, “le cadre buceal est tout-a-fait triangulaire,’ and on
p- 102 of the Calappidee, “il n’y a point d’épistome distinct.” But a study
of these crabs shows that the triangular “cadre buccal” is chiefly made up
of the antennary sternum, 7.e., it is the epistome, which is said to be absent !

Among more recent authors, Calman (23, p. 257) writes: “the antennal
sternum is mainly represented by the epistome, a plate of varying shape
lying between the labrum and the bases of the antenne,”’ and further on
‘‘the buccal frame ..... is more or less distinctly delimited by the epistome
itself or by a transverse ridge which divides the epistome into two parts,
the epistome proper and the endostome or palate.’ He does not, with
Huxley, recognise the inclusion of the mandibular sternum, but there can be
no doubt that Calman identifies the greater part of the “ epistome ” with the
antennary sternum. <A little further on the same author makes the following
statement: “In all the Brachyura the rostrum or frontal plates ends down-
wards in the middle line a process which unites in front of the ophthalmic
and antennular sterna with the epistome and separates from one another
the basal segments of the antennules.” This represents accurately enough
the prevailing opinion on the subject, and indicates quite clearly that the
antennulary sternum is not to be regarded as a component part of the
“epistome.” But, whilst this part of the statement is accurate, the re-
maining part is most inaccurate, for an examination of a large number of
crabs belonging to diverse families shows that, with very few exceptions, the
downward process of the front or rostrum does not unite with the antennary
sternum, that is with the epistome, but with the antennulary sternum. The
fact is so obvious that it is difficult to understand how the error originated,
the more so because Milne Edwards, however obscure he may have been on
the subject of the epistome, is quite definite and exact on this point. ‘“ Le
front se prolonge au-dessus de l’anneau qui porte les yeux. Dans le jeune
Age, cet anneau reste 4 découyert antérieurement, et les yeux ne sont pas
logés dans des cavités orbitaires completes; mais, plus tard, la partie
inférieure du front se réunit, sur la ligne mediane, 4 une prolongement de

A STUDY IN CARCINOLOGY. 45

Parceau inférieure du second anneau, de fagon & entourer complétement le
segment oculaire quwon n’apercoit plus qu’a Vintérieur de la carapace ” (25,
vol. i. p. 250). ;

It is not necessary to push the question further, for the above quotations
show that the terms ‘‘ epistome ” and ‘“* mouth-frame ” (cadre buccal) have no
definite connotation, and being incapable of exact application, must be dis-
carded in fayour of a more precise terminology founded on an appreciation
of the morphoiogical elements entering into the composition of this region.
Tt is, however, no easy task to give such an appreciation.

A consideration of the more simple and primitive arrangements obtaining
in the Macrura must precede a discussion of the highly modified relations of
homologous parts in the Brachyura, and for this purpose Vephrops norvegicus
may be selected as an example. Fig. 19 is a frontal and fig. 20 a lateral
view of the cephalic segments of this species. In both the rostrum has been
cut through near its base, the ocular peduncles have been cut through near
their origins, and in fig. 19 the edge of the carapace has been cut away
where it overlaps the external boundary of the right antennary fossa. The
first point for consideration is, what constitutes the anterior extremity of the
cephalothorax ? Not, I think, the rostrum. Huxley (88) has given reasons
for regarding the rostrum as a forward prolongation of the third or antennary
somite, and although, following Milne Edwards, he based his identification on
the somewhat doubtful evidence afforded by the anterior head seginents of
adult Stomatopods, there is no reason to go so far afield to obtain evidence
in support of this view. In the deep-sea Penida, Benthesicymus altus and
Hemipeneus spinidorsalis, the ocular and antennulary segments are sufficiently
distinct and freely moveable on one another to afford abundant justification
for it. It is also evident from an inspection of fig. 20, that in Nephrops the
ocular segment lies in front of the base of the rostrum and has its proper
tergum in the form of a median sclerite of cartilaginous consistency, little
if at all calcified, but none the less definite in form and extent and clearly
recognizable as a distinct element in the exoskeleton of this region. The
side-walls or epimera of the ocular segment are thin and membranous, but
the sternum is a distinct triangular piece, scarcely calcified and of much the
same cartilaginous consistency as the tergum. It has been recognised by
all previous authors. Immediately behind the ocular is the easily recogniz-
able antennulary sternum, and to the right and left of it are the articular
sockets of the antennules, separated by the arthrophragms ii/iii from the
antennary sockets. The antennulary segment is completed, as has been
shown by previous authors, by the aliform calcified plates which form the
greater part of the walls of the shallow excavations in which the ocular
peduncles rest when turned outwards and represent the epimera of the
antennulary segment. I agree with Huxley (88, p. 157) in identifying a

46 DR. G. C. BOURNE ON THE RANINIDZ:

narrow transverse bar at the base of the rostrum with {the tergum of the
antennulary segment. This bar is easily seen from the inside, but does not
appear in a front view, being concealed by the base of the rostrum. Right
and left of it are two deep erescentic grooves which are the external indica-
tion of two deep aliform apodemes to which the anterior gastric muscles are
attached. The apodemes in question are known-as the procephalic lobes,
and various suggestions have been made as to their significance, but, in my
opinion, they are simply the well-developed pleural apodemes of the inter-
seoment i/ii. Thus far the interpretations I have given are hardly, if at
all, at variance with those of previous authorities, but the homologies of
the antennary and mandibular segments present greater difficulties. The
so called epistome of Nephrops, as also that of the lobster and the crayfish,
is clearly made up of two parts: an anterior broad plate the lateral regions
of which are produced right and left into broad divergent wings, giving the
whole the shape of a broad inverted VY. Behind this, fitting closely into
the angle of the V, but separated from it by a distinct groove, is a somewhat
projecting narrow curved bar the outer extremities of which end in incurved
knobs, which knobs are excavated internally to form the strophidia for articu-
tion with the inner articular processes of the mandibles. The presence of
these strophidia is of itself evidence that the bar in question is the mandibular
sternum. The transverse groove in front of it deepens in the middle line to
form a triangular pit or depression and on the inner surface is a correspond-
ing projection—in fact, an apodeme for the attachment of muscles. As
transverse apodemes are always intersegmental, this groove with its median
apodeme indicates clearly enough the boundary between the antennary and
mandibular sterna. The antennary epimeron of either side is represented by
the very narrow carved bar forming the outer boundary of the antennary
socket. This bar is united by a membranous fold to and is overlapped by
the fold of the anterior margin of the carapace forming the commencement
of the branchiostegite. Posteriorly this bar bends inwards to form the
thickened interned rim of the antero-lateral margin of the antennary
sternum, and though it is completely fused with the latter, I do not doubt
that this thickened rim represents the intersegmental arthrophragm iii/iv.
Hxternally the antennary epimeron expands and passes, without any obvious
line of demarcation, into the calcified plate marked « in fig. 18. In
Nephrops this plate is firmly fused to the outer edge of the V-shaped
antennary sternum, and is set back at an angle to it in such wise as to form
the inner and upper wall of the anterior part of the exhalant branchial canal,
but in both the lobster and the crayfish it lies more nearly in the same
plane as the antennary sternum and takes little or no part in the formation
of the branchial canal. The postero-internal margin of this plate is thickened,
produced backwards, and engages in a groove on the front surface of the

A STUDY IN CARCINOLOGY. 47

mandible, thus forming the hinge-line about which the mandible rotates.
The postero-external edge of the plate is ill-defined and passes into an
imperfectly calcified area forming the roof of the branchial canal. The
‘membrane narrows posteriorly, and arriving at the external articular pro-
cess of the mandible, again becomes calcified and thickened to form a rather
large strophidium for articulation with the mandible. After comparison
with many other species, I have satisfied myself that the whole of this area,
ealcified in front and behind, but imperfectly calcified or membranous in
the middle, represents the epimeron of the mandibular segment, which
in the Astacura is fused to the antennary epimeron, but in some primitive
Brachyura, as I shall show subsequently, is quite independent of it and
also of the antennary sternum. Relying on these interpretations which, I
venture to think, will be accepted by anyone who makes a careful compara-
tive study of this region, I shall henceforth discard the misleading and
purely topographical names in general use and speak of the sterna, epimera,
ete., of the various seements.

Before bringing the Raninidze into comparison it will be well to consider
the characters presented by some other Oxystome crabs, particularly of
the Dorippide from which, according to some authors, the Raninide are
descended. I have studied Dorippe lunata, M. Hdw., dthusa ciliatifrons,
Fax., and #Hthusina gracilipes, Thiers. Of these the last-named species
shows the most primitive and interesting features, and its mouth-frame and
antennary region are depicted in fig. 21. The basal joints of the antennules
are enormously inflated and interposed between the exposed portions of the
ocular peduncles and the rostrum. The antenne have a four-jointed peduncle
and a fairly long flagellum ; the basal joint is small but freely moveable.
The downward process of the front or rostrum articulates with the antennulary
sternum, the latter being produced right and left into wings which are
adapted to the shape of the inflated tasal joints of the antennules. In this
species the antennulary sternum is largely exposed, only its posterior margin
is overlapped by the front end of the spout-shaped antennary sternum.
The latter sclerite is well defined and the sutures uniting it to adjacent
parts can be seen clearly. It consists of a central somewhat scutiform plate
with antero-external projections of which the edges are inrolled ventrally to
form the spout-shaped Oxystomatous mouth. It is to the hinder ends of
these lateral infolded projections that the pterygostomial processes of the
carapace are united. Behind the body of the antennary sternum is the
mandibular sternum, a very slender but distinet and calcified curved bar.
Right and left, intervening between the lateral wings of the antennary
sternum and the outer ends of the mandibular sternum, are two distinctly
defined oval plates forming the roof of the anterior part of the branchial canal
and extending so far backward that they supply the articalar strophingia for

48 DR. G. C. BOURNE ON THE RANINIDZ:

the external articular processes of the mandibles. There can be little doubt
that these are the epimera of the mandibular segment, and that they are
homologous withjthe less sharply defined areas similarly identified in the
Astacura.
In Athusa ciliatifrons the arrangements are very similar to those in
thusina, but the individual sclerites are not so easily distinguished. In
this species the spout-shaped anterior end of the antennary sternum is pro-
duced so far forwards that very little of the antennulary sternum can be
seen in a surface view. The mandibles also are reduced in size and the
mandibular sternum is membranous. In Dorippe lanata the spout-like
anterior end of the antennary sternum is produced forward io such an extent
that it articulates with and overlaps the downward process of the rostrum,
tlus completely concealing the antennulary sternum. ‘This is one of the
rare instances in which union is effected between the rostral process and
the antennary sternum. In Dorippe as in Athusa the mandibular sternum
is almost wholly membranous, exhibiting only two small calcifications in the
region of the strophidia. The mandibular epimera re large and distinct,
triangular in shape, the apices of the triangle produced backwards. The
body of the antennary sternum exhibits a large central boss in front of the
mandibular sternum, which might, at first sight, be taken for a forward
median extension of the latter, but it obviously bas no connection with it.
After this survey of one of the families of the Oxystomatous crabs, the
Raninidee may be brought into comparison. Fig. 22 is a front view of
the present region of Votopus dorsipes, all the appendages being removed
with the exception of the right mandible and the left ocular peduncle.
Fig. 23 is a similar view of Ranina dentata, and figs. 24 and 25 illustrate the
same regions in Votosceles and Lyreidus. For descriptive purposes Votopus,
a less modified genus than Ranina or Lyreidus, will be taken as the example
of the family. The first thing to be observed is that the antennary sockets
are relatively large and situated behind rather than to the sides of the
antennulary sockets. In this respect Notopus more nearly resembles
Homarus than Nephrops, for in the latter the antennary sockets are dis-
placed laterally. Reference to fig. 29 shows that in Notopus, as in all
Raninidee, the antennary peduncle is tive-jointed, the basal joint being freely
moveable, short antero-posteriorly but of relatively considerable width ven-
trally, and it bears the excretory aperture on a distinct prominent tubercle.
In Notopus and in Ranilia, M. Mdw., the antennary flagellum is multi-
articulate and of considerable length, with special features which will be
referred to later. All these characters are Macruran. The antennules.
(fig. 28) with their enlarged basal segments and abbreviated few-jointed
external and internal flagella are more caneroid in character, but the basal
joints are not inflated to anything like the same extent as in the Dorippidee

A STUDY IN CARCINOLOGY, 49

and the antennary sockets lie well behind the ocular peduncles ; there are no
antennulary fossee internal to the orbits. The ocular peduncle of Ranina, as
is well known, exhibits these calcified segments bent at an angle to one
another when the peduncle is retracted into its orbit. Reference to fig. 22
shows that there are similarly three segments in the ocular peduncle of
Notopus, but the two proximal are very short, the greater part of the
peduncle being formed by the elongated distal segments which, when turned
backwards and downwards, rests in an elongated orbital excayation fringed
with hairs. There is no division of the orbit by a downward growth of the
front external to the antennules, and the proximal segments of the ocular
peduncles are inserted close to the middle line on either side of the rostral
downgrowth. In all these respects the Raninidze have departed very little
from the Astacuran type and differ from the more cancroid characters
exhibited by the Dorippidee. In Wotopus the downward process of the
rostrum is narrow and strongly keeled ventrally ; its dovetail-like articu-
lation with the antennulary sternum is well seen in fig. 22. The antennulary
sternum is escutcheon-shaped, strongly keeled in the middle line, and narrow
posteriorly. It has a certain amount of mobility on the antennary sternum,
which latter structure is a broad triangular or rather V-shaped plate the
apex pointing forward, the antero-lateral margins raised and thickened but
not infolded and forming part of the inner wall of the antennary sockets :
more posteriorly these margins are united to the pterygostomial lobes of the
carapace. The ventral surface of the antennary sternum is deeply excavated
to form the exhalant orifice of the branchial canals ; its anterior moiety is
divided into right and left channels by a low median ridge, posterior to which
the right and left branchial canals are separated from one another by a large
ventrally projecting hood. This hood, which is characteristic of the Rani-
nidee, is the greatly developed median part of the mandibular sternnm, and its
postero-external corners are produced right and left into curved horns at the
extremities of which are the strophidia for the articulation of the mandibles.
The-suture between the antennary and mandibular sternum is well marked
in all the Raninidee I have examined. ‘he shape and size of this hood-like
mandibular sternum and its relations to the antennary sternum in front and
the labrum behind are shown for Ranina in the perspective sketch (fig. 26).
The figure also shows the curved lines of hairs guarding the exit of the
branchial canal ; the posterior and more prominent line of hairs marks off
an oblong plate forming part of the roof of the branchial canal, and as a
distinct suture can be observed internally corresponding to the external line
of hairs, I identify this oblong plate with the mandibular epimeron. In
Notopus the exit of the branchial canal is guarded by a diffuse hairy
patch, and the area behind this, drawn out into the triangular projection
shown in fig. 22, must be taken to represent the mandibular epimeron,
LINN. JOURN,—ZOOLOGY, VOL. XXXV. 4

50 DR. G. C. BOURNE ON THE RANINID@:

but it is so intimately fused to the antennary sternum that no suture is
visible.

Posteriorly the mandibular epimeron is continued into the flexible mem-
brane forming the roof and outer wall of the branchial canal. In Notosceles
(fig. 24) the antennary sternum is very short, is easily separable from
the antennulary sternum, and the intersegmental arthophrragm ili/iv in
front, and the mandibular epimera appear to be wholly membranous. In
Lyreidus, on the other hand (fig. 25), in keeping with the great elongation
of the pre-oral region of the carapace, the anterior part of the antennary
sternum is greatly produced and to a certain extent overlaps, but does not
conceal, the antennulary sternum. The regions of the mandibular epimera
are also produced far backwards on either side of the mandibular sternum
and are calcified, but I cannot find any line of division between them and
the antennary sternum. It should be observed that in Lyreidus the pterygo-
stomial lobes of the carapace are produced far in front of the mandibles
and are united for a considerable distance with the edges of the antennary
sternum, a condition very different from that in the Leucosiide, with which
Lyreidus has so often been compared.

From what precedes it is evident that the Raninide diverge so much from
the Dorippidze and are so much more primitive in several important par-
ticulars, that they cannot be descended from the latter family. Nor does a
comparison of the pre-oral segments give any support to the descent of the
Raninidze from the Dromiacea. Fig. 27 is a frontal view of the antennary
and oral regions of Dromia vulgaris. The antennee and antennules have
been removed and also the left ocular peduncle, but that of the right side is
left in place. The whole facies is more cancroid than in the Raninide. The
proximal ends of the ocular peduncles, concealed behind the inflated basal
joints of the antennules when the latter are in place, are membranous. The
antennulary and antennary fossee are small and lie in nearly the same trans-
verse line. The orbits so far resemble those of the Raninide that there are
no downward processes of the front external to the antennules. The median
triangular shield with the apex directed forward is the antennular sternum,
and evidently is what Milne Edwards called the “ epistome” (loc. cit.). It
is separated by a distinct but shallow membranous apodeme from the
antennary sternum behind, and there is a certain amount of mobility between
these sterna. The apex of the antennulary sternum fits into a deep depres-
sion of the downward process of the rostrum, but can easily be pulled out of
it. The antennary sternum forms the anterior boundary of the so-called
mouth-frame, and consists of a median bilobed area and two lateral wing-
shaped areas marked off from the former by grooves. The two lobes of the
median area are nearly square in outline and separated from one another by
a deep median fissure : their anterior margins are thickened and truneated,
forming a prominent ledge behind the base of the antennulary sternum.

A STUDY IN CARCINOLOGY. 51

Each of the lateral wings is armed anteriorly with a prominent spine ; its
outer border is nearly straight and makes nearly a right angle with the
transverse ledge formed by the median lobes; its inner border is thickened
and curves round the outer extremity of the mandibular sternum. ~The last-
named structure is a projecting curved bar intimately fused to the median
area of the antennary sternum, but marked off by a groove which in the
middle line deepens to form a triangular fossa. Laterally the mandibular
sternum seems to terminate in a pair of incurved prominences for articula-
tion with the inner articular processes of the mandibles, but is clearly
prolonged beyond these as a pair of diverging horns intimately fused to the
postero-internal edges of the aliform external areas of the antennary sternum
but projecting slightly beyond the latter. The mandibular epimera are
largely membranous, but their antero-internal ends are calcified to form the
two plates marked in fig. 26 which form the roof of the anterior part of
the exhalant branchial canals. A comparison of figs. 19, 22,and 27 will, I
think, convince the reader that in the whole make-up of the pre-oral region
the Dromiacea have departed further from the Astacuran type than have the
Raninidee, particularly in the reduction of the rostrum, the greater develop-
ment of the suborbital lobe of the carapace, the membranous condition of
the inner moieties of the ocular peduncles, the size and relative positions
of the antennulary and antennary fosse. Further, the modifications of the
antennary and mandibular sterna are widely divergent in the two groups.
It may be objected, and there would be some force in the objection,
that the genus Dromia, on which I have relied for study of details, is a
much modified and specialised genus of the Dromiacea, and that I should have
directed my attention rather to the more primitive genera, Homolodromia and
Dicranodromia, on which Bouvier so largely relies in establishing his theory
of the Astacuran origin of crabs. Unfortunately examples of these rare
and instructive forms were not at my disposal, but they have been deseribed
in sufficient detail by A. Milne Edwards and Bouvier (28), and a reference
to pl. i. fig. 2 and pl. iii. fig. 2 of their admirable memoir will convince the
reader that I am justified in extending the results of my study of the genus
Dromia to the more primitive members of the group and in asserting that in
the make-up of the pre-oral region, as in other characters previously cealt
with, the Dormiacea have departed more widely from the Astacuran type
than have the Raninidee, and therefore cannot, be enrolled in the ancestry of
the latter.

I submit that, by a detailed study and comparison of the neryous system,
the endophragmal system of the thorax, and the cephalic segments, I have
proved the first part of my thesis and have shown that, whilst there is plenty
of evidence for deriving each group separately from an Astacuran ancestor,
the Raninide cannot be directly descended from a Dromiacean stock. The

4%

52 DR. G. C. BOURNE ON THE RANINIDA:

paleontological evidence available, as far as it goes, supports this thesis.
The Raninide are ancient forms among crabs, and when they make their
first appearance have their characteristics fully developed. he earliest
fossils of which I can find a sufficiently exact account are Laninella Trigert
from the grés verts of Maine (France) and 2. elongata from the cretaceous
beds of the Sarthe. Both deposits belong to the Turonian division of the
Cretaceous. In these early Raninide the carapace is of elongated oval
shape, broadest in the anterior third; the “buccal frame” is so much
elongated that it is nearly half the length of the body ; the thoracic sterna,
wide between the first pair of legs, become narrowed between the second
pair and are reduced to linear dimensions between the posterior pairs of
pereiopods ; the four pairs of pereiopods are compressed and adapted for
digging in sand (A. Milne Edwards 27). This is the description of a
very typical Raninid of fossorial habit showing no greater affinity to the
Prosoponide or to Protocarcinus (Palwinachus,-Woodw.) than do existing
members of the group. The presence of Ranina marestiana and Notopus
Beyrichi in the Hocene show that modern genera of the Raninidee were then
differentiated and well established. Bittner’s (8) excellent figure of the
latter species might pass for a representation of the anterior half of the cara-
pace of a modern Notopus. Perhaps the geological evidence proves no more
than the antiquity of the Raninidz, but that antiquity affords support to
the argument that they had an independent origin from the Astacura.

If, then, they are not descended from a Protocarcinid-Prosoponid stock by
way of the Dromiacea, a fortiori the Raninide cannot be descended from
the Dorippide or Cyclodorippidee, which are themselves derived from the
Dromiacea. From what has preceded it is evident that they cannot be;
their primitive nervous system and endophragmal skeleton is sufficient
evidence of that. On the same reasoning, the Dorippidee cannot be descended
from the Raninidee, for it has been shown that their ancestry is altogether
different. So one of the families, and that the most primitive, of the
remainder of the Oxystomata is excluded from relationship with the
Raninidee. What, then, of the two remaining Oxystome families, the
Calappide and the Leucosiide. They are classified with the Raninide
because the majority of carcinologists have found it difficult to believe that
the Oxystome “mouth frame” could have been acquired independently by
different lines of descent. But since the Dorippidee are excluded from
relationship with the Raninidé this objection no longer has any force. But
there is room for the opinion that the Calappidee and Leucosiide are
descended from the Raninidz, and I have already quoted (p. 26) de Haan’s
positive opinion that Lyreidus is an annectant form with the Leucosiide.
There are, of course, many points of difference between the Raninidee and
the two above-named families. It would be an easy though a lengthy task

A STUDY IN CARCINOLOGY. 53

to enumerate them, but the labour may be spared because the majority of
them could plausibly be explained on a theory of descent with modification
from a Raninid ancestor. This explanation could be given satisfactorily in
such matters as the concentration of the nervous system, the disappearance
of the sternal canal, the broadening of the posterior thoracic sterna, and a
large number of other characters. But it cannot apply in the case of
structural features which have been profoundly altered or have disappeared
altogether in the Raninide but are present and exhibit normal relations in
Calappidee and Leucosiidee.

To take first a feature peculiar to the Raninids, the marked reduction in
verticai depth of the posterior part of the branchiostegite whereby a con-
siderable area of the epimera of the eleventh, twelfth, and thirteenth
segments is left uncovered. This is by no means primitive but a definite
specialisation, and I have attributed it to the burrowing habits of the family.
The Calappinee are certainly and the Matutine largely sand-burrowing
crabs, butin both the lower edges of the branchiostegite fit very closely to the
coxee of the pereiopods. In the Leucosiide, which are supposed to resemble
the Raninidee more closely than other Oxystomes, the adaptation of the lower
edges of the branchiostegite to the cox of the pereiopods is particularly
close and elaborate. It cannot be argued that the original and more
primitive relations of the branchiostegite to the epimera were re-established
when the necessity for enlarged muscle-cavities for the pereiopods disap-
peared with the assumption of new habits by the Calappidee and Leucosiide,
for the muscle-cavities are very large in these families but their enlargement
is provided for in a very different manner. In the Leucosiide the arrange-
ment of the elongated abductor muscle-cavities is peculiar and interesting,
but there is no room to describe it in this place.

In all the Raninidee the posterior margin of the pterygostomial region of
the carapace is closely united to a broad offset of the tenth sternum in front
of the cheliped. Consequently there is no inhalant branchial orifice in
front of the cheliped, and the epipodite of the third maxilliped is aborted,
though a trace of it remains in the form of a vestigial setobranch. In the
Calappinee and Matutinee the pre-chelipedal inhalant aperture is conspicuous
and the epipodite of the third maxilliped well developed. It cannot seriously
be maintained that these structures have been re-acquired in these two sand-
burrowing sub-families. In the Leucosiidz, however, the posterior margin
of the pterygostome is as intimately fused with the plastron as in the
Raninidee, and with the disappearance of the pre-chelipedal branchial orifice
the epipodite of the third maxilliped has disappeared even more completely
than in the Raninidee, for there is not even a vestigial setobranch. A similar
state of things occurs in some of the Cyclodorippidee, but need not be
discussed here because the Dorippidee have already been excluded from

D4 DR. G. C. BOURNE ON THE RANINIDA :

relationship with the Raninidze. De Haan laid great stress on this point of
resemblance between the Leucosiidze and the Raninide, but a detail exnmina-
tion shows that the result is arrived at in a different manner in the two
families and must be regarded as a case of homoplasy rather than homogeny.
In all the Raninidee the free end of the sternal plastron is formed by the
more or less prominent lanceolate median projection of the tenth (chelipedal)
sternum. The ninth sternum is narrow and projects upwards at an angle
of 130° to 135° above the lanceolate projection of the tenth sternum. The
coxee of the third maxillipeds are articulated to the hinder end*of the narrow
ninth sternum, and therefore close to the median line, and they effectively
exclude any part of the sternum from participation in the formation of the
broad plastron (see fig. 27,ix). Further, in the Raninidz, the connection
between branchiostegite and plastron in front of the cheliped is effected by
lateral outgrowths of the tenth sternum which meet the branchiostegal
margin, not by any marked ingrowths of the latter. In the Leucostide
(1 have studied the arrangement in Jha nucleus, Plilyaia undecimspinosa,
and Philyra levis) the ninth sternum is short antero-posteriorly, but broad
and forms the anterior end of the plastron. The coxee of the third maxill-
peds are articulated to its outer ends and are therefore far apart. The tenth
sternum does not give off any prominent outgrowths in front of the chelipeds,
but is united to an inwardly directed process of the edge of the branchio-
stegite on each side, this process being the most anterior of a series of
similar downgrowths of the branchiostegal margin which pass between the
coxie of the pereipods and effect the very close and elaborate fit of branchio-
stegite to coxee already referred to. It may be mentioned here that
A. Milne Edwards and Bouvier (28) have described a shallow pterygostomial
gutter closed in below by the third maxillipeds in Cyclodorippe. This
gutter occupies the same position as the well-known pterygostomial inhalant
canal of the Leucosiidee, and the similarity of the structural appointments is
so great as to suggest the derivation of the Leucosiidee from a Cyclodorippid
rather than from a Raninid ancestor.

Finally some importance is to be attached to an apparently minor feature.
In all the Oxystomatous Crabs the floor of the exhalant branchial canal is
formed by an expanded spoon-shaped operculum furnished by the first
maxilliped. In the Raninidz (see figs. 31, 45, 50, & 53) both the exopodite
and endopodite are expanded and modified, and co-operate in forming the
operculiform floor in question, but it is the exopodite which is the longest,
most expanded, and the most effective agent in forming the operculiform
floor of the anterior part of the exhalant canal. This modified exopodite
never bears a flagellum. In the lLeucosiide, the Calappinee, and the
Matutinze (Garstang is in error in stating that the exopodite is operculiform
in Matuta banksit) it is the endopodite of the tirst maxilliped that is elongated.

A STUDY IN CARCINOLOGY. BY)

expanded, and modified to form the operculiform floor of the exhalant canal.
The exopodite is somewhat modified and to a varying extent in different
genera, but it is always shorter than the endopodite, does not share in the
formation of the opereulum, on the whole retains the characteristic shape of
an exopodite, and invariably bears a flagellum—I hope I may not be criticised
for attaching undue importance to a trifling character. It is just such
instances as this, in which the same purpose is effected by somewhat different
means, that afford the best criteria as to whether some particular structural
resemblance is homogenetic or homoplastic. It was the discovery of this
difference in the structure of the first maxillipeds that finaliy confirmed my
opinion that the Raninidz are not genetically connected with the other
Oxystomatous Grabs. My reliance on thiss item of evidence was somewhat
shaken by A. Milne Hdwards and Bouvier’s statement that the opercular
floor of the exhalant canals in Cymonomus is formed by the dilated exopodites
of the first maxilliped, but an examination of their figure (28, pl. xv. fig. 6—
there is a mistake in the numbering of these figures—) shows that the
resemblance to the Raninidee is only superficial, and the exopodite of the first
maxilliped of Cymonomus has a long six-jointed flagellum furnished with long
sete. My test case, therefore, holds good, and the evidence produced is
sufficient to justify Boas’ doubt as to the inclusion of the Raninidse among
the other Oxystomata, and A. Milne Edwards and Bouvier’s decision to
exclude them from their monograph on the Oxystomata of the ‘ Blake’ and
‘Hassler’ Expeditions. The Raninidee, therefore, must be removed from
their present systematic position and assigned to some other place in classi-
fication. They cannot be excluded from the section Brachyura for reasons
given on p. 27, but in conformity with more recent and generally accepted
classifications they must rankasa separate tribe, equivalent to the Dromiacea,
Brachyenatha, and the rest of the Oxystomata. For this new tribe I propose
the name Gymnopleura*, which directs attention to one of the most charac-
teristic features of the group, namely the exposure of the epimera of the
posterior thoracic segments. ‘This new tribe may be defined as follows.

Tribe GYMNOPLEURA.
Anterior thoracic sterna broad, posterior thoracic sterna narrow and keel-
like ; posterior thoracic epimera largely exposed by reduction of branchio-
stegite ; female openings on coxe ; last pair of pereiopods dorsal in position,

* Mr. 1. R. R. Stebbing in ‘ Nature,’ Jany. 1922, has pointed out that Latreille (Cours
d’Entomologie, 1881) instituted the tribe Notopterygia for Raina. At first sight it seemed
advisable to adopt Latreille’s name for the tribe comprising the Raninide instead of my
own. But the rules of nomenclature do not apply to tribal names; Latreille’s system of
classification differs largely from that followed in this paper and his reasons for placing
Ranina in a separate tribe differ widely from mine. Confusion rather than perspicuity
would result from the adoption of Latreille’s tribal name.

56 DR. G. GC. BOURNE ON THE RANINIDA:

normal or reduced in size; sternal canal present; thoracic nerve ganglion-
chain elongate ; antennary sternum triangular, spout-shaped ; branchiz 8 on
CACM BCI, cocacvcs coosasoac Faminy: Raninide.

We may now deal with some features of general and special interest
presented by the tribe Gymnopleura. It has been shown to have originated,
independently of other ‘‘crabs,” from the Astacura. It is but a small
assemblage of animals, comprising some eight or nine living genera,
exhibiting a very small range of variation, and evidently very well adapted
to their environment, for the tribe emerges in the Cretaceous with its special
adaptive characters fully established, and two of the recent genera date back
to the Eocene. But although not descended from crabs, but from lobsters,
the Raninidee have been so similarly modified, in one direction and another,
that they must be classed with the crabs with which they have no relation-
ship save that of a remote common ancestor which was not itself a crab.
It is by no means an isolated phenomenon. We know of several crab-like
forms, Porcellana, Lithodes, Hippa, and the interesting Porcellanopagurus,
whose assumption of a crab-like form has been described in a very lively
manner by Borrodaile (20), which have undoubtedly been derived, quite
independently in each individual case, from Macrurous ancestors. The
question is, what causes have operated to produce so great similarities in
animals so remotely related to one another? Without doubt, I think, the
answer is that the efficient cause has been the assumption of the habit of
burrowing in sand or mud. Bohn (7) has shown that various members of
the Nephropsidea, when living on sandy bottoms, protect themselves by an
investment of sand, and to a limited extent take cover in it. But their
pereiopods are ill-adapted for digging; they must retreat backwards into the
sand to leave their antennze and eyes free to keep watch against enemies,
and the elongated abdomen is a great obstacle to the effective and rapid
concealment of the posterior parts of their bodies. A suitable modification
of the pereiopods and reduction and infolding of the obstructive abdomen
are prerequisite to retrofossorial efficiency, and any mutations in these
directions must have had a high selective value. It is not surprising that
favourable modifications should have presented themselves and have been
selected several times over, nor is it surprising that, once the burrowing
habit was adopted, similar adaptive modifications to the new condition of life
should have established themselves. In the first place the Decapod, buried
in sand or mud, must adapt its respiratory mechanism to the changed
conditions. The inhalant spaces extending along the posterior and ventral
edges of the branchiostegite would be choked and rendered useless unless
some provision against the entrance of sand were developed, and the
researches of Garstang and Bohn have shown by what various means this
form of suffocation is obviated in different genera of the sand-burrowing

A STUDY IN CARCINOLOGY. oN

crabs. Most commonly the edges of the branchiostegite are titted closely to
the epimera just above the cox of the pereiopods and to the tergum of the
first abdominal segment, so that no water ean find ingress or egress that way.
As a consequence the podobranchs or arthrobranchs of the three posterior
pereiopods disappear ; there is no longer room for them, and as the branchial
cavity is reduced posteriorly by the great development of the muscle-cavities
of the digging legs, the posterior pleurobranchs follow suit. The branchiz
are reduced in number and those that remain are massed in the anterior half
of the branchial cavity, nearest to the newly developed entrance of the
respiratory current in front of the chelipeds. In the Raninide, however,
the arrangements differ from those usually observed in crabs. There is, in
most of them, a pair of posterior respiratory orifices situated between the
tergum of the first abdominal segment and the coxz of the last pair of
pereipods. When the abdomen is extended or only slightly flexed, water
can pass freely into or out of these orifices, but they are pretty effectively
closed when the abdomen is closely flexed under the thorax. The orifices
were accurately described and figured in Ranma by Milne Edwards, but
many subsequent authors, particularly Ortmann (42), seem to have failed to
recognise them. Borrodaile (14) gives an accurate account of their relations.
These posterior branchial orifices are not, however, peculiar to the Raninidee :
I have found them in precisely the same position in Corystes, in which genus
they can easily be seen by bending the abdomen upwards and looking between
the lower side of the tergum of the first abdominal segment and the coxa of
the last pair of legs. Garstang (30) failed to observe these apertures in
Corystes, and I suspect, but have not yet had the opportunity of proving it
by experiment, that they form the main entrance for water into the branchial
chamber during the operation of the normal current. ‘There is also a pair of
posterior apertures in Zhia polita, but these are in a somewhat different
position, close under the edge of the branchiostegite in front and to the out-
side of the articulation of the first abdominal segment withthe carapace, and
well in front of the coxee of the last pair of pereiopods. The persistence of a
posterior branchial opening is a primitive feature, never to be seen, as far as
my observations go, in crabs in which the abdomen is permanently flexed
and kept closely applied to the sternal plastron.

To return to the Raninide: the posterior part of the branchial chamber,
into which the posterior orifice opens, is reduced to a narrow passage by the
reduction in vertical depth of the branchiostegite and the close adherence of
its edge to the thoracic epimera. The edge of the branchiostegite is held in
place by two prominences, one on the anterior edge of the eleventh, the
other on the posterior edge of the thirteenth epimeron, and a close fit is
ensured by a well-marked ridge running upwards and backwards along the
eleventh, twelfth, and thirteenth epimera (figs. 7 & 17). Thus the entrance
of water into or its egress from the branchial chamber at the sides of the

58 DR. G. C. BOURNE ON THE RANINIDA:

thorax is effectively prevented. ‘The only entrances and exits are by way of
the anterior so-called exhalant branchial canals and by the posterior canals
described above. Bohn (6) has shown that reversal of the respiratory current,
first observed in Corystes by Garstang, is a normal phenomenon among
Decapod Crustacea, and is manifested even when they are buried in the sand.
There must, therefore, be some apparatus in forms which, like the Raninidee,
have posterior branchial orifices for filtering the sand from the water entering
by these orifices during the “normal” phase of the respiratory current ;
that is when it is setting from behind forwards. In most of the Raninide,
notably in Ranina, Notopus, Notopoides, Notosceles, and Zanclifer, this
apparatus is furnished, partly by the chelipeds, but principally by the manner
in which the flattened pereiopods, the edges of which are fringed with long
and closely set hairs, are bent upwards and forwards in such a manner as to
form a water-chamber on either side of the posterior thoracic segments.
The part played by the pereiopods would never be guessed from the con-
ventional representations of these animals in systematic works, where they
are depicted, usually from the dorsal surface, with the legs extended sym-
metrically on either side of the body in order to display as much as possible
of their structure and the chelipeds extended forwards in front of the
body. The adaptive characters of the thoracic limbs can be studied with
equal advantage in Notopus, Notopoides, Zanclifer, Notosceles, and Ranina, but
I will take the last-named genus as an example for descriptive purposes.

In Ranina there is a conspicuous triangular patch of short dense hairs
extending forward from the articulation of the cheliped over the postero-
lateral area of the pterygostomial region on either side of the thorax. The
conjoined basis and ischium of the cheliped is very short and immoyeably
fused to the merus, the two forming a relatively long curved segment of the
limb, dilated on its external aspect but smooth and flattened internally so as
to fit closely against the hairy patch on the pterygostome. It is evident,
from its smooth and polished inner surface, that the ischiomerus is normally
held close to the body and slides forwards and inwards or outwards and
backwards over the above-mentioned hairy patch. Whatever its position, so
long as it is pressed against the hairy patch, there is no room for the passage
of water, much less of sand or mud, between it and the pterygostome.
When the ischiomerus is rotated as far forward as possible, the somewhat
inflated carpus lies beneath the outermost of the large frontal spines of
the carapace, and the flattened propodus and dactylus are folded back
under the anterior part of the pterygostomial region, but do not fit closely
and accurately to the latter as in the case of Calappa and Matuta, Their
function is quite different. It is evident that instead of forming the floor of
what Garstang has called an exostegal canal the propodus is so articulated
to the carpus that without either the latter or the ischiomerus being shifted
from their positions, it can be rotated outwards in such a manner as to rake

A STUDY IN CARCINOLOGY. 59

sand or other material from beneath the anterior part of the carapace, the
raking action being facilitated by the five large spines on the lower margin
of the propodus. Thus a more or less clear water-way is kept on the under
side of the anterior side of the thorax.

When the ischiomerus is drawn as far back as possible, the carpus can be
flexed inwards to form an acute angle with it, and the flattened propodus may
then be folded back so far that its posterior margin fits into a groove in the
merus and both it and the dactylus are pressed against the basal joints of
the third maxillipeds. In this position also the propodus can be rotated
outwards with a raking action so as to clear away sand from beneath the
body, without any corresponding moyement of the ischiomerus and carpus.
A close examination of the chelipeds leaves no doubt that their main function
is to keep open a passage for water down to their basal joints. From this
point backwards water must pass in two streams to the posterior branchial
orifices along passages roofed in, partly by the overhanging edges of the
branchioslegite, partly by the forwardly directed last pair of pereiopods.
The inner walls of these passages are formed by the epimera of the eleventh,
twelfth, and thirteenth segments, which are somewhat excavated and over-
hung by the edge of the branchiostegites. Their outer walls are formed hy
the first three pairs of pereiopods, which are tucked up under the roof formed
by the last pereiopods, the merus in each case being directed forwards, the
carpus, propodus, and broad lanceolate dactylus downwards and backwards.
The segments of the pereiopods being broad and flattened and their margins
furnished with fringes of stiff closely set hairs, they form a sort of basket-
work or sieve the meshes of which are filled in by the fringes of hairs, and the
whole apparatus is adapted to admit to the interior water from which sand
and other solid particles have been strained. The strainer is made more
effective by the dense hairy fringes on the edges of the branchiostegite, and
the whole apparatus is completed and roofed in posteriorly by the flattened
segments of the abdomen, which are likewise provided with dense fringes of
stiff hairs.

Whatever may be the position of the chelipeds, there is a ventral gap on
either side between their basal joints and the first pair of the pereiopods.
This gap is partly filled by the ‘epaulettes” projecting from the sterna of
the eleventh segment ; the rest of the gap is bridged over by the stiff hairs
projecting from all round the margins of the epaulettes and forming a very
effective strainer. It is through these two gaps furnished with this efficient
straining apparatus that the greater part of the respiratory current of water
must find its way from the cavity raked out in the sand by the chelipeds
underneath the anterior part of the thorax.

The structure and arrangement of the pereiopods and their relations
to the thoracic epimera and abdomen being as above deseribed, it may be
inferred that Ranina buries itself in the sand by the digging action of the

60 DR. G. C. BOURNE ON THE RANINIDA:

first three pairs of pereiopods, the fourth pair being used to shovel the dis-
placed sand outwards. When dug in, the animal may be inferred to assume
an oblique position, the frontal spines of the carapace just breaking the
surface of the sand; the chelipeds are folded up under the pterygostomial
region of the carapace and the long three-jointed eye-stalks may be extended
well above the sand on the look-out of, on the approach of danger, may
be folded back and concealed in the orbital cavities. In this position the
normal respiratory current will be provided for by the raking action of
the chelipeds and the lateral water-passages enclosed between the
pereiopods and the thoracie epimera, as described above. These afford a
mechanism whereby filtered water is supplied to the posterior branched
openings, and the normal exhalant current will pass out by the narrow
anterior passage bounded below by the merus of the third maxilliped and
laterally by the flattened basal joints of the second antenne.

As the Raninide are tropical and sub-tropical crabs living at considerable
depths it has not been possible for me to observe the habits of the living
animals, much less to make experiments on their respiratory mechanisms.
But in the absence of direct evidence I consider myself justified in giving
the foregoing account of the course of the normal respiratory current, the
more so because an examination of numerous specimens of Wotopoides latus
and of several examples of Zanclifer caribensis in the British Museum of
Natural History gives the clearest evidence that these species are sand
burrowers, and that an inhalant current passes into their branchial chambers
through the posterior branchial orifices, which are exceptionally large and
conspicuous in these cases. In nearly all the individuals of these two species
the conspicuous hairy fringes of the pereiopods, of the abdominal pleura, of
the edges of the branchiostegite, and of the epaulettes of the eleventh sterna,
are more or less heavily clogged with sand, a fact which bears witness to
their function as a filtering apparatus. On the other hand, the water-
passages lying between the posterior thoracic epimera and the pereiopods,
and therefore guarded by these hairy fringes, are remarkably clean and free
from sand. The tiltration, however, has not been perfect, for the walls of
the posterior branchial passages are encrusted with fine particles of sand,
the distribution of which leaves no doubt that the current which deposited
them set in from behind forwards. The evidence in these two species is
remarkably clear, and the inferences drawn from it may be extended to
Notopus dorsipes and to Notosceles chimmonis, though, in the last-named
species, in which the last pair of pereiopods are greatly reduced in size, there
is evidence that the posterior inhalant respiratory current is becoming of less
importance and that the incurrent supply of water to the branchial chamber
is chiefly provided for by special modifications of the antennary region.

In the absence of opportunities for observation of and experiment on living

A STUDY IN CARCINOLOGY. 61

animals, the exact course of the anterior inhalant-respiratory current in the
Raninidze mast remain largely a matter of conjecture. Nevertheless I have
not hesitated to put forward the following interpretation based upon a
detailed study of anatomical facts in the hope that, whilst my conclusions must
necessarily be subject to revision and correction, naturalists who have oppor-
tunities of studying the live animals may be so far interested as to devote
some time and trouble to the elucidation of the workings of their respiratory
mechanisms. ‘The anatomical characters are most easily studied in Notopus
dorsipes, and it is probable that what is true of this species is also true of
Ranilia muricata, M. Edw., but as I have only been able to examine a single
dried example of the latter in the British Museum of Natural History,
I cannot say anything certain about it. I have stated elsewhere (p. 26)
that, in my opinion, Fanilia, M. Edw.= Notopus, de Haan. In Notopus the
antennee are more primitive, that is to say, they depart less from the
Macruran pattern, than in any other Raninidee. The peduncle (figs. 29 & 29.)
is made up of five distinct segments cf which the proximal is admedian in
position, small, largely concealed by the sub-antennary lobe of the pterygo-
stome and bears at its inner angle the aperture of the excretory gland. The
second segment is also small and largely concealed by the sub-antennary
lobe: it lies on the same level as the first and is external toit. The third
segment is large, sub-quadrangular in outline, its extero-ventral surface
slightly convex and tuberculated, its intero-dorsal surface smooth and slightly
concave. Its admedian margin is prominent and curved, forming a pro-
jection which meets its fellow in the middle line. It bears a conspicuous
tuft of long forwardly directed setee. Its extero-anterior angle is produced
into a process which embraces the proximal part of the fourth peduncular
segment; this process is largely developed in other Raninidze but remains
small in Notopus. The fourth and fifth segments are fairly long, their
external and ventral surfaces granular and convex, their admedian surfaces
smooth and flat or slightly concave. ‘The dorsal and ventral edges of these
smooth admedian surfaces are fringed with long closely set plumose sete
which, when the two antenne are approximated, interlock with the corre-
sponding setee of the antenna of the other side. The antennary flagellum
comprises twenty-six joints, is relatively longer than in any other Raninidee
(except Ranilia, M.-Edw. and Cosmonotus) and bears a dorsal and a ventral
row of strong setee which are directed ohliquely inwards so as to interlock
with those of the opposite side when the antennary flagella are approximated.
Thus there is formed an antennary water-tube (figs. 1 & 56), shorter indeed
but otherwise similar to and fully efficient as that of Corystes. Water sucked
down the antennary tube passes into a shallow chamber lying behind the bases
of the antenne, the floor of which is formed by the meri of the third maxilli-
peds, its roof by the spoon-shaped expanded anterior ends of the exopods of

62 DR. G. C. BOURNE ON THE RANINID@:

the first maxillipeds. These last are concave dorsally and convex ventrally,
and fit so closely and accurately to the converging exhalant channels of the
antennary sternum as to constitute an efficient horizontal partition between
an exhalant passage above and an inhalant passage below. The incurrent
stream of water, taking the more ventral course, must pass right and left
‘along the conduits provided by the grooves on the ventral sides of the exopods
of the second maxillipeds; thence under the edge of the inflated pterygostome
into the channel which lies parallel to that in which the scaphognathite
works, but separated from the latter by the vertical partition formed by the
mastigobranch of the first maxilliped. The floor of these inhalant passages
is formed by the flat and closely opposed meri and ischia of the third
maxillipeds the outer edges of which fit close to the mouth frame. There is
no “exostegal” groove in the pterygostome forming a definite inhalant canal
as in the Leucosiide, but the relation of the incurrent to the excurrent stream
of water must otherwise be much the same in the two groups. That is to
say, in both cases there must be two opposite currents, the one setting
inwards to, the other setting outwards from the antennary region of the front.

In both the Raninidee and the Leucosiidee the excurrent stream is set in
motion by the “normal” action of the scaphognathite, and passes forward
between the converging channels of the antennary sternum and the exopods
of the first maxillipeds to emerge between the basal joints of the antennules.
The course of the incurrent stream in the Raninide is as described above for
Notopus, but it can hardly have escaped the reader’s attention that the
channels in which the opposite currents flow are not very distinctly separated
from one another. In the Leucosiide the existence of the exostegal canals
makes the separation much more complete, and the respiratory mechanism is
in this respect more highly specialised than it is in the Raninide. In
this connection it should not be forgotten that the latter have additional
apertures of ingress to the branchial chambers, viz., the posterior branchial
apertures of which there is no counterpart in the Leucosiide.

It may be surmised that Notopus makes use of the posterior branchial
apertures only when it is lightly covered by the loose upper stratum of sand,
and can make use of the raking action of the chelipeds to clear a passage
for water below the thorax. As it digs down into the more compact deeper
layers of sand, this source of water-supply must become more and more
restricted, and the antennary tube is then brought into action. It is of
course possible and even probable that, when the antennary tube is used,
the action of the scaphognathite is reversed and the posterior branchial
apertures become exhalani, the frontal exhalant passages being closed and
thrown out of action so long as the respiratory current is reversed, only to be
brought into use again when the normal respiratory current, from behind
forwards, is restored. But the evidence is strongly in favour of my contention

A STUDY IN CARCINOLOGY. 63

that in the Raninide an anterior incurrent stream of water is produced by
the “normal” action of the seaphognathite.

In no other member of the group (with the possible exception of Cosmo-
notus, which I have not had the opportunity of studying sufficiently closely)
do the flagella of the antennee form a water-tube as in Notopus and Ranilia.
Indeed, the antennee are modified in other directions but, none the less, are
subservient to the establishment of an inhalant water current, the course of
which can best be described by taking anina as an example.

In this genus both the antennee and the antennules are modified in con-
nection with the respiratory currents. The antenna (figs. 35 & 35 a) is short
and thick, the flagellum reduced to a rudiment comprising at the most seven
joints ; often it is missing. The two basal joints of the peduncle are fused
together, but the suture between them remains distinct. The first joint is
triangular in outline, with the apex directed forward ; its admedian basal
angle projects inwards and slightly forwards and the excretory aperture is
placed somewhat to its dorsal side, in such a’ position that the excretory
products are discharged into the excurrent rather then into the incurrent

‘stream. The two basal joints, which have a very limited movement in the
antennary socket, lie directly in front of the sub-antennary lobe of the
pterygostome, but the third peduncular segment is flexed inwards to form a
somewhat acute angle with them. The shape of the third segment can best
be understood by reference to figs. 35 and 35a, which represent external and
admedian views of the right antenna. The most noticeable feature is the
great development of the external lobe. Small and inconspicuous in Votopus,
this lobe in Ranina forms a large sinuous ridge which projects forward as far
as the anterior limit of the fourth segment; its outer margin is fringed with
long plumose setee. When the third peduncular segment is flexed towards
the middle line, the lobe in question is brought into a position athwart the
incomplete lower boundary of the orbital cavity, and. forms its ventral and
internal wall, the long fringe of hairs on what is now its anterior margin
extending upwards across the orbital cavity and interlocking with a similar
fringe spreading downwards from the supra-orbital ridge of the front of the
carapace. Tor brevity’s sake this lobe will be referred to as the crest of the
third peduncular segment. The fourth segment is sub-triangular in section :
its external and ventral surfaces are convex and granular, but its internal
surface is smooth, slightly concave, and flabelliform in outline. Its margins
are surrounded with long plumose sete the disposition of which is shown in
fig. 35a. The fifth peduncular segment is small and the flagellum rudi-
mentary or absent. When the antennz are flexed inwards, the concave
flabelliform surfaces of the fourth segments are brouglt nearly into contact
in the middle line, their marginal sete interlock, and thus a narrow but deep
space is enclosed which is continued posteriorly into the wider space bounded

64 DR. G. C. BOURNE ON THE RANINIDA:

laterally by the smooth and somewhat concaye inner surfaces of the third
peduncular segment; ventrally by the meri of the third maxillipeds; dorsally
by the densely setose expods of the first maxillipeds. This may be called the
inhalant chamber.

The antennules are inserted to the inside of and somewhat above the
antennee, As shown in figs. 34 and 34a, their basal joints are expanded
distally into a flabelliform lobe very similar to that of the fourth peduncular
segment of the antenne and, like the latter, their margins are garnished
with a fringe of long sete. The slightly concave inner surfaces of these
flabelliform expansions of the antennules, when opposed in the middle line,
form a second vertical cleft or passage within the larger passage formed by
the antennee, and this internal and somewhat dorsal antennulary passage is
so disposed that its hinder opening coincides with the spout formed by the
convergence of the exhalant canals of the antennary sternum. The cleft
between the basal joints of the antennules therefore serves as a conduit for
the excurrent stream of water. Thus far the apparatus for directing the
courses of the excurrent and incurrent respiratory streams differs from that
of Notopus chiefly in the fact that in Ranina the long tube formed by the
antennary flagella is absent, but in the latter genus a large part of the
incurrent stream must find its way into the above-mentioned inhalant
chamber through the orbital cavities. I have already shown how these
narrow and elongated cavities are covered in by fringes of setee which meet
over the retracted eyes and eye-stalks but are pushed aside when the latter
are extended. Though the orbit appears to be closed below by the crest of
the third segment of the antennary peduncle, a sufficiently wide space is left
between this segment and the basal joint of the antennule to allow of the
free passage of water from the orbit into the inhalant chamber, and much of
the inhalant current must pass through the orbits. It is evident that when
the animal is deeply dug in, the broad front of the carapace just breaks the
surface of the sand; the tips of the largest frontal spines perhaps project
very slightly above it. The hairy fringes of the orbits and of the fourth
joints of the antennze are then just awash between sand and water, and are
admirably adapted for filtering solid particles from the streams setting
inwards through the orbits and between the flabelliform fourth segments of
the antennee. Water is also admitted into the orbits on either side through
a considerable cleft lying between the long and narrow sub-orbital lobe of
the pterygostome and the innermost of the large frontal spines. These clefts
are guarded by dense hairy fringes. The arrangements by which the
incurrent stream is directed from the inhalant chamber into the branchial
chamber by the channel formed by the exopod of the second maxilliped are
the same in Ranina as in Notopus.

IT would not have presumed to give an account unsupported by direct

A STUDY IN CARCINOLOGY. 65

observation and experiment of such complicated functions as those dealt with
in the foregoing paragraphs if there were not very clear evidence that such
an anterior inhalant current as I have described does in fact exist. One
such piece of evidence is afforded by the matted feltwork of hairs covering
the ventral surfaces of the expanded anterior extremities of the exopods of
the first maxillipeds. These hairs lie flat and point backwards : they are as
evidently laid back by a current of water passing over them as the sea-grass
on the mud-banks of an estuary is laid back by the tide that has swept over
it. Again, in most examples of Ranina a considerable amount of sand is
entangled in the setee with which both the exopods and endopods of the
second maxillipeds are abundantly furnished. This gives evidence that these
appendages are bathed by a current of water not wholly freed from sand-
particles by the filtering apparatus of the orbits and antenne. But perhaps
the clearest evidence of all was furnished by the large Ranina sent me by
Prof. Kishinoye. In this specimen the inner surfaces of the third pedun-
cular segments of the antenie, the ventral surfaces of the exopods of the
first and second maxillipeds, the inner surfaces of the mastigobranchs of the
first maxillipeds, the proximal part of the mastigobranchs of the second
maxillipeds, and the bases of the podobranchs of the second and third maxilli-
peds were infested by numerous small pedunculate cirrhipedes belonging
to some as yet undetermined genus and species of the family Lepadide.
In other words, these intruders had established themselves along what I
have described as the course of the incurrent respiratory stream, but none
were to be found in the upper part of the gill-chamber, nor on the scapho-
gnathite, nor on any part of the walls of the exhalant canals. It is evident
that their larvee had been swept by the incurrent stream into the inhalant
chamber, and had subsequently fixed themselves in positions where the adults
could obtain a constant supply of nourishment borne by the incoming current.
But, where the currents were setting outwards, conditions were unfayour-
able and none had established themselves.

From what precedes, it follows that Ranina, no less than Notopus, is highly
adaptable in respect of its respiratory arrangements, and can make use
of different mechanisms as the circumstances of the moment may require.
When half buried in loose sand or when wandering on the surface these
animals probably respire through the apertures at the posterior end of the
thorax. When deeply buried in compact sand they are forced to rely on the
antennary apparatus. In point of efficiency and specialisation their
respiratory mechanisms fall far short of those exhibited by other “ Oxysto-
matous” crabs, the Leucosiids, the Calappine, and the Matutinze, but
adaptability to varying conditions implies a high survival value, and it is to
be remarked that Notopus and Ranina are among the oldest of the fossil
Raninide.

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 5

66 DR. G. C. BOURNE ON THE RANINID A:

Among other genera, Zanclifer and Notopoides, though their antennary
flagella are not so much reduced, are in all essential respects so similar to
Ranina that it can hardly be doubted that they have similar habits and are
equally adaptable as regards their respiratory processes. Zanclifer differs
from all other Raninide in having normal, 7.e. not feathered and differenti-—
ated, chelipeds, and it may be that it is more nearly related to the ancestral
form than Notopus. It has all the external characteristics of a burrowing
crab. Notopoides, on the other hand, is much more closely related to
Notopus, from which it differs mainly in the elongation of the twelfth
sternum, whereby the second, third, ard fourth pereiopods are shifted back-
wards and form a posterior group of legs, separated by a space from the first
pereiopods: this feature is exhibited to a more marked degree in otosceles
(fig. 3) and is carried to an extreme in Raninoides (fig. 6). In each genus
the increase in length of the twelfth sternum is correlated with a reduction
in size of the last pair of pereiopods, a progressive narrowing of the abdomen,
a reduction in the size and functional importance of the posterior branchial
orifices, and progressive modifications of the second and third pereiopods
indicative of their increasing use as swimming organs. A comparison of the
series Ranina, Notopus, Notopoides, Notosceles, Raninoides, leads me to the
conclusion that, whilst all of them are to a greater or less extent sand-
burrowers, the first-named during its periods of activity progresses mainly
by crawling and is a poor swimmer. Notopus and Notopoides crawl and
swim; Votosceles is a fairly efficient and Raninoides a very efficient
swimmer. This conclusion I will now endeavour to justify.

Notopoides is obviously a burrowing crab ; the sandy condition of the
specimens in the Natural History Museum bears testimony to this habit.
In most of the structural features subservient to the respiratory processes
Notopoides bears a strong resemblance to Notopus. Attention may be
directed specially to the following points : the last pair of pereiopods are not
much reduced in size and all their joints are densely fringed with hairs ; the:
tergum of the first abdominal somite is as broad as the posterior margin of
the carapace ; the posterior branchial orifices are patent and give evidence ©
of their function as inhalant passages ; the exposed portions of the epimera
of the eleventh, twelfth, and thirteenth somites are concave and overhung by
the edge of the branchiostegite ; the chelipeds are adapted for raking sand
from under the thorax. In short, all the adaptations described in Notopus
and Ranina for maintaining a respiratery current through the posterior
respiratory orifices are present to an equal degree in Notopoides. This
genus must be equally addicted to the burrowing habit, but the articulations
and characters’ of its last three pairs of pereiopods suggest that, when active,
it is to some considerable extent a swimmer. :

In Notosceles the adaptations for maintaining a_ posterior inhalant
respiratory current are still recognizable, but are obviously less efficient.

tied

A STUDY IN CARCINOLOGY. 67

The last pair of pereiopods, though much reduced in size, is sufficiently
liberally provided with hairy fringes to form an effective covering for the
roof of the water-chambers enclosed between the other pereiopods and the
thoracic flanks. The epimera forming these flanks, however, are not so
deeply concave as in Notopoides, and the posterior branchial orifices, though
distinct enough, are smaller and obviously of less functional importance.
The dactylus of the last pair of pereiopods is small, elliptical, and clearly
better adapted for natation than for digging.

Before entering into a discussion of the transition from digging to
swimming limbs it should be premised that in the pereiopods of all the
Raninidze the articulation of the dactylus to the propodus is singularly like
that of the tarsi of the whirligig-beetle Gyrinus, familiar to all naturalists
from the illuminating description of Miall. Generally, in the Raninidz, the
carpus is moderately long, triangular in shape, with an extensive basal
articulation for the propodus. The propodus is a short and broad oblong,
more or less flattened, with a very small articulation for the dactylus close
behind its postero-external angle. The dactylus is very variable in shape.
Generally in the first two pereiopods it is lanceolate and obviously efficient
in digging, but in the last two pereiopods something between crescentic and
cleaver-shaped like an oriental “kukri.” In either case it is so articulated
by a narrow pedicle to one end of the oblong propodus that it can be slid
behind the latter as the sticks of a lady’s fan slide over one another. A
very similar form of limb with yet more elabofate adaptive details is seen in
Matuta which, like the Raninidze, is at once a swimming and a burrowing
erab. It is presumed that the lanceolate form of dactylus is more uselul as
a pick, the kukri-shaped dactylus as a shovel, but the latter may be service-
able in swimming, just as a shovel may on emergency be used as a paddle.
In Zanelifer the dactyli of all the pereiopods are narrow and sickle-shaped ;
they seem ill adapted for swimming but effective instruments for digging.
The criterion for deciding whether a limb is utilised for swimming or for
digging consists in the length and mobility of its joints. The short strong
limbs of Ranina showing restricted mobility at the joints are clearly fossorial
and ill adapted for natation : the somewhat longer and more mobile limbs of
Notopus and Notopoides are, however, serviceable for both purposes. In
Notosceles (figs. 2 & 3) the dactylus of the second pereiopods is sickle-shaped
as in Zanclifer, and it may be inferred that this pair of limbs is specialised
for digging, but it can hardly be doubted that the broad kukri-shaped
dactylus of the third pair, hinged fanwise to the outer angle of the trans-
versely elongated propodus and provided with a marginal fringe of long sete,
is specially adapted for swimming. The reduced fourth pereiopods of this
genus, with their small flat elliptical dactyli, can hardly be of any value in
digging and are rather suggestive of steering paddles.

5*

for)
[o2)

DR. G@. C. BOURNE ON THE RANINID# :

Though Notosceles is in several respects intermediate between Notopoides
and Raninoides, the last-named genus (figs. 5 & 6) has many distinctive
features, most of which may be explained as adaptations to a swimming
habit. The carapace is elongated, and its flanks taper so evenly posteriorly
that it may fairly be described as boat-shaped. The first abdominal tergum
is notably narrower than the posterior margin of the carapace, and the whole
abdomen is narrow and more distinctly flexed under the thorax than is usual
in Raninide. On the ventral surface there is a broad and long flat plastron
in the formation of which the eleventh and twelfth sterna take an even larger
share than the tenth (fig. 6). The twelfth sternum is of considerable length
antero-posteriorly and is also broad and nearly flat. The articular cavities
of the second pair of pereiopods are placed at its hinder border and face
backwards. The thirteenth sternum being very short, the articular cavities
of the third pereiopods are close to those of the second, and the two pairs of
limbs, whilst closely contiguous to one another, are separated by the whole
length of the twelfth sternum from the first pereiopods. The last pair of
pereiopods are so reduced and slender that they are usually described as fili-
form ; they terminate in minute pointed dactyli. The reduction of the last
pair of legs and the narrowing of the base of the abdomen seems to be cor-
related with the suppression of the posterior branchial orifices ; at any rate,
I can find no trace of these orifices in Raninoides. Nor could they be of
much service if present, for the external water conduits which in other
forms admit of a flow of filtered water to these orifices are here wanting.
Though the eleventh, twelfth, and thirteenth thoracic epimera are as much
exposed to the surface in Raninotdes as in other Raninide, they are not con-
cave and are scarcely overhung by the edge of the branchiostegite. The
epimeron of the eleventh somite is relatively small, and so far from being
concave that it is slightly convex. The epimera of the twelfth and thirteenth
somites are obliquely grooved, and into these grooves the meri of the second
and third pereiopods, when bent forwards and pressed against the sides
of the body, fit so closely that no water can pass between. The first
pair of pereiopods are slender, and when at rest are directed forwards,
their large cordate dactyli being pressed against the bases of the cheli-
peds. Their function is obscure, but probably they are used for digging.
The characters of the second and third pereiopods are clearly shown in
fig. 6. In these limbs the coxa and basis are directed backwards: the short
ischium is nearly transverse to the long axis of the body: the merus is long
and slender: the carpus normal, but it is to be observed that both it and the
merus are scantily furnished with short hairs. The propodus and dactylus
are flat and greatly expanded and the ‘“fan-hinge” between them
is a noticeable feature. A fringe of long stiff hairs is borne on the
hind edge of the propodus and on the inner margin of the dactylus. The

A STUDY IN CARCINOLOGY. 69

efficiency as organs of natation of these broad blades borne at the end of
slender and very mobile limbs is apparent. Hvidently Ranznoides is a
strong swimmer and,a poor digger, and as such stands at the end of
the series opposite from Ranina. As the posterior branchial orifices are
absent, one might expect to find special arrangements for an incurrent
respiratory stream in the frontal region, but, though I have looked carefully
for such, I have been quite unsuccessful. However, I will deal with
this question further on.

Lyreidus (fig. 4) exhibits so many resemblances to Raninoides that one
can hardly doubt that the two are closely related, yet, as I shall show, they
are clearly adapted to different conditions of life. In Lyreidus the abdomen
is narrow; its first three segments lie nearly in a straight line with the
carapace, but the fourth is of peculiar shape, bears a strong median dorsal
spine, and constitutes as it were the knuckle of a sharp ventral flexure. The
sixth abdominal segment is long and narrow ; its posterior angles are pro-
duced into small aliform processes which at first sight might be mistaken
for vestigial uropods, but they are only processes of the tergum having on
their ventral surfaces small coneayities which, when the abdomen is flexed to
its fullest extent, engage with small knobs on the two pterygoid processes
extending backwards from the twelfth sternum. No such apparatus for
locking the flexed abdomen to the sternum is seen in any other Raninid.
The last pair of pereiopods are so slender that, like those of Raninoides, they
may be described as filiform: they terminate in small flattened elliptical
dactyli. Correlated with the reduction of the abdomen and of the last pair
of pereiopods is the absence of posterior branchial orifices. I have studied
this point carefully and am certain that these orifices are non-existent in
LL. tridentatus, nor could I find any trace of them in the large specimen of
LL. channeri in the Natural History Museum. As there are no posterior
respiratory orifices the water conduits of the flanks are, as might be expected,
absent. The epimera of the posterior thoracic somites are nearly flat and
the edge of the branchiostegite is but slightly prominent and bare of sete.
In the frontal and oral regions there are also many points of resemblance.
Though the “front” is truneated and scarcely narrower than the broadest
part of the carapace in Raninoides, the distance between the extra-orbital
spines and the lateral spines shows that this region has undergone elongation.
Further examination shows that it is the antennary somites that has been
lengthened, for the antennary sternum, which has more or less the shape of an
equilateral triangle in Notopus and WNotosceles, is an isosceles triangle in
Raninoides. In Lyreidus the elongation of the antennary somite is carried
to an extreme, and the front being narrowed, the fore part of the carapace is
produced into the snout characteristic of the genus, the lateral spines being
situated far behind the orbits. As I shall show further on, the details of the

70 DR. G. C. BOURNE ON THE RANINID# :

antennules and antenne are extraordinarily similar in Raninoides and
Lyreidus (figs. 36-39) ; so also are details in the first and second and even in
the third maxillipeds. In all these things the relationship seems very close,
but in other respects the two forms are divergent. In Lyretdus the eleventh
sternum is large but narrowed between the articulations of the first pereio-
pods and produced backwards into two small pterygoid processes which
form the sides of a hollow in which the last segment of the abdomen can be
lodged, as described above. Owing to the presence of these pterygoid
processes the twelfth sternum appears to be deeply fitted, but it is really
short, narrow, and flat, in this differing much from Raninoides. The thirteenth
sternum being also short the coxee of the first three pairs of pereiopods are
close together, and in this respect Lyreidus resembles Notopus rather than
Notopoides, Notosceles, or Raninoides. In the first and second pereiopods of
Lyreidus the ischiomerus is long, slender, and scantily fringed with hairs ;
the carpus father short ; the propodus nearly twice as long as it is broad ;
the dactylus elongate and almost styliform, with a strong external ridge. In
the third pereiopods the propodus is about as broad as it is long and the
dactylus cultriform. These are clearly digging and walking legs, and one
may conclude that the descendants of a form in which reduction of the
abdomen and last pair of legs led to the suppression of the posterior branchial
orifices, diverged as they adopted a more exclusively swimming habit into
Raninoides, as they became more exclusively burrowers into Lyreidus.
Lyreidus must bury itself deeply, with the tip of its narrow elongated snout
just breaking the surface of the sand. Having no posterior branchial orifices
it must be dependent on an inhalant current setting in somewhere in the
region of the snout. One would expect some specialised inhalant apparatus,
but there is very little evidence of such. Indeed, it is a singular thing that
Ranina, Notopus, and Notosceles, all of which have well-developed posterior
branchial orifices, have in addition much more specialised orbital and anten-
nary arrangements for directing the flow of an incurrent respiratory stream
than have Raninoides and Lyreidus, in which posterior respiratory orifices are
absent. In Notosceles the antennary structure is not very different from
that of Ranina. The flagellum is longer (figs. 41 & 41a), but the thick
shortened segments of the peduncle, the flabellate shape of the fourth joint
with its fringe of plumose hairs, the greatly developed crest of the third
joint also fringed with hairs, are very similar. So also is the flabellate shape
of the basal joint of the antennules. Tig. 57 is a frontal view of the “face”
of Notosceles showing the antenne as nearly as I can draw them in their
natural position. In the centre, below the rostrum, is the narrow exhalant
passage bounded by the basal joints of the antennules. The orbits are closed
below by the crests of third segments of the antennary peduncles, and the
spaces below are filled in by the proximal segments of the antennze which in

A STUDY IN CARCINOLOGY. al

their turn are overlapped by the broad anterior margins of the meri of the
third maxillipeds. The left antenna is closed nearly as far as it will go
against the antennule, and this is the more normal position: that of the right
side (the spectator’s left) has been pulled outwards to show the passage
between the third and fourth joints of the antennary peduncle and the basal
joint of the antennules. Through this passage water can pass from the
orbits to the inhalant chamber covered in below by the meri of the third
maxillipeds. The arrangements are very much the same as in Ranina though
somewhat less highly specialised. In Raninoides (figs. 36 & 37) and Lyreidus
(figs. 38 & 39) the basal joints of both antennules and antennz are longer
and more slender, and to that extent more primitive than in Notosceles and
Ranina. In the antennules the admedian surfaces of the basal joints of the
antennules are expanded and somewhat concave, so that when opposed they
form a conduit for the exhalant current. The third segment of the anten-
nary peduncle is long; a good deal longer in Lyreidus than in Raninoides.
The crest is prominent but narrow, and extends nearly as far forward as the
end of the fourth segment. The proportions differ slightly in the two species,
but the general character is the same. The nature and distribution of the.
hairy fringes are clearly shown in the drawings. ‘The relative positions of
the antennules and antennz are shown in fig. 58 for Raninoides: they are
hardly different in Lyreidus. It will be observed that the “spout” of the
antennary sternum is produced to open well forward between the basal joints
of the antennules: it hardly extends as far forward in Lyreidus. In both
genera, but to a greater extent in Lyreidus than in Raninoides, the third
pedunecular segment of the antenna is overlapped by the tip of the elongated
merus of the third maxillipeds, and the portion not overlapped is covered by
a tuft of long hairs projecting forward from the tip of the merus. The inner
faces of the third peduncular segments of the antennze and their crests are so
shaped that when bent inwards they fit very exactly to the outer surfaces of
the basal joints of the antennules, and when in this position completely bar
the passage of water from the orbits to the inhalant chamber above the third
maxillipeds. But when the antennz diverge slightly outwards a clear
passage is left between the antennules and the antennze, and the intrusion of
sand is guarded against by the overlapping bristles of both. Through this
passage, I think, the inhalant current must pass when the scaphognathite is
working normally, and through it the excurrent stream must flow when the
action of the scaphognathite is reversed. But the subject is very obscure,
and I am by no means content with this explanation. I give it for what it is
worth in the hope that some favourably placed naturalist may read it, find
fault with it, and straightway proceed to make observations and experiments
on the living animals. The results cannot fail to be of interest. I feel
certain only of this, that there is an inhalant current between the third and

72 DR. G. C. BOURNE ON THE RANINIDZA:

the second and first maxillipeds. It is to direct this current under the edge
of the mouth frame into the branchial chamber that the exopod of the second
maxilliped is elongated and modified by being channelled along its ventral
surface. The modification is carried furthest in Wotosceles (fig. 44) and
Ranina (fig. 42), two forms in which the accessory antennary modifications
are highly developed, and in these the exopod has lost its flagellum. But in
Notopus, in which genus the antenne are most highly specialised for respira-
tory purposes, the exopod in question (fig. 32) retains a large flagellum,
and the flagellum is present but small in Lyreidus (fig. 52) and Raninoides
(fig. 49). This may seem a trivial character, but it gains importance when
comparison is made with the same appendage in the Leucosiidee. In Philyra
levis for example (fig.54) the exopod of the second maxilliped is of simple
shape, shorter than the endopod and convex ventrally. In this species, as in
all the Leucosiidee, there is a definite exostegal canal covered in below by the
exopod of the maxilliped: the second maxilliped, therefore, does not lie in
the course of the incurrent stream and is not modified. I have already
(p. 54) called attention to the difference between the first maxillipeds of
the Raninidee and those of other Oxystomatous crabs. Indeed, as f have
already indicated when dealing with the endophragmal skeleton, the anterior
thoracic sterna, and other structures, the Leucosiide differ in a hundred ways
from the Raninidee, and I regard the comparison with Lyreidus as wholly
illusory. The exostegal canal of the Leucosiide is most probably derived
from the shallow groove in the pterygostome occupying the same position
in Cyclodorippe. It would be interesting and not wholly irrelevant to
attempt to trace out the various lines of descent in the families of Oxysto-
mata other than the Raninide, and I have collected much evidence on this
subject based on a study of the endophragmal skeleton and other anatomical
features not taken into account by M. Edwards and Bouvier. But it would
almost double the length of this memoir if I were to make the attempt.
I have already written enough, and haye, I hope, proved my main thesis that
the Raninidee cannot have descended, by way of the Dorippide and the
Dromiacea, from the Macrura. They have originated independently from
the last named, and must therefore be separated from other Oxystomes as a
separate tribe Gymnopleura. I have endeavoured to show that the numerous
brachyuran features exhibited by this tribe must be explained by their having
adopted the same habits as other crabs, and that such modifications as the
reduction of the abdomen; the loss of the uropods; the adherence of the
branchiostegite to the thoracic epimera (exhibited, however, in a quite
peculiar manner in the Gymnopleura) ; the expansion of the ischium and
merus of the third maxillipeds; the broadening of the front, with which
is correlated the formation of orbital cavities and the reduction of the -
rostrum; the great development of the muscle-cavities of the thorax,—are all
characters of high survival value in decapods which have acquired the habit

A STUDY IN CARCINOLOGY. 73

of burrowing in the sand for protection and concealment. A decapod
crustacean which habitually passes a large part of its existence buried in the
sand, must undergo a considerable modification of its respiratory arrange-
ments, and I have attempted to give an intelligible account of these modifi-
cations in the various members of the Gymnopleura. In the end they lead,
as is natural enough, to the suppression of the inhalant current at the hind
end of the thorax and to the substitution of inhalant currents in the antennary
region. I have said, in the earlier part of this memoir, that the Gymnopleura
are crabs by definition. They retain so many Macruran features that they
cannot be said to be crabs by ancestry, and there is one small Macruran
feature which I have omitted to mention before because I only recognised it
when the greater part of this paper was written. The statocyst, usually
closed in the Brachyura, is openin the Gymnopleura. Certainly so in Ranina,
Notosceles, and Lyreidus, but | have not been able to satisfy myself of the
existence of the aperture in all the species examined because the antennules
are very small and I have not been able to cut sections. The position of the
aperture of the statocyst is clearly shown in fig. 34 for Ranina.

I must conclude with a description and definition of the new genus and
species WVotosceles Chimmonis.

There can be little doubt that Raninoides serratifrons of Henderson (36)
belongs to the genus Notosceles and not to Raninoides, but his species,
from Ceylon, is different to mine. He mentions another example of
serratifrons from Holothuria Bank, N.W. Australia, as being in the collec-
tions of the British Museum of Natural History, but I was unable to find
it there. There are considerable differences between Notosceles and
Raninoides, many of which have been referred to in the course of this

paper.

NoTosvELEs, noy. gen.

Carapace ovate, convex from side to side, minutely punctate or smooth
for the most part but granular towards the frontal region; fronto-orbital
border little more than half as wide as the broadest part of the carapace; the
frontal region marked off by a slight transverse ridge joining the bases of
the extra-orbital spines; the rostrum pointed, fairly prominent with two
flat Jateral and basal teeth. Ocular peduncies rather short, their terminal
joints inflated, their corneze of moderate size. Antennal peduncle short and
broad, the flagellum small. Merus of the third maxilliped little more than
half as long as the ischium. The second and third sternal elements of the
sternal shield (sterna xi & xii) broad and convex in front, but narrowed
posteriorly so that the bases of the first as well as the second pereiopods are
approximated to the middle line. Last pair of pereiopods reduced in size,
situated above and in front of the penultimate pair, their dactyli small, oval,

74. DR. G. C. BOURNE ON THE RANINIDA:

and flattened. First abdominal tergum not much narrower than the posterior
margin of the carapace, about equal in breadth to» the second tergum, the
remaining terga progressively narrower ; the telson small and triangular.

NoroscELES CHIMMONIS, n. sp. (Pl. 4. figs. 2 & 3.)

Carapace minutely punctate to smooth over the greater part of its surface,
but granular towards the frontal region: proportion of length to breadth of
carapace about 5:3. The frontal region is marked off by a distinct trans-
verse ridge joining the bases of the extra-orbital spines. _The median frontal
region is fairly broad and produced anteriorly into a median pointed rostral
spine at either side of the base of which is a triangular flattened and some-
what upturned spine, the margins of all three spines minutely serrated. On
either side a fissure separates the median frontal projection from a short
triangular tooth or spine, and outside the latter is a prominent incurved
extra-orbital spine. A single lateral spine occurs on either side of the cara-
pace a short distance behind the extra-orbital spine, and nearly of the same
size as the latter. On the upper surface a slight carina runs from the
median rostral spine to the transverse granular ridge separating the fronto-
orbital region from the rest of the earapace. In the chelipeds the anterior
edge of the ischium is serrated ; the merus largely dilated externally at its
base; the carpus granular above and without a distal spine ; the propodus
bears two subequal spines on its lower margin, and on its upper surface are
two parallel ridges separated by a narrow groove ; the immobile claw bears
four denticles on its inner margin, In the third maxillipeds the relative
length of the merus to the ischium is as 6:10; the lower surface of the
merus is granular and hairy. ‘The pterygostomial regions are hairy. In the
sternal region the chelipedal sternum is of the shape characteristic of
the Raninide, the following sternal element flat, broad in front and narrow
posteriorly ; the third element of the sternal plastron triangular, convex, the
apex of the triangle anterior. The articular cavities of the first and second
pereiopods approximated to the middle line. The first abdominal tergum is
flat and quadrangular, not much narrower than the posterior margin of the
carapace; the second abdominal tergum is nearly equal to it in width; the
remaining abdominal terga narrow rapidly and the telson is small and sub-
triangular in shape.

Two specimens from the Sulu Sea, both males. Length of the larger
specimen with the abdomen fully extended 28 mm.: length of carapace
20 mm.: greatest width of carapace 13 mm.

The species is named after Captain Chimmo, R.N., of H.M.S. ‘ Nassau,’
whose collections from the Sulu Sea were presented to the Oxford University
Museum in 1872.

Lam of the opinion that Raninoides serratifrons, Henderson (36) should

o : 6

A STUDY IN CARCINOLOGY. 75

be placed in the genus Notosceles, which differs from Raninoides in the pro-
portions of the carapace, the shape of the rostrum, the lesser width of the
fronto-orbital region, the Jarger corneze of the eyes, the proportions of
the joints of the antennal peduncles, and in other characters, but especially
in the retatively much greater width of the base of the abdomen, and the
proportions and shape of the sternal shield.

LITERATURE.

. Aucock, A.—Natural History Notes from H.M. Indian Marine Survey Steamer

“Investigator.” Ser. ii.no.1. Ann. Mag. Nat. Hist. (6) xiii. 1894, p. 406.

. Ibid. Materials for a Carcinological Fauna of India. Journ, Asiatic Soc. Bengal, Ixy.

1896, p. 134.

. Brrrner, A.—Beitrige zur Kentniss tertidirer Brachyuren Formen. Denkschr, Kais.

Wiss. xlvyili. 1853.

. Boas, E. V.—Studier over Decapodernes Slaegtskabsforhold. Kong. Danske Videnskab.

Skrifter, i. Copenhagen, 1880-85.

. Boun, G.—Sur la respiration du Careinus menas Leach. Comptes Rendus de l’Acad.

Sei. exxy. 1897, p. 441.

. Ibid. Sur la renversement du courant respiratoire chez les Décapodes. Loc. cit. cxxy.

1897, p. 5389.

. Ibid. De l’enfouissement chez les Homaridés et les Thalassinidés. oc. cit. exxyii.

1898, p, 781.
Borrapaitr, L. A.—A series of papers in Professor J, Stanley Gardiner’s ‘Fauna
and Geography of the Maldive and Laccadive Archipelagos,’ including :—
Land Crustaceans. Vol. i. pt. 1. :
Marine Crustaceans: Introduction. Vol. i. pt. 2.
Varieties in the Decapod Crustaceans. Vol. i. pt. 2.
Portunidee (Swimming Crabs). Vol. i. pt. 2.
The Xanthidz and some other Crabs. Vol. i. pt. 3, p. 287.
Some Remarks on the Classification of Crabs. Vol. i. pt. 4, p. 424.
The Crabs of the Catametope families. Vol. i. pt. 4, p. 429.
The Sand Crabs (Oxystomata). Vol. i. pt. 4, p. 434.
The Sponge Crabs (Dromiacea). Vol. ii. pt. 1, p. 574.
The Spider Crabs (Oxyrhyncha). Vol. ii. pt. 2, p. 681.
On the Classification and Genealogy of the Reptant Decapods. Vol. ii. pt. 2, p. 690.
The Hippidea, Thalassinidea, and Scyllaridea. Vol. ii. pt. 3, p. 750.

. Ibid. Crustacea. Pt, I. Decapoda. Brit. Antarctic (Terra Nova) Exped. 1910. Nat.

Hist. Rep. Zool, vol. iii. 1916, p. 75.

. Ibid. Crustacea. Pt. IL. Porcellanopagurus: an instance of Carcinization. Loe. cit.

vol. iii, 1916, p. 111.

. Bouvier, E. L.—Le systéme nerveux des Crustac s Décapodes. Ann. Sci. Nat. Zool.

(7) vii. 1889, p. 73.

. Ibid. Sur Vorigine homarienne des Crabes. Bull. Soc. Philomath. Paris, (8) viii. 1895,

pl.

, CALMAN, W. T.-—Crustacea in Ray Lankester’s ‘Treatise on Zoology.’ London, 1909.
. Dana, J. D.—U.S. Exploring Expedition: xiii. Crustacea.

. Epwarps, H. Mirne.—Histoire Naturelle des Crustacés, 3 vols. Paris, 1837.

. Ibid. Crustacea in Cuvier’s ‘Régne Animal.’

76 DR. G. C. BOURNE ON THE RANINIDE :

27. Epwarps, ALPHONSE.—Note sur l’existence de Crustacés de la famille des Raniniens
pendant la période Crétacée. Comptes Rendus, lv. 1862, p. 492.

28. Epwarps, A. Mrine, and Bouvier, BH. L.— Report on results of dredging by U.S.
Survey Steamer ‘ Blake,’ xxxix.: Les Dromiaiés et Oxystomes. Mem, Mus.
Comp. Zool. Harvard, xxvii, 1902.

29. Ibid. Exped. Sci. du ‘Travailleur’ et du ‘Talisman’: Crustacés Décapodes.
Brachyures et Anomures. Paris, 1900.

30. Garstanc, W.—The habits and respiratory mechanism. of Corystes cassivelaunus.
Jour, Mar. Biol. Assoc. iv. 1896, p. 223.

31. Ibid. On some Modifications of Structures subservient to Respiration in Decapod
Crustacea which burrow in the sand. Quart. Journ. Mier. Sci. xl. 1897, p. 211.

32. Ibid. The Function of Antero-lateral Denticulations of the Carapace in Sand-
burrowing Crabs. Journ. Mar. Biol. Assoc. iv. 1896, p. 396.

33. Ibid. The Systematic Features, Habits, and Respiratory Phenomena of Portumnus
nasutus. Journ. Mar. Biol. Assoc. iv. 1896, p. 402.

34, Haan, W. pre—Crustacea in Siebold’s ‘Fauna Japonica.’ 1850.

35. Hrnperson, J. B.—Report on the Anomura: ‘Challenger’ Reports, Zoology, xxvii.
1888.

36. Ibid. A Contribution to Indian Carcinology. Trans. Linn. Soc. Lond., Zool. (2) v.
1393, p. 408.

37. Hrreicx, F. H.—Natural History of the American Lobster. Bull. Bureau of Fisheries,
xxxix. 1911,

38. Huxuery, T. H.—The Crayfish, an Introduction to the Study of Zoology. London,
1880.

39. Int, J. EK. W.—Decapoda Brachyura der ‘Siboga’ Expedition: iii, Oxystomata.
Monogr. xxxixb*. Leiden, 1918.

40. Laurie, R. D.—Reports on the Marine Biology of the Sudanese Red Sea: Brachyura.
Journ. Linn. Soc., Zool. xxxi. 1915, p. 407.

41, OrTMann, A.—Die decapoden Krebse des Strasburger Museums. Zool. Jahrb, Abth.
f. Systematik, vi. 1892.

42. Ibid. Crustacea in Bronn’s ‘Thierreich.’ 1901.

43. Ratupun, Mary J.—Marine Brachyura. Rep. Percy Sladen Expedition, vol. in. 1912,
p. 191.

44. Wuirr, ApAM.—Short descriptions of new or little-known Decapod Crustacea. Proc.
Zool. Soc. 1847, p. 227.

EXPLANATION OF PLATES 4-7.
LETTERING,

A,;. Antennules. A». Antenne. abr. arthrobranch. ad. apodeme. aph. arthro-
phragm. at. antennal respiratory tube. dst. branchiostegite. cr. crest of antenna. edpl.
endopleurite. edst. endosternite. en.p. endopodite. Lp. Epimeron. ep.ad. epimeral
apodeme. epl. epaulette of the tenth sternum. 2. excretory pore. exp. exopodite.
g. ganglia of nerve chain. 7b. labrum. mbr. mastigobranch. md. mandible. oe. ocular
peduncle. oc.t. tergum of the ocular segment. a. cesophagus. P. Pereiopods. pbr. podo-
branch. pgt. pterygostome. . plbr. pleurobranch. 2. Rostrum. shy. rudimentary setobranch
of the third maxilliped. sey. aperture of statocyst. spd. strophidium. spng. strophingium.
st.a. sternal artery. s¢.c. sternal canal.

The roman numerals refer to segmental structures, sterna, epimera, ganglia, etc.
Intersegmental structures are indicated thus ix/x.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig, 5
Fig. 6
Fig. 7
Fig. 8
Fig. 9
Fig. 10.
Fig. 11
Fig. 12
Fig. 13.
Fig. 14.
Fig. 15
Fig. 16.

A STUDY IN CARCINOLOGY. 717

Prater 4,

. Ventral view of Notopus dorsipes, de Haan. The chelipeds have been turned

outwards and the pereiopods are extended to show their structure as far as
possible.

. Dorsal view of Notosceles chimmonis, noy. gen. et sp. The chelipeds and pereiopods

have been extended as in fig. 1.

. Ventral view of Notosceles chimmonis.
. Ventral view of Lyreidus tridentatus, de Haan. Note the elongated digging dactyli

of the first and second pereiopods.

. Dorsal view of Raninoides personatus, White MS., Henderson. The first and third

pereiopods of the left side are missing.

. Ventral view of the same specimen as fig. 5. Note the large size of the sternal

plastron, due to the great enlargement of sternum xii and the adaptation of the
second and third pereiopods for swimming.

. Left side view of Notopus dorsipes. The first three pairs of pereiopods have been

pulled downwards to show the large extent of epimera xi and xii left uncovered
by the edge of the branchiostegite.

. La. . , il. yj .d
Central nervous system of Ranina dentata. i. ocular, ii. antennulary, iii. antennary

nerves; iv. mandibular nerves; y—ix. nerves to maxillee and_maxillipeds issuing
from the subcesophageal ganglion; x. ganglion of cheliped; xi-xiy. ganglia of
the four posterior thoracic segments; xy-xx. ganglia of the abbreviated
abdominal nerve chain. st.a. aperture for passage of the sternal artery.

. Right half of the endophragmal skeleton of Runina dentata as seen from within

after division into two halves by a sagittal cut. The more anterior thoracic
somites are omitted. The thoracic portion of the central nervous system is
shown zm situ. *Note the great depth of the sagittal apodemes of sterna xii, xiii,
and xiv. g.11-g. 14, the four posterior thoracic ganglia.

Central nervous system of Lyreidus tridentatus. Lettering as in fig. 8.

. A drawing from the left side of the eleventh and twelfth segments of the common

lobster to show the relations of the endophragmal to the exoskeleton. xi and
xii. sterna of the eleventh and twelfth segments. The preparation has been
slightly tilted to the right to show the endopleurites edp/., consequently the
epimera Ep. xi and Zp. xii are foreshortened.

. A left side view of the thoracic skeleton of Dromia vulgaris. The ventro-lateral

projections of the sterna have been omitted, and the preparation is slightly tilted
over to the right. xi-xiy. the four posterior thoracic sterna.

PLATE 5,

A posterior view of the thoracic skeleton of the common lobster, showing the large
size of the penultimate and the structure of the last thoracic sternum.

A posterior view of the thoracic skeleton of Ranina dentata, illustrating the shape
of the last three thoracic sterna and the formation of the sella turcica posterior.
sg.a. sagittal apodemes,

. A similar view of the thoracic skeleton of Dromia vulgaris. In the absence of

sagittal apodemes in the posterior thoracic sterna the arthrophragms xiy/xv
do not meet in the middle line, and there is no sella turcica.

The endophragmal skeleton of Notopus dursipes as seen from above after removal
of the upper parts of the epimera. 6, 7, 8, the articular cavities of the
second maxilla and first and second maxillipeds. The endosternites edst. and
endopleurites edpl. are clearly shown,

oS

be

18.

ig. 19.

27

DR. G. C. BOURNE ON THE RANINIDA:

Thoracic skeleton of Notopus dorsipes seen from the left side, illustrating the great
depth of the epimera and the oblique raised ridge on the 11th, 12th, and 13th
epimera, against which the lower margin of the branchiostegite fits: ix, x, the
ninth and tenth sterna; vi/vii to ix,x. intersegmental arthrophraems.

Frontal view of the thoracic skeleton of Notopus dorsipes. The articular socket of
the second maxilla of the left side is omitted.

The antennary region of Nephrops norvegicns. The rostrum is cut short and the eyes,
antennules, and antenn are removed from their sockets. i. The membranous
ocular sternum; ii. the antennulary sternum ; iii. the antennary sternum ; iv. the

antennary fossee. ., sclerite identified as tle mandibular epimeron.

. Side view of the ocular and the antennary regions of Nephrops norvegicus. The

rostrum is cut short and the antennules and anténne are removed. oc.t. sclerite
representing the tergum of the ocular segment; i. the sternum of the ocular
segment: the epimera of this segment are membranous.

. Antennary region and buccal frame of Aithusina gracilipes, Miers. Ep. iy. epimeron

of the mandibular segment.

2. Antennary region and buccal frame of Notopus dorsipes. The appendages have been

_ removed, with the exception of the right ocular peduncle and the left mandible.
Ay, Ao,®sockets of the antennules and antenne ; pgt. sub-antennary lobe of the
pterygostome.

. A similar view of the antennary region and buccal frame of Ranina dentata.

PLATE 6,

A similar view of the antennary region and buccal frame of Notosceles chimmonis.

25. A similar view of the antennary region and buccal frame of Lyreidus tridentatus,

showing the great elongation of the antennary sternum iii. pg. paraglossee.

. Perspective sketch of the buccal frame of Rania dentata, to show the hood-shaped

mandibular sternum iv.
Antennary region and buccal frame of Dromia vulgaris, The appendages have been
removed, with the exception of the right ocular peduncle.

Figs. 28-83. Notopus dorsipes. Fig. 28. Right antennule: admedian aspect.
Figs. 29 & 29a. Right antenna, admedian and outer aspects. 1-5, segments of the peduncle.

Fig.
Fig.

30.
31.

34,
_ Bb,

». 36.

89.
ig. 40.
g. 41,
43,

Second maxilla with scaphognathite.

First maxilliped. Fig. 32. Second maxilliped: note the relatively large flagellum
of the exopodite. Fig. 33. Third maxilliped: note the oblique line of hairs on
the ischium also seen in Ranilia, M. Edw.

outer and Fig. 34a, admedian views of the right antennule of Ranina dentata.
Note the external opening of the statocyst, scy.

outer and Fig. 85a, admedian views of the right antenna of Ranina dentata.
1-5, segments of peduncle. f

Outer aspect of the right antennule of Raninordes personatus. Note the infolding
of the cuticle to form the statocyst, and the external aperture of the latter at scy.

. Admedian view of the right antenna of Raninotdes personatus. In this genus and

in Lyreidus the two hasal segments of the peduncle are indistinguishably fused
together.
Admedian view of right antennule of Lyreidus tridentatus.
Admedian view of right antenna of Lyreidus tridentatus.
Admedian view of right antennule of Notosceles chimmonis.
outer and Fig. 41a, admedian views of the right antenna of Notosceles chimmonis.
Notosceles chimmonis. Third maxilliped.

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A STUDY IN CARCINOLOGY. 19
6
PLATE 7.

Fig. 42. Right second maxilliped of Ranina dentata, The exopodite is deeply grooved on
its surface and has no flagellum.

Figs. 44-47. Notosceles chimmonis. Fig. 44. Second maxilliped, the exopodite of which, like
that of Ranina, is deeply grooved on the ventral surface and bears no flagelluin.
Fig. 46. First maxilliped. Fig. 46. Second maxilla with scaphognathite.
Fig. 47, First maxilliped.

Figs. 48-50. Raninotdes personatus. Fig. 48. Third maxilliped. Fig. 49. Second maxilliped,
the expodite of which bears a minute flagellum. Fig. 50. First maxilliped.

Figs. 51-53. Lyreidus tridentatus. Fig. 51. Third maxilliped. Fig. 52. Second maxilliped,
the exopodite of which bears a minute flagellum. Fig. 53. First maxilliped,
the exopodite of which is elongated in correlation to the elongation of the au-
tennary sternum.

Fig. 54. Second maxilliped of Philyra levis. The podobranch and mastigobranch are
wanting: the exopodite is not grooved on the ventral surface and bears a short
flagellum.

Fig. 55. First maxilliped of Philyra levis, The mastigobranch is of great length and the
exopodite is of normal shape and bears a flagellum. These appendages of Philyra
should be compared with the corresponding appendages of the several genera of
Raninide.

Fig. 56. Notopus dorstpes: ventral view of the antennary region to show the relation of the
respiratory tube to the antennary sternum. The pterygostomial portions of the
carapace have been cut away.

Fig. 57. Frontal view of Notosceles chimmonis to show the antenne in their natural position
and their relation to the orbits. The right antenna has been pulled slightly
outwards to expose the antennules.

Fig. 58. Ventral view of the anterior part of thorax and head of Raninoides personatus. The
right third maxilliped has been removed and the left displaced from the middle
line to expose the mouth frame and to show the elongated exopodites and endo-
podites of the first maxillipeds forming the floor of the spout-shaped antennary

sternum. The latter extends well forward between the basal joints of the
antennules.

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Vou. XXXV. ZOOLOGY. No. 232.

CONTENTS.
Page
I. The Theory of Recapitulation: a Critical Re-statement of the

Biogenetic Law. By Waurer Garstanc, M.A., D.Se.(Oxon),
Professor of Zoology in the University of Leeds. (Communi-
cated by Dr. W. Baruson, F.R.S., F.L.8.) (Witha Text-figure.) 81

II. On the Terrestrial Isopod Eluma celatum (Miers) = purpurascens,
Budde-Lund. By Watrer EH. Cottines, D.Sc., F.L.8., Keeper
of the Yorkshire Museum, York. (Plate 8.) .................0055 103

III. On Two new Terrestrial Isopods from Madagascar. . By WALTER
Hi. Continex, D.8c., F.L.S., Keeper of the Yorkshire Museum,
BYCorissan Ckyla hen Oh) Re peta cane a ey een tcl ananaNWare umicsiinr leaned on 107

TV. On the Mouth-parts of the Shore Crab. By L. A. Borrapaize,
Se.D., Fellow and Tutor of Selwyn College, Cambridge, and
Lecturer on Zoology in the University. (Communicated by

Prof. H. 8. Goopricu, M.A., F.R.8.,Sec.L.8.) (Plates 10,11.) 115

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1922.

LINNEAN SOCIETY OF LONDON.

LIST OF THE OFFICERS AND COUNCIL.
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PRESIDENT.
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LIBRARY COMMITTEE.
The Officers ex officio, with the following in addition :—
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E. T. Browne, M.A. B. B. Woodward, Esq.
Dr. W. T, Calman, F.R.S.

THE THEORY OF RECAPITULATION. 81

The Theory of Recapitulation: A Critical Re-statement of the Biogenetic
Law. By Watrer Garstanc, M.A.,- D.Se.(Oxon), Professor of
Zoology in the University of Leeds. (Communicated by Prof. W.
Bareson, F.R.S., F.L.S.)

(With a Text-figure.)
[Read 2nd June, 1921.]

Mone than half a century has passed. since Haeckel propounded his “ funda-
mental Biogenetic Law” (1866). It played a great part in the campaign
for the recognition of Evolution, has inspired and still inspires much good
work in Paleontology, but, as a working hypothesis in Embryology, is
admitted to have evoked little but controversy and confusion. This history
alone renders it probable that the law is a mixture of sound and questionable
elements, but the two have never been satisfactorily disentangled. The late
Dr. C. H. Hurst (1893), Adam Sedgwick (1894 & 1909 (a)), and Geoffrey
Smith (1911), Oscar Hertwig (1898 & 1896), and Morgan (1908), among
others *, have criticised particular aspects of it, but no one has presented a
complete theoretical scheme capable of replacing Haeckel’s as an explan-
ation of the relations between ontogeny and phylogeny. Lately MacBride
(1914 & 1917), from the embryological side, and Bather (1920), from the
standpoint of paleontology, have revived the full Haeckelian doctrine ; and
the former has even considerably extended it, though neither, so far as I can
see, has refuted, or even appreciated, the force of the criticisms made by
their predecessors. As it is not to the credit of science that Zoology should
harbour a “law” which, like a creed, may be accepted or rejected at
pleasure, and as I believe the basis of this law is demonstrably unsound,
I venture to make a renewed attempt to define the pointsat issue. The most
satisfactory way of doing this appears to be to re-state, in accord with
modern knowledge, the theoretical relations of ontogeny to phylogeny, and
then to subject the alternative theories to verification by test-cases. As the
old Jaw was essentially morphological, I exclude from present consideration
all bionomical and etiological questions not directly involved.

1. The two aspects of Haeckel’s doctrine—the statement of fact and the
theory of causation—were summed up by himself in the phrases: ‘* Onto-
genesis is the recapitulation of Phylogenesis” and ‘“ Phylogenesis is the
mechanical cause of Ontogenesis.” In these now familiar terms the new

* Bateson’s criticism of the law of von Baer, though not specially referring to Haeckel’s
modification of it, should be included here (1894, pp. 8-10).

LINN. JOURN.— ZOOLOGY, VOL. XXXV. 6

82 PROF. W. GARSTANG ON THE THEORY OF REOAPITULATION :

conception of evolution was wedded, fifty years ago, to current ideas of
ancestry, heredity, and development: Ancestors created, heredity transmitted,
and development repeated the order of creation.

2. To Haeckel, phylogenesis meant “‘the chain of manifold animal forms
which represent the ancestry” of an organism, i.e. the phyletic line of
succession of adults.. Ontogenesis was, and is, the succession of form-
changes between zygote and adult of the same organism. The ontogenetic
sequence was regarded as resembling, and actually caused by, the phyletic
sequence of adults, which had preceded it.

3. But Haeckel overlooked the other evolutionary sequence, the phyletic
line of succession of zygotes, running more or less parallel with the adult
sequence, step by step, though steadily diverging. Hvery elaboration of
adult form, even of its degree of pliability under environmental influence
(for there are great differences among animals, as among plants, in this
respect), was preceded by a corresponding elaboration of zygotic structure *,
nuclear or cytoplasmic or both, determining, under suitable conditions, the
form and character of the ontogenetic changes and their result. Through
the whole course of Evolution, every adult Metazoan has been the climax of
a separate ontogeny or life-cycle, which has always intervened between adult
and adult in that succession of forms which Haeckel terms ‘“‘ Phylogenesis.”
The real Phylogeny of Metazoa has never been a direct succession of adult
forms, but a succession of ontogenies or life-cycles.

4, This was so from the very beginning, when zygote and adult were
indistinguishable in form as ancestral Flagellate Protozoa. Zygosis must
have been followed, then as now, by successive cell-divisions, corresponding
to the cell-divisions of Metazoan ontogeny, though they led to no single
multicellular adult. A stage further on, the corresponding cell-divisions
gaye rise to adherent colonies, fixed or free, arborescent or epithelial, each
type established by its own ontogeny. The very first, most ancestral
Metazoan of all—at whatever grade of evolution the dividing line may be
drawn—must be admitted to have been built up by a full ontogeny from
unicellular zygote to multicellular adult, so that, in the first, as in the latest
Metazoan, ontogeny eame first, leaving the first adult Metazoan as its original
achievement. The next generation, through a new ontogeny, produced a
second adult, and so on. Ina word, Haeckel’s causes and effects must be
inverted. Phylogeny (in Haeckel’s sense) is the product, the “record ”—
not the precedent cause—of successive ontogenies ; and neither the first, nor
the second, of Haeckel’s phrases can any longer express the basis of true
biogenetic law. Ontogeny does not recapitulate Phylogeny: it creates it.

* Of. Hertwig (1906, i. p. 56): “Die Hizelle z. B. eines jetzt lebenden Siiugetieres ist
kein einfaches und indifferentes, bestimmungsloses Gebilde....sondern.... das ausser-
ordentlich komplizierte Endprodukt eines sehr langen, historischen Entwickelungsprozesses.”

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 83

5. Does this involve, then, the loss of the doctrine of Recapitulation ?
To which question I reply, first, with Goethe, ‘“ Was fruchtbar ist, allein ist

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wahr,” and, secondly, as Balfour wrote on another matter: “If the above
position be admitted, it is not permissible to shirk the conclusions which seem

6*

$4 PROF. W. GARSTANG ON THE THEORY OF RECAPITULATION :

necessarily to follow, however great the difficulties may be which are
involved in their acceptance” (1885, ii. p. 32). But the “parallelism” of
ontogenetic and phyletic sequences, which was incorporated by Haeckel
in his “law,” was noticed by many a “good physiologist” before him
(cf. Meckel, Von Baer, L. Agassiz, &c.), and cannot disappear with his inter-
pretation of it. Perhaps now we shall see its true extent and meaning more
clearly. Ontogeny proceeds through successive grades of differentiation by
which layers, tissues, organs, and parts together with ordinal, family, generic,
and specific characters, are more or less successively established. As differ-
entiation increases, the combination of layers, tissues, organs, and parts
exhibited at successive stages resembles more or less distinctively the com-
binations characteristic of successive grades of evolution represented in our
schemes of phyletic classification. To that limited extent the ontogeny of
a given animal i$ an epitome of its phylogeny, and may be said, in the
true sense of the word, to recapitulate phylogeny, 7.e. to sum it up,
recall the main phases of it. This is the parallelism observed by Meckel,
Von Baer, and many others, expressed in evolutionary terms. It exists and
is undeniable.

6. This parallelism exists because phylogeny is itself the creation of
successive ontogenies, and ontogenies of necessity run parallel with one
another from zygote to adult. For ontogeny is the expression of zygotic.
power, the function of zygotic structure ; and zygotic change involves no.
radical departure froni the routine of ontogenetic method. One ontogeny is,
in this sense, a modification of its predecessor. The ontogeny which first.
established the Coelenterate grade was the basis of a later ontogeny which.
established the Coelomate grade. The life-cycle was extended accordingly,.
but never by the simple addition of a substantial unit or stage, distinctively
Ccelomate, to the final adult stage of a Coelenterate ontogeny. A house is
not a cottage with an extra storey on the top. A house represents a higher
grade in the evolution of a residence, but the whole building is rilioret—
Poandations timbers, and roof —even if the bricks are the same. You may
begin by building a cottage a little larger than its predecessor, cutting off
an entrance passage from the parlour, and adding a back kitchen ; but when
your ambition rises to an entrance hall, three reception rooms, two staircases,.
and so on, you are forced to a mutation in your building plans which affects
operations from the start. The ontogeny of a Coelenterate adds, in a certain
sense, on a simple diploblastic base, certain effective, workaday, adult.
features by which it copes with the conditions of its life; but the replace-
ment of these effective characters by others suited to the more adventurous.
career of a Coelomate (e.g. development of prehensile mouth instead of tentacles)
involves their disappearance altogether ; and there remains of Ccelenterate
organisation only that diploblastic residuum of differentiation out of which
the Coelomate may be economically and directly built up. Nor is the end of

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 85

the life-cycle alone effected. The Ccelenterate had a larva fitted both to
distribute it and to build it up. It must be changed so as to perform this
double function for a quite different creature, probably of very different size,
habits, and requirements. And then the changes at the larval end must be
fitted and co-ordinated with the changes at the adult end, so that every phase
of the life-cycle is modified in some way or other. Yet it is oniy this much-
pruned Ceelenterate sequence that survives as building material out of which
a specifically Annelid ontogeny may give rise to a Crustacean, and so on.
Inevitably the Coelenterate sequence in the Crustacean’s ontogeny is reduced
to the simplest terms, and is as far from “ mirroring” any functional Coelen-
terate type, or the original mode of its formation, as possible. Nevertheless,
the grades persist as stepping-stones from zygote to adult; and, having been
successively pruned of unessentials as they ceased in turn to furnish directly
the equipment of the adult stage, they have become very constant features
of the ontogeny in along line of evolutionary progress. Tor there is an
irreducible minimum beyond which even ontogeny cannot abbreviate. The
zygote is always unicellular, the larva multicellular and fitted for swimming,
and the adult a multilaminate complex of interdependent parts; and even
the Ctenophore, with its elaborate pree-organisation of the zygote, cannot
escape the rule that 8=4x2. Ontogeny repeats the necessary successive
grades of ancestral differentiation, but no ontogenetic stage is ever more
than an immature adumbration of a particular adult type in the phyletic
chain*. It reproduces those successive grades, not because successive adult
types have been included in it, but because each ontogeny is a modification,
within limits, of its predecessor; and by those predecessors the phyletic
chain of adults was organised and equipped.

7. Thus Ceelenterate, Coelomate, Protochordate, Gnathostome, and Tetrapod
are successive grades of differentiation both in the ontogeny and phylogeny
of a Frog; but at none of these grades does the ontogeny recall the form
and structure of a possible adult ancestor. This is obvious enough in each
of the first three grades ; and in the fourth, which is held to “recapitulate ~
the Fish, the tadpole lacks dermal skeleton (both scales and fin-rays), paired
tins, and biting jaws, which the adult ancestral Fish undoubtedly possessed.
The tadpole, in fact, is not a modified reproduction of an adult Fish-ancestor,
but a modification of the larva which that ancestral fish undoubtedly
possessed—still recognisable, in less modified form, in the larvee of Polypterus
and Dipnoi to-day. In other words, the life-cycle of the Frog is a modifi-
cation of the life-cycle of an ancestral freshwater Fish ; and adjacent terms
in the old life-cycle (larva and adult) have undergone parallel and correlated”
modifications, as well as some independent specialisations.

* Cf. Von Baer (1828, p. 230): ‘Der Embryo geht nie durch eine andere Tierform
hindurch, sondern nur durch den Indifferentzzustand zwischen seiner Form und einer
anderen.” ¢

86 PROF. W. GARSTANG ON THE THEORY OF RECAPITULATION :

8. That “recapitulation” does not require the reproduction of adult stages
n the ontogeny in order to be exhibited is plainly seen in the development
of many Geometrid moths. Hvyeryone knows the Geometrid or “looper ”
type of caterpillar, provided with prolegs only on two of the hindmost
abdominal segments (the 6th and last). This type is admittedly derived
from a prototype which possessed the full Lepidopteran equipment of prolegs
on segments 3 to 6, as well as the last, the prolegs on the three first segments
having subsequently disappeared. But many Geometrid caterpillars possess
vestiges of one or more pairs of these missing prolegs : in the March moth *
(Erannis escularia, Schiff.) there are traces of the last pair (South, ii. pl. 125);
in the common Brimstone (Opisthograptis cratwgata (Linn.)) and Scalloped
Hazel (Gonodontis bidentata, Cl.), clear rudiments of the last two pairs (I. c.
pl. 115) ; while the Orange Underwing (Brephos parthenias (Linn.)) has the
first two pairs rudimentary and the third pair fully developed and functional
(Meyrick, 1895, and South, 1908, ii. pl. 39). In the Feathered Thorn, Colotois
(Himera) pennaria (Linn.), the single pair of vestigial prolegs arises and
disappears between the Ist and 4th moults (Buckler, 1897). Now, the time
has long passed when it was possible to regard these prolegs as homologues
and derivatives of the true legs of some Scolopendroid ancestor. They were
“cenogenetic” larval features, adaptive interpolations, modifications of the
middle stages of a life-cycle which originally, in the earlier phases of Hndo-
pterygote history, exhibited larvee lacking prolegs altogether, asin Coleoptera.
Yet these examples of vestigial organs are as reminiscent of ancestral (though
larval) structure as the larval foot of the oyster, the larval stalk of Antedon,
the transitory feet of the parasitic Portunion, or any other of the familiar
examples that are held to prove the theory of adult recapitulation. They
demonstrate, as Morgan has already urged (1919), that recapitulation is
merely the static aspect of inheritance, and that, in this aspect, inheritance
is not primarily the reproduction of adult characters, but the reproduction
of the characters of each part of the whole life-cycle—the sequential
expression of the full train of zygotic potencies.

9. It may be urged that such hereditary changes in the middle phases of
the life-cycle do not affect the proposition that evolutionary changes usually
take place at the end, and that the case for adult recapitulation rests on the
evidence for this proposition. Nevertheless, to clear up the misunder-
standings of the past, it is necessary to leave no margin for ambiguity.
If “recapitulation,” in the special sense of partial reproduction of the past,
is hereby shown to be independent of the characters of adult ancestry, that is
something gained: the axe is laid at the root of the tree. For much of the
glamour of the old biogenetic law is due to its appeal to such idols of the
market-place as the assumption that “like begets like,’ and that, as adults

' #* Meyrick’s classification and nomenclature are followed here, but the English names
have been added for convenience of reference to South’s figures.

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 87

only are capable of begetting, the thing begotten is built up of successive
adults. That ‘“cenogenetic” interpolations *, without any adult ancestral
significance, are a normal feature of almost every life-cycle, is often verbally
admitted, though the recapitulationist rarely realises how profoundly such
interpolations may affect his phylogenetic conclusions, and how dangerously
subtle become his arguments when engaged in proving that such larve as the
Trochosphere, the Nauplius, and other distributive stages of the life-cycle
are at bottom “recapitulative.” I propose to deal more fully with the origin
and significance of such larval forms in another communication ; but, to
illustrate the principle, I may add one example of the origin of an inter-
polation in the most progressive of sequences known to us, viz. that of
Vertebrates. There is no doubt that Birds are descended from Reptiles.
It is beyond question that Reptiles are hatched in a form and with a somatic
organisation which is that of a miniature adult Reptile in all respects. Yet
the Bird is hatched in a form and with sundry details of organisation
different from those of the adult, e.g. its downy plumage. Now, the
“typical”? down-feather is an open hollow tube, splayed out at its free
extremity into a ring of soft barbs (or barbules) of equal size, and I ask if
such a tubular feather is to be regarded as an intermediate stage in the
phyletic derivation of feathers from scales. Isukmit that there is not ascrap
of evidence, or of probability, that any adult ancestor of Birds, along the
whole route from Reptiles to Sparrows, was ever clothed in anything except
scales, feathery scales, and finally contour-feathers. The chick is an inter-
polation in the life-cycle of Birds, and its down is a ‘“‘ secondary” modif-
cation of complete contour-feathers. The Duck, the Fowl, and the Pigeon
represent three successive grades of differentiation in the phylogeny of
Birds. Anyone who will examine under a microscope the nestling-down of
these three birds in the order mentioned, will see that they exhibit successive
phases in the degradation (a) of the primitive rachis of a contour-feather
and (b) of the barbs of such a feather, 7.e. that the chick stage, with its
peculiarities of organisation, has been evolved, step by step, within the group
of Birds alone, and is an interpolation that has no relation with, and throws
no light on, the pre-Avian adult ancestry, or on the way in which scales
were transformed to feathers +.

* H.g. Weismann’s discussion of the evolution of markings in larvee of Sphingidz, much
of which is probably sound, though unnecessarily complicated by the assumption that
primitive longitudinal markings have been ‘‘shunted back” into earlier stages of the
ontogeny, instead of being simply replaced in the later stages by patterns more suited to
increased size or special conditions of exposure (1904, pp. 177-185).

+ The subsequent publication of Prof. Cossar Ewart’s valuable paper on “The Nestling
Feathers of the Mallard” (P. Zool. Soc. 1921) renders this discussion inadequate and I hope
to amplify it. In the meantime I would merely remark that, on the relation of feathers to
scales, the association of several feather-germs with single scales on the foot of the Owl is

no disproof of my thesis, since the feathers here represent a secondary extension, like that
of the scales on the head of Ceratodus.

88 PROF, W. GARSTANG ON THE THEORY OF RECAPITULATION :

10. Let us now take an equally unambiguous case of evolutionary change
at the adult end of the life-cycle. There is a group of genera of Geometrid
moths in which the life-cycle terminates with normal winged males and
more or less wingless females (Apocheima, Hybernia, Theria*, &e.). It is
un adaptive change, for, unlike their congeners which hibernate as pups and
emerge in early summer, these moths emerge in winter when the trees are
leafless, and the normal method of repose is much more dangerous. The
males, like both sexes of other related genera, rest by day exposed with
wings outspread on tree-trunks, palings, &e. The wingless females
hide in the crevices of bark. Both are active at night, the females
creeping out of their crannies, and the males hunting for them up and
down the woods. In the Early Moth (7. rupicapraria) and Dotted Border
(H. marginaria), which emerge in February and March, the wings of
the females are half as long as the body (South, ii. pl. 120); in the Scarce
Umber (4. aurantiaria), which emerges as early as October or November,
the wings are mere stumps (J. ¢. pl. 120); while in the Mottled Umber (H.
defoliaria) and various other species the wings are completely lacking (J. ¢.
pl. 122). The wings of the males are of full size throughout (Meyrick, and
South, J. c.). -Now here is a case of evolutionary change of the adult form,
and in one sex only; but, with these facts before him, and with our know-
ledge of the origin and breeding of similar mutations in Morgan’s Drosophila
experiments, who can assert that this abnormal adult has been added to the
life-cycle of its normally-winged ancestors, and that the old adult has been
“pushed back” to an earlier phase of the life-history. The wingless female
is the exact counterpart of the normal male, and, though I do not know if
any change has already taken place in the pupal characters of the female, it
is a safe deduction from our knowledge of the pupal condition in more
extreme cases to assert that the only changes likely to ensue will be in the
direction of still further reducing the size of the pupal wings. The ontogeny
will be influenced in the direction taken by the new adult, and without regard
to the ancestral adult at all. The new adult is just a modification of the old
adult. There is no addition, no “tacking on” of a new stage ; no “ pushing
back” or “tachygenesis” of the old adult stage—merely a substitution of
one adult type for another, and, sooner or later, some correlated changes in
the stage which immediately precedes it. Zygotic mutations have caused the
changes; natural selection has controlled the breedings of successive
generations ; and heredity has perpetuated the results of the selection.
Certain ancestral adult characters are disappearing from the ontogeny ; and
the condition of a flea, ontogenetically, as well as finally, without a trace of
wings at any stage, is likely to be the end result.

11. I have selected this example, not because it is representative of all
evolutionary changes that manifest themselves in the final stages of onto-
geny, but because of its bearings on the most recent exposition of the theory

* Meyrick’s nomenclature (1895); English names and figures in South (1908).

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 89

of recapitulation from the embryological side. I have already shown that,
in its original and general sense, recapitulation is a fact which was recognised
long before there were any theories to account for it. But this generalised,
or Meckelian, recapitulation needs to be clearly distinguished from the
specifically adult recapitulation of Haeckel and his school, who could not
understand the origin of the former except on a theory of catenary ancestral
inheritance, each term in the ontogeny (except the last) being moulded after
_the likeness of a specific adult ancestor—though, of course, condensed,
abbreviated, telescoped, and secondarily modified by adaptive changes. Now,
the only way that I can see of establishing this theory by purely embryo-
logical methods, is to show that the penultimate stage of the ontogeny of
a given type of adult resembles the final (adult) stage of the ontogeny of
some theoretically ancestral type more closely than it resembles the corre-
sponding penultimate stage of the same, and similarly with regard to the
antepenultimate stage, and so on. I cannot find that this has been done, or
even attempted, in any case—certainly not in any of the cases recently
selected by MacBride for discussion. Yet this is his thesis: *‘ When we
assert that a Metazoan recapitulates in its life-history the past history of the
race or stock to which it belongs, we mean that the stages intervening
between the egg and the adult form resemble in some of their prominent
features the adult animals which belonged to the same stock at different
epochs in the past history of the race” (1917, p. 425) ; and he is concerned
to show both that the adult stage of the ontogeny of a new species is an
addition to the ancestral ontogeny (1914, pp. 23, 650), and that the adult
stage of the ancestral ontogeny is reproduced (“recapitulated”) in the
ontogeny of the new species as the last larval (or “neanic”) stage (J. ¢.
pp. 21, 22). But his method of establishing these points is merely to select
a number of cases in which the adult deviates considerably from the normal,
and to show that “the young form resembles the type of the order to which
the parent-belongs and not the parental type itself” (1917, p. 428).
“Thus the young Hermit-Crab swims freely about in the water and has a
symmetrical abdomen like that of Shrimps and Prawns” [but so have the
young stages of these creatures!]; “the young Flatfish swims with its
ventral edge down and its dorsal edge up, and has an eye on each side of the
head” [but so have the young of all Teleostei!] ; “the young Comatulid
is fixed to the bottom by a stalk like other Crinoids [and their young too, in
all probability !]; and the young American Oyster possesses a foot like that
~ of other bivalves by which it crawls about” [and, I may add, as the young
of nearly all other Lamellibranchs crawl about! ]. Nowhere does he show,
or claim to show, that the young stages of any of these animals resemble
the adult more closely than the young stage of typical members of their
respective orders. He does not show it because he cannot. In every case
that he discusses, whether the above, or the cases of the parasitic Portunion

90 PROF. W. GARSTANG ON THE THEORY OF RECAPITULATION:

and Actheres (1914, pp. 22, 206), the young resemble the young stages far
more closely than they resemble the adult stages of their respective “normal”
relatives *. The symmetrical larva of Pleuronectids is scarcely distinguishable
from many Teleostean larvee of other families : it is distinct from any existing
or fossil adult Teleost. It is the adult Oyster which has lost its foot, not the
young Oyster which has acquired it. Itis the adult Portwnion which has
lost its legs, not the young Portunion which has acquired them by tachy- (or
any other kind of) genesis from its adult ancestors! These cases are all in
the same category as the case of the wingless Moths already discussed. No
new stage has been added to the life-cycle. One adult stage has been trans-
formed into another, but the penultimate stages remain as before. The
protagonist has missed his point, and the riposte is obvious. It was not his
task to prove that Oysters were Mollusks, that Hermit-Crabs were Crustacea,
or Pleuronectids Fishes. Comparative Anatomy did that long before the
science of Embryology staked its claim. His province was to show that by
virtue of Haeckel’s Biogenetic. Law he could reconstruct the prominent
features of an adult ancestor from a developmental stage. All he has done
is imperfectly to confirm Von Baer’s pree-Haeckelian doctrine, that animals
resemble one another more closely in their young stages than in their adult
stages t. For his own illustrations show how greatly the adult may differ
from the larva. He has merely shown the resemblance between the larvze
of a given class. It follows that, for all he has shown to the contrary, the
“typical” or “normal” larvee, which the Pleuronectid larva resembles,
might have grown into Cod, Mackerel, or any other type of Teleost, and
that the adult ancestors of Pleuronectids, so far from being “ normal,” may
have carried themselves upside down like a Remora, or stood on their tails
like Pipefishes. If no more relevant evidence than this is forthcoming,
I claim that the old Biogenetic Law of adult recapitulation is dead, and that
Morphology is henceforth free from a delusive and cramping hypothesis.
Ontogeny is not a lengthening trail of dwarfed and outworn gerontic stages.
Youth is perennially youth and not precocious age.

12. It is true that ontogeny could not exhibit its normal progressive
differentiation of structure if evolution had always been of the type exhibited
by these examples of metamorphic Insects, Lamellibranch Mollusks, parasitic
Crustacea, and Pleuronectid Fishes. Hvolution within these groups to-day
partakes mostly of the nature of an adaptive radiation of the various types,
whereas the general lines of ontogeny correspond rather with that kind of
evolution which involves morphological and physiological progress.
Although a detailed examination of any of these various advances falls

* Note especially the absence of the 8th pair of thoracic limbs in Epicarid, as in all

other Isopod larvae.
+ “Im Grunde ist also nie der Embryo einer hoheren Tierform einer anderen Tierform

gleich, sondern nur ihrem Embryo” (1828, p. 220).

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 91

outside the purview of the present communication, the slightest survey of
Vertebrate evolution shows a series of triumphs over limiting environments
of medium, temperature, space and time which has been based as much on the
substitution of new for old organs as on the continuous elaboration of
particular ones. When a given organ is wholly transformed in the course
of evolution, it rarely shows traces in ontogeny of the original steps of
its transformation (e.g. bony scales to fin-rays, horny scales to feathers,
lobate fins to pentadactyle limbs, pentadactyle limbs to wings). The final
form alone is inherited and develops directly. But when originally separate
organs are ultimately united into one organ, some stages of the process of
amalgamation are necessarily repeated (e.g. branchial arches to hyoid,
vertebral elements to vertebrae, muscle-buds for paired limbs, anchylosis of
limb-bones, segmental tubules of kidney, &c.). And when a new organ has
arisen in intimate dependence on an old organ, the old organ may still
remain necessary for the development of the new (Kleinenberg, 1886).
Thus backbone replaces notochord, and bone replaces cartilage in present as,
doubtless, in past ontogeny, for the former organ or tissue is still necessary
as scaffolding for the later one ; and the constant development of gill-slits in
the ontogeny of terrestrial Vertebrates is but another illustration of the same
phenomenon, as Sedgwick has already pointed out (1894)—for a complex
double circulation that has been elaborated along channels determined by a
branchial cireulation cannot readily depart from the phyletic steps of its
formation. It is this formative dependence of one organ, or set of organs,
on another that confers on Vertebrate ontogeny its marked recapitulative
character.

13. But it is equally clear that the whole succession is explicable without
recourse to the theory of successive adult incorporations, and that the onto-
genetic stages afford not the slightest evidence of the specially adult features
of the ancestry. So far as notochord and gill-slits are concerned, they make
their appearance in the earliest larval stages of every animal that presents
them, including Amphiowus itself. Their phyletic origin is still wrapped in
obscurity. The case is hardly different as regards cartilage, bone, scales,
feathers, hairs, lungs, limbs, and all the other organs concerned. No example
can be adduced of any of these organs arising in an adult stage of ontogeny.
Until that evidence is produced, it is idle to claim that recapitulation which
involves any of these organs is a repetition of specifically adult ancestral
features: Moreover, it is impossible to overlook the fact that some of the
most pregnant changes in the characteristics of the higher Vertebrates are
directly or indirectly traceable to changes in the earliest stages of the
ontogeny. The elaboration of the brain in Birds and Mammals, and the
development of their social and esthetic senses, are connected with the inter-
polation of the helpless chick, puppy, or baby stage in the ontogeny, which
from the simplest beginnings has led to the development of educability and

92 PROF. W. GARSTANG ON THE THEORY OF RECAPITULATION :

preferential action in place of the limited range of reflex and automatic
mechanisms of more primitive types. That “little twist of brain,” which dis-
tinguishes one philosopher from another, is not more striking in its effects
than are those trifling touches to the structure of the heart which transformed
the cold-blooded Reptile and Stegocephalan into the warm-blooded Bird and
Mammal respectively. Yet these are changes which, however graduated
through successive generations at the outset, were not of a character to have
been completed, or even initiated, in any adult stage of ontogeny. They must
have been first manifested as a series of embryonic mutations, subjected
continuously to selective tests of their relative physiological efficiency. Age
bears the buffets of the world, but youth regenerates it.

14. It is, however, the paleontologists who are the real defenders of the
Biogenetic stronghold. With them the Law is a faith that inspires to deeds,
while to the embryologist it is merely a text for disputation. The difference
is striking and worth defining. When the embryologist sets up his larval
images and worships them as pree-Cambrian ancestors, the real ancestors
cannot be produced to demonstrate his folly. But the paleontologist’s aim
is to trace lineages directly, and he is not satisfied until he has produced his
ancestors, or at least the most substantial remnants of them. I confess that
I have been tempted many times before to-day to attack a theory which has
led so many of us into blind alleys, but always Hyatt’s Ammonites recurred
to present an unanswered, and seemingly unanswerable case for Haeckelian
recapitulation. A priori it seems absurd that senile characters should be the
beginnings of a line of evolution (Hyatt, 1897, p. 221 &e.), but the for-
midable array of evidence, the wide range of unfamiliar material to be studied,
and, not least, our ignorance of the habits and conditions of life of this type
of Mollusk, have all conspired to render these Ammonites to me a real obstacle.
The following case, however, has reeently impressed me with its remarkable
analogies, and justifies me, I hope, in presenting a general argument without »
directly tackling the Ammonite problem itself, at any rate for the present.

The curious Prosobranch Gastropod Lamellaria, which mimics and devours
Compound Ascidians, produces veliger larvee of a unique type known as
Echinospira (Krohn, 1853, 1857). The hyaline shell first produced is dilated
so that it is far larger than its oecupant—resembling in this respect the
gelatinous house of an Appendicularian. It is coiled like the shell of an
Ammonite, being in some species discoidal, with perfect symmetry, in others
spiral, and in the related Onchidiopsis more simple and sac-like (Bergh, 1887).
The larva can withdraw himself completely, or, with his mantle-edge clasping
the mouth of the shell, he can protrude a large 4- or 6-lobed velum, and
swim about with it on his excursions with wonderful grace and ease. The
mouth of the shell is regularly extended at its margin, the successive
additions being marked by transverse lines of growth and generally by one
or two pairs of longitudinal (spiral) rows of tubercles or spines as well.

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 93.

Inside this rapidly growing house (the roof of earlier whorls being periodically
destroyed to make room for their successors), the larva secretes a second
flatter shell (the rudiment of the adult shell) which fits his visceral hump.
and adheres to it, but is temporarily fixed in the cavity of the outer shell
until he shifts his position, when it is carried forward—as though he were
trying to improve on an Ammonite’s arrangements by the device of a
portable septum *. Now, the point of the analogy is this: that the spiral rows.
of tubercles on the outside of the shell are variable in different species, and
that in the same individual they may go through a cycle of changes exactly
like the progressive changes of an Ammonite. The shell may pass through
a smooth stage, a unituberculate stage (the outer row of tubercles), a di-
tuberculate stage (with both rows developed), a spiny stage, and lastly a
ribbed stage, in which cross-ribs join the tubercles of the two rows together.
Unfortunately for the completeness of the analogy, Hehinospira does not (so.
far as I have yet seen) present a gerontic stage, for, being only a larva, and
usually very lively and vigorous, he quits his cage before old age comes.
over him, and transforms himself into a torpid Ascidian-eating Lamellaria.
I hope to publish shortly some figures of the remarkable process of meta-
morphosis, of which I was lucky enough to be an eye-witness last year at
Plymouth, as well as someé further details of the growth of the larval shell ;
but for my present purpose I refer to Simroth (1885, text-fig. 5; Taf. xvi.
figs. 1 & 2; xviii. figs. 1-2, 6-8), whose excellent figures sufficiently
illustrate my immediate points. Meanwhile I submit (1) that the characters.
of the larval shells of Lamellaria and its allies are purely cenogenetic, with
no relationship to the characters of any adult ancestors; (2) that gradual and
progressive changes in the shell of the same individual, from one type of
“ornament” to another, oecur regularly, and are apparently determined by
the constitution, size, and vigour of the larva under the particular conditions
of its existence ; and (3) that different degrees of the power of tubercle- and
spine-formation characterise the larvee under different conditions of existence.

In this case, from which all specific influence of adult ancestry is excluded,
there is no escape from the conclusion that the power to perform these
variously graduated operations, and the extent of that power, are essentially
functions of the zygotic constitution, though there is a considerable margin
for the direct influence of conditions. I conclude that, if this is so for the
“cenogenetic” larvee of Lamellaria, it is not likely to have been different.
for the “ palingenetic ” stages of Ammonites.

15. I return to the keynote with a direct comparison between the ontogeny
and phylogeny of an animal in which the skeleton has been an important
index of racial structure throughout geological time—the Crinoid Antedon.

* The two shells correspond to the two dayers of an ordinary Molluscan shell, dislocated
from their original union, The outer or larval shell, corresponding to the prismatic layer,

is formed by mantle-edge alone; the inner or adult shell, corresponding to the nacreous.
layer, is formed by the visceral surface of the mantle alone.

94 PROF. W. GARSTANG ON THE THEORY OF RECAPITULATION :

In selecting this type from the few that fulfil the necessary conditions, I have
naturally not overlooked the fact that in the two most doughty advocates of
Haeckel’s law in this country, we also possess two’of the foremost experts in
Hehinoderm embryology and Crinoid paleontology respectively. If I err in
my selection, or statement, of facts to be brought into prominence, they will
know, I think, that it is from inadvertence and not from intention. The
test of paleontology cannot, of course, be brought to bear on the origin of
the stalked condition, or at present, at any rate, of the primal torsion of
the internal organs of Echinodermata; but I regard it as established by the
form-sequences which Bather and his colleagues have traced, that radial
symmetry was imposed upon the skeleton of an original pear- or sac-shaped
body by the extension of superficial food-grooves leading to the mouth from
food-collecting tentacles—a view which I understand is shared by MacBride
(1911, p. 248). The hypothesis that Cystoid, Blastoid, and Crinoid were
successive and independent offshoots from an unknown stock that lacked a
skeleton seems to me to involve the negation of precise morphological evidence.

Accepting as my basis Bather’s masterly sketch (1900) of the phyletic
classification of these groups, and bearing in mind his own cautions (J. c. p.138),
as well as the slenderness of the geological record of Permian and Triassic
forms, the main outlines of the adult ancestry of Antedon cannot have
deviated far, I think, from the following sequence (the Roman numerals in
brackets refer to certain figures of special significance in Bather’s work) :—

(?) Pree-Cambrian.—Pree-brachiate ancestors, first Cystoid, with numerous
irregular thecal plates, then reduced and approximating to Blastoid
regularity. Finally an immediate ancestor of Cysto-Blastoid structure
exhibiting an ill-defined separation between calyx and stalk (cf.
Cystoidea, viii., xviii.), but with fixed pentameral symmetry and com-
position of the firm cup, as in Blastoids*, from the Basals upwards
(Stephanocrinus, ii.). From such an ancestor, after development of
arms, the Monocyclica and Dicyclica, distinguished at first only by
the exclusion, or inclusion, of Infra-Basals in the cup (7. e. the position
of the growth-zone), diverged. I pursue further only the Dicyclic
series, and neglect the Camerata..

Cambrian.—The primitive Inadunate: 5 simple arms, distinct from the
cup ; disk firmly plated with 5 Deltoids (Orals), supporting ambulacra
above their conjoined edges (cf. Hybocrinus, xxxvi.). Slightly
modified, this type survived among Ordovician Cyathocrinoids
(Porocrinus, lxxxvi.): Anal plates (X & RA) present in the circlet of
Radials; Posterior Oral, the only madreporite.

* The stereotyped monocyclic constitution, sharply separated stalk, and late geological
development of Blastoids suggest that this group may be composed, in reality, of peedo-
genetic Crinoids, and it would be worth while to extend this hypothesis to some of the
Cystids themselves.

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 95

Ordovician.—Dendroecrinoid modification. Arms still non-pinnulate, but
dichotomously branched ; the disk now flexible by interpolation: of
small plates between Orals and Radials, but Ambulacra remain supra-
teominal. Anals as before, supporting a great anal turret (xxvi.).
Madreporite lost.

Silurian.—Dendrocrinoid arms forked and pinnulate (Dendrocrinus, iii.).

Dendrocrinoids lose RA, and then X from cup (Graphio-

crinus and Hrisocrinus,c) ; their arms become biserial ;
Infrabasals reduced and covered by stem.

Devonian.
Carboniferous.

Trias.— From Graphiocrinidee arise Pentacrinidee : cup (patina) shallow,
the disk bulging up between the arms; no persistent proximal
columnal as yet; stem cirriferous, its ossicles changing from round
to pentagonal, with petaloid furrows and radiating ridges ; no Anal ;
arms (again?) uniserial. Various members of the family swim about,
and re-anchor themselves by distal cirri of stem. Obscurely leading
to Flexibilia Pinnata, but exact links missing (cf. Bathyerinus, with
bi-fasciate stem-ossicles, like Bourgueticrinide and Antedonide,
figs. xlix., exv.).

Jurassic.—The first “ Pinnata,” with persistent proximal columnal and
reduced Basals; stem-ossicles of modified Pentacrinid or Bathycrinid
types. Adillericrinus (no cirri) broke away from bottom of its stalk
for swimming, the stem being slowly absorbed (lii.).. Thiolliericrinus,
the first Antedonid, with cirriferous compound Centrodorsal, fairly
stout stem, and bifasciate joints of stem-ossicles.

Lias.—-Antedon and later types break away from top of stalk early in life.

With this sketch of the adult ancestry of Antedon before us, let us now
see how it is recapitulated in the ontogeny. Certain sequences of form-
change take place in the same order. The larva on fixation exhibits
successively an armless (‘‘Cystid”) stage, and stages with simple arms,
forked arms, and pinnulate arms respectively ; the Oral plates at first are
co-extensive with the disk, as in Blastoids * and Cyathocrinoids, and then a
peripheral growth-zone (perisome) is established between them and the
Radials as in Ordovician Dendrocrinoids ; an Anal plate appears within
the circlet of Radials, rises above it and disappears, as in Carboniferous
Dendrocrinoids ; the Infra-Basals arise separately and then fuse with one
another and the proximal columnal, marking the change from Dendro-
crinoidea to Flexibilia Pinnata.

* Another Blastoid or pre-Crinoid relic appears to be involved in the abortive attempt
of the 5 primary tentacles to branch and grow as radial canals in the vestibulate stage, each
peristomial tentacle (in spite of its lack of food-grooves and skeleton) representing a Blastoid
brachiole. If Blastoid brachioles are homologous with Crinoid pinnules, the view that
‘simple unbranched arms are primitive in Crinoids becomes untenable.

96 PROF, W. GARSTANG ON THE THEORY OF RECAPITULATION :

Now note the discrepancies. Except possibly in the anal interradius,
there is no trace of pra-Cambrian irregularity in the number and arrange-
ment of the skeletal plates, of the earlier acquisition of radial symmetry by
the ambulacra and its later imposition upon the plates of the calyx; no sign
of the derivation of the stalk by constriction of a pyriform base ; no evidence
of the oral plates having originally formed a solid disk, above the sutures of
which the ambulacra ran. Except for certain additional dislocations to be
referred to in a moment, it is just as in the development of the skeleton of a
Vertebrate limb: the number of the skeletal elements is fixed from the
beginning (even the pattern of the stem-joints) and ontogeny reveals no signs
of their past history—with two exceptions: the migrations of the Anal plate
and the composition of the Centro-dorsal. The former is a precious record
of the change exhibited by the ancestral Dendrocrinoids, when, as arm-
structure changed and flexibility increased, the diminishing anal chimney
(fig. iii.) no longer required a buttress in the calyx wall to support it. But
its retention in the ontogeny of Antedon is no proof of the normality of so.
precise a record of ‘ancestral change: rather is it the exception which proves
the rule of absence of such records. It is comparable with the “useless”
notochord of the Vertebrate embryo. It has no part to play in the adult,
because, as growth proceeds, the bases of the arms take over the main support
of the body; but in the larva the patina is the sole support, and, as an Anal
plate (and a Radianal as well) was a constant inherited element of the cup.
through nearly the whole of Paleozoic time, it is scarcely surprising that it
should be retained in that part of the life-cycle where it is still conceivably
useful. On the other hand, it is squeezed out of the cup as soon as the anal
tube, by remaining small, withdraws any demand for its retention, and when
the flexible incorporation of the five arm-bases in the eup sets up a counter-
demand for strict pentameral symmetry. This demand, so far as the Radianal
is concerned, has long since been met by the complete elimination of
that plate from the ontogeny. ‘One thing at a time” is nature’s rule.
Similarly the ontogenetic history of the Centrodorsal is a physical necessity
if one plate is to be made by the amalgamation of a number (¢/. development
of vertebrae, pore-plates of Hchinus, &e.); and the other recapitulative
features of the ontogeny (arm-development &c.) are examples of other
necessities of differentiation, since you cannot get 2, except by duplicating 1.
Adult recapitulation demands that the arm-branches should extend to the
full length of the arms (as in Cyathocrinoids) before they reduce themselves
alternately to the dimensions of pinnules. This they do not do. They take
the shortest route to their goal, so far as constitution, not ancestry, will
allow them.

But these discrepancies with phylogeny are trifles beside the phenomena
of development of the Ora] and Radial plates. In the whole series of adult
ancestors from Cambrian times to the present, not one possessed Oral plates

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 97

which rested on the Basals ; nor, if we overlook this developmental modifi-
cation, did one of those ancestors in its adult condition ever possess Orals
which were hinged to the Radials on the edge of the cup, capable of opening
and shutting over the entire disk like the valves of a trap-door (Bather,
fig. xxxiii.; MacBride, 1914, figs. 408-110). Where does the sure, frail
“ Ariadne-thread”” conduct us now? If no fossil evidence were avail-
able, anyone who should attempt to reconstruct the ancestral Crinoid on
the common embryological assumption that the stalked larva of Antedon
represented an ancestral adult stage would go inevitably astray, as many
have done already in spite of paleontological knowledge*. For the
remarkable thing is that several existing Crinoids possess an arrangement of
oral valves in the adult precisely or closely similar to that of the larval
Antedon (Lolopus, l.c. xxxiv.; Hyocrinus, Sedgwick, 1909, fig. 209; Thau-
matocrinus, Carpenter, 1884, pls. iii., vi., lvi.). The relations of the oral
plates in <Antedon to the vestibular roof of the larva, as well as the
temporary suppression of Radials, are clearly ‘“‘cenogenetic” features.
If a vestibule was a feature in the development of the earliest Crinoids,
the oral plates must have been deposited beneath its floor, and not in
its roof. Their relation to the roof (which alone enables them to split
apart and function as valves) is an embryonic mutation. It is, therefore,
scarcely open to doubt that the condition of the oral yalves in the adult
Holopus (and Hyocrinus?) is due to the retention of a feature that was
purely. embryonic, not adult, in origin, and that, in this respect, these
interesting Crinoids are as “‘pedomorphic” as any Perennibranchiate
Amphibian.

This brings me to my last point. When the common argument is urged
that the stalked larva of Antedon “recapitulates” the adult stage of its
stalked ancestors, it seems to be forgotten that every type of Pelmatozoan,
from pree-Cambrian Cystids to the present time, must also have possessed a
tiny fixed stage of simple structure following a free-swimming larval life,
and that the main features of the skeleton must have been laid down in that,
or a still earlier, stage of its ontogeny. It follows from what has been said
that the modern Pentacrinoid larva of Antedon is a modification of the
corresponding stage of the ancestral ontogeny, not of the adult stage, and
that the adult Antedon is not an addition to the ontogeny of any preceding
Crinoid, but just a modification of the adult phase of the same ontogeny—
partly by loss (e.g. anal turret and plates, Oral plates, stalk, &.), partly by

* P. H. Carpenter (1884, p. 145) compared the vestibulate condition of Antedon with
the Camerate condition of Haplocrinus (Bather, xxxy.) and the Platycrinide (/.c. xl.). It
is a tempting suggestion, especially as the Carboniferous Platycrinide possessed bifasciate
oval stem-ossicles. But these types are Monocyclic; the relations of the “Orals” are only
superficially similar, and it is very doubtful if these plates are other than enlarged “ proxi-
mal ambulacrals”’ (see Bather, pp. 127-129, and fie. xli.).

LINN. JOURN.—ZOOLOGY, VOL. XXXV, 7

98 PROF. W. GARSTANG ON THE THEORY OF RECAPITULATION :

elaboration of pre-existing structures (arms, stem-ossicles). Nature does
not build up a new type by addition or abstraction of “stages,” but of
organs, or parts of organs*. Moreover, when she makes a change, she does
not do so by altering these organs, or parts of organs, when fixed or rigid,
but when plastic and growing. In particular cases this may be late in life,
but it is not usually so, and it is not likely to have been so with respect to
the patinal skeleton of the Crinoids under consideration. The Anal plate
within the circlet of Radials is a feature inherited from earliest Silurian
ancestors. It is claimed to “recapitulate” an adult feature of those
ancestors. I submit that no Anal or other plate was ever interpolated within
the patina except in the formative stage of growth when the Radials them-
selves were loose and unsutured. The first Anal plate that entered the
Radial cirelet from the disk (if that was its origin) must have done so as the
result of an embryonic, not an adult, mutation. Once let the towering anal
chimney of an adult Dendrocrinoid (cf. Bather’s fig. iii.) effect a breach in
the wall of the patina, and*the whole cup would split asunder. Nature
underpins when it is safe to do so. She usually builds the foundation first
and the superstructure afterwards.

16. The following summary, omitting illustrative detail, recapitulates in
closer logical sequence the chief points of this attempt to re-define the
foundations of Morphogenetic Law.

RECAPITULATION.

I. Ontogeny is the sequential expression of zygotic powers of cell-division
through simple to complex grades of cell-grouping and differen-
tiation.

II. Phylogeny is the procession of ontogenies along a given phyletic line
of modification. It is expressed in terms of adult structure, but
the zygotes of successive ontogenies have also undergone a parallel
elaboration of nuclear or cytoplasmic structure, or of both, which
determines the sequence of the ontogenetic form-changes.

III. The phyletic succession of adults is the product of successive onto-
genies. Ontogeny does not recapitulate Phylogeny : it creates it.

IV. An individual confronts the world before his ontogenetic processes
are completed, and often at a very early period of his life-cycle.
Only those individuals reproduce who have survived the ordeal of
larval conditions. Adaptation of the larva accordingly plays a
prime part in determining the modification of successive ontogenies.

* Cf. Weismann (1904, ii. p. 174): “.... itis impossible to compare a particular stage
in the embryogenesis of a species with a particular ancestral form, Only the stages of
individual organs can be thus compared and parallelized.”

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 99

V. The life-cycle is extended, not by addition of a new adult stage at the
end of the old adult stage, but by further differentiation of organs
or parts of organs. Old adult characters are eliminated from the
ontogeny unless required as temporary bases for the new characters.

VI. -As the individual, through all the form-changes of his life-cycle, is an
evolutional and functional unity, modifications manifested in his
larval or adult phases involve co-ordinating changes in the more
passive and formative phases (embryonic, post-larval, pupal stages).

VII. Thus, while a given ontogeny, under normal conditions, tends to
repeat the form-sequences of its predecessors, it is liable to changes
in every part of the life-cycle—positively, by equipping the larval
and adult stages for the changing conditions of their various careers,
or with greater efficiency for the same conditions, and negatively,
by abbreviating the formative processes to the uttermost.

VIII. The idea that form-changes in ontogeny were preceded by similar
changes in adult ancestry is an illusion, since adult Metazoan
ancestors never directly gave rise to their successors, but to gametes ;
and these, blended with other gametes, were the real heralds of
successive ontogenies. Plainly the first Metazoan was not produced
by a Metazoan. He was the result of a Protozoan ontogeny, the
tour de force of a genius among Protozoan zygotes. The first Bird
was hatched froma Reptile’s egg. We can speak of earlier and
later, original and modified, ontogenetic processes ; but the possi-
bility of a distinction between ancestral and ontogenetic processes
is out of the question. All changes are ontogenetic.

IX. In the same way the contrast between “ palingenetic ’
adult ancestry) and ‘‘cenogenetic” (foreign or non-repetitive)
characters, which was originally based by Haeckel on an assumed
hereditary difference between adult changes and embryonic adap-
tations, has lost its significance. Both types of character were of
ontogenetic origin, and equally hereditary, but the one set arose
earlier in the phyletie history than the other. Morphology will not
recover exactitude of outlook until it is entirely freed from the
hypnotic influence of Haeckel’s terminology. I propose in future
to use palwogenetic and neogenetic when referring to ontogenetic
processes, and palwomorphic and neomorphic when contrasting
primitive and moditied types of structure.

X. There is a general correspondence between the successive grades of
differentiation in ontogeny and the successive types of organisation
which characterise the steps of phyletic progress (Meckel’s
law). This general correspondence exists because each series—the
ontogenetic and the phyletic—was preceded and caused by the same

Ra

r)

(repetitive of

100 PROF. W. GARSTANG ON THE THEORY OF RECAPITULATION :

phylogenetic series of ontogenies. The outcome of each successive
ontogeny was an adult representative of one of the successive types
of organisation. The last ontogeny of the whole series is the one
under consideration. Inevitably there is recapitulation of sueces-
sive grades of differentiation, but repetition of adult ancestral
stages is necessarily and entirely lacking. Ontogeny is not an
animated cinema show of ancestral portraits; but zygotes may be
likened to conjurers playing the old tricks for the most part, and
oceasionally opening a surprise packet—nor do they always keep
their novelties back until the end of the performance, as Antedon
and Holopus bear witness.

In other articles I propose to deal with the origin and significance of larval
forms, and to draw attention to some further examples of the influence of
larval characters upon adult organisation, to which I apply the term ‘‘ Peedo-
morphosis.”

LITERATURE.

Barr, C. E. von. 1828.—Ueber Entwickelungsgeschichte der Tiere, i.

Batrour, F. M. 1885.—Comparative Embryology.

Bateson, W. 1894.—Materials for the Study of Variation. Macmillan.

Batuer, F. A. 1893.—The Recapitulation Theory in Paleontology. Natural Science, ii.
pp, 275-281 (ef. Hurst, enfra).

Batuer, F. A. 1900.—Lankester’s Treatise on Zoology, iii. Echinoderma.

Barner, F, A. 1920.—Presidential Address to the Geology Section, British Association.
Cardiff Meeting.

Brereu, R. 1887.—Die Marseniaden. Semper’s Reisen, ii. 2 Suppl. p. 157.

Buckier. W. 1891.—The Larve of British Butterflies &c. vii. p. 8 (Ray Society).

CarpPEntER, P.H. 1884.—Report on the Crinoidea. ‘Challenger’ Reports, xi. pp. 145 &e.

Dourn, Anton. 1875.—Der Ursprung der Wirbelthiere u. das Princip des Functions-
wechsels. Leipzig.

Harcxet, E. 1866.—Generelle Morphologie der Organismen. (The essential points of the

“ Biogenetic Law ” appear in all Haeckel’s later books, e.g. ‘The Evolution of
Man,’ Watts & Co., 1906-~a translation of his ‘ Anthropogenie,’ 1891.)

Hertwia, Oscar. 1906.—Handbuch der... . Entwickelungslehre der Wirbelthiere, i.

Hurst, C. H. 1893.—- The Recapitulation Theory. Natural Science, ii. pp. 195-200 &
364-369.

Hyarr, A. 1897.—Cycle in the Life of the Individual (Ontogeny) and in the Evolution of
its own Group (Phylogeny). Proc. Amer. Acad. Arts & Sciences, xxxii.
pp: 209-224, (hicludes sketch of the history of the doctrines of Recapitu-
lation and “ Palingenesis,” with quotations from Meckel, Vogt, L. Agassiz,
&c.)

KLEINENBERG, N. 1886.—Ueber die Entwicklung durch Substitution yon Organen. (Die
Entstehung des Annelids, &c., Kap. vi.) Zeit. f. wiss. Zool. xliv.

Kroun. 1853.—Ichinospira. Arch, f. Naturgeschichte, xix. pp. 223-285, & 1855, xxi.

A CRITICAL RE-STATEMENT OF THE BIOGENETIC LAW. 101

Kroun. 1857.~—Loe. cit, xxiil.

MacBripr, E. W. 1911.—Two abnormal Plutei of Echinus &e. Quart. Journ. Mier.
Sci. Lvii.

MacBripk, f. W. 1914.—Heape’s Text-Book of Embryology, i. Invertebrata.

MacBripn, Hh. W. 1917.—Recapitulation as a Proof of the Inheritance of Acquired
Characters. Scientia, xxii. pp. 425-484,

Marswaty, A. Minnes. 1890.—Presidential Address to the Biology Section, British
Association, Leeds Meeting. (The most complete of all summaries of evidence
bearing on the Recapitulation Theory.)

Meyrick, Ef. 1895.—Handbook of British Lepidoptera. Macmillan.

Morean, T. H. 1908.—EKyolution and Adaptation. Macmillan.

Morean, T. H. 1919.—A Critique of the Theory of Evolution.

Sppewick, ApAM. 1894.—On the Law of Development commonly known as Von Baer’s
Law &c. Quart. Journ. Mier. Sci. xxxvi.

Sepewick, Adam. 1909 (a).—The Influence of Darwin on the Study of Animal Embry-
ology. Darwin and Modern Science.

Smpewrex, Avam. 1909 (6).—Studeut’s Text-Book of Zoology, iii.

SmrotH, H. 1895.—Die Gastropoden der Plankton-Expedition. Ergebnisse der Expedi-
tion I]. Ff. (Mechinospira, pp. 29-52, Taf. xvi., xvii., xviii., text-fig. 5.)

Situ, Grorrrey. 1911.—Primitive Animals. Cambridge University Press.

Souru, R. 1908.--The Moths of the British Isles, ii. Warne & Co.

Weismann, A. 1904.-—The Evolution Theory, ii, pp. 159-191. - Edward Arnold.

Rie

So An

THE TERRESTRIAL ISOPOD ELUMA CALATUM. 103

On the Terrestrial Isopod Hluma celatum (Miers)=purpurascens, Budde-
Lund. By Watrer H. Continan, D.Sc.,: F.L.8., Keaper of the
Yorkshire Museum, York.

(PLATE 8.)

[Read 15th June, 1922. }

THe genus Elwma was described by Budde-Lund* in 1885 for a species of
Terrestrial Isopod which he named LH. purpurascens. The description of
both the genus and species is imperfect, and no figures accompany it.

As a doubtful synonym Budde-Lund gives the Armadillidium cvlatum of
Miers, which was described and figured by Miers in 1877 from specimens
obtained at Cayenne, French Guiana, South America f.

A careful comparison of the description and figures, both imperfect,
with specimens received from Mr. D. R. Pack-Beresford, from the Hill of
Howth, Co. Dublin, Ireland, leaves no doubt, in my mind, as to the identity
of Miers’s species with these. His specific name celatum therefore has
priority, and must replace that of purpurascens.

Neither Miers or Budde-Lund gave any figures of the structure, and their
references are very brief and in some points scarcely accurate ; I am there-
fore venturing to redescribe and figure the genus and species.

Mr. Pack-Beresford } figures the external appearance of this species, but
his illustrations are not correct in all details. Verhoeff$ has given two
figures of parts of the exoskeleton.

The Rhacodes inseriptus of Koch ||, regarded by Dollfus and Budde-Lund
as a synonym of Zylos latreillei, Aud. & Say., was thought by Eaton | to be
referable to this species.

Euuma, Budde-Lund.

Body oblong-ovate, strongly convex, setose, and closely and minutely
punctured. Cephalon strongly marginate, with median aud lateral lobes ;
epistome with sloping dorsal portion and keeled. Eyes simple, very small.
Antennule small, 3-jointed, terminal joint conical. Antenne somewhat
short, flagellum bi-articulate. Pleural plates of mesosomatic segments 2-7
slightly excavate anteriorly, ventral margin indentate on segments 2-4, trun-
cate on 6-7. Coxopodite of first segment separated from the pleuron and

* Crust. Isop. Terr. 1885, p. 48.

+ Proc. Zool. Soc. Lond. 1877, p. 665, pl. 67. figs. 3-3 6.
{ Irish. Nat. 1908, p. 255, pl. 10. i

§ Arch. f. Biontologie, 1908, Bd. ii. p. 371.

|| In Rosenhauer’s ‘ Die Thiere Andalusiens,’ 1856, p. 422.
4] Ann. & Mag. Nat. Ilist. 1882 (s, 5), vol. x. p. 360.

104 — DR. W. B. COLLINGE ON THE
forming a notch on the posterior margin. ‘Telson triangular, width greater
than the length, not extending beyond the uropoda. Uropoda short, extend-
ing slightly beyond the telson ; basipodite robust, thickened, antero-dorsal
surface expanded, articulating ventro-anteriorly ; exopodite flattened, ex-
panded, laminate ; endopodite styliform, elongated.

This genus is undoubtedly most nearly related to the genus Armadillidium,
Brandt, as is readily seen in the form of the body generally, in the structure
of the cephalon and its appendages, and in the form of the uropoda and
telson. It differs, however, from Armadillidium in the following characters :—
The cephalic lobes are more feebly developed and the epistome is keeled in
the middle line, and has a sloping dorsal portion, and auricula-shaped pro-
minences above and lateral to the antennal sockets; the eyes are small and
simple ; the pleural plate of the first mesomatic segment exhibits a notch on
the posterior angle formed by the protrusion of the coxopodite ; finally the
uropoda extend beyond the telson.

In the sloping dorsal portion of the epistome and in the form of the first
mesosomatic seoment, there is a resemblance to the condition found in certain
South African Cubaridee.

HnumMa caLatum (Miers). (PI. 8. figs. 1-12.) :
Armadillidium celatum, Miers, Proc. Zool. Soc. Lond. 1877, p. 665, pl. 67. figs. 3-36.
Eluma purpurascens, Budde-Lund, Prosp. gen. spec. Crust. Isop. Terr. 1879, p. 6;

Crust. Isop. Terr. 1885, p. 48.

> 5 Dollfus, Ann. Soc. Espan. Hist. Nat. 1892, xxi. p. 164; 3 Con-
eres Inter. Zool., Leyden, 1895, p. 857; Mérm. Soc. Zool.
France, 1906, ix. p. 528.

% Ms Norman, Ann. & Mag. Nat. Hist. 1899 (s. 7), iii. p. 69.

s Verhoeff, Arch. f. Biontologie, 1908, ii. p. 371, t. 31.
figs. 68, 69.

0 5 Pack-Beresford, Irish Nat. 1908, p. 255, pl. 10.

0 a1) Pack-Beresford & Foster, Proc. Roy. Inish Acad. 1911, xxix.

(s. B), p. 185, pl. 8. figs. 4 a-0.
Lluma celatum, Collinge, Check-List Brit. Terr. Isop. 1917, p. 115.

Body oblong-oval, strongly convex, smooth and somewhat shiny, minutely
punctured, clothed with short, thick sete. Cephalon (figs. 1 & 2) strongly
marginate anteriorly, median lobe extending beyond the lateral lobes, which
are small; epistome with sloping dorsal portion and keeled in the median
line, with auricula-shaped prominences above and lateral to the antennal
sockets. yes simple, very small. Antennul (fig.3) small, 3-jointed, with
three or four short sete on the sid& of the third joint. Antenne (fig. 4)
somewhat short, 1st joint small, 2-4 almost subequal, 5th elongated ; flagellum
biarticulate, proximal joint less than half the length of the distal one, with
fine terminal style. Wirst maxillee (fig. 5) with the outer lobe terminating in
ten spines, the first three of which are strong and curved, the 4th, 5th, and

TERRESTRIAL ISOPOD RLUMA CASLATUM. 105

6th smaller, the 7th a long fine spine, 8th and 9th very small, 9th and 10th
faintly denticulate, setose on the outer border of the lobe; inner lobe pro-
longed on the outer side terminally, with two short setose spines. Maxilli-
pedes (fig. 6) with comparatively small lobes, outer lobe 3-jointed, 1st joint
short, with two large spines, 2nd joint also with two large spines, 3rd joint
terminating in a number of spinous processes ; inner lobe with a single long
pointed spine, and two tooth-like spines towards the inner border. ‘The seg-
ments of the mesosome (figs. 7-9) strongly convex, subequal, pleural plate
of the Ist segment flanking the cephalon anteriorly, strongly curved, lateral
margin thickened, posterior angle notched. Pleural plates of 2nd to 7th
segments excavate anteriorly, ventral margin of 2nd to 4th indentate, 6th to
7th truncate. Uropoda (figs. 10 & 11) short, but longer than wide, extend-
ing beyond the telson ; basipodite robust and thickened, antero-dorsal surface
expanded, articulating ventro-anteriorly ; exopodite flattened, expanded, and

“laminate, protruding on the inner side ; endopodite styliform, ionger than
the exopodite, widest just above the middle, with terminal setaceous pad and
spinous setee. Telson (fig. 12) roughly triangular, with rounded apex, width
greater than the leneth, not extending beyond the pleural plates of the last
segment of the metasome. ;

Length 11°5 mm.

Colour (in alcohol) a light coffee-brown.

Hab. Hill of Howth, Co. Dublin, Ireland (). R. Pack-Beresford).

In the figure given by Mr. Pack-Beresford (op. eit. pl. 10) the distal
joint of the flagellum of the antenn is too short and no style is shown,
whilst the pleural plates of the metasome are scarcely long enough ; further,
the last segment extends more posteriorly than is shown in his figure (c/.
Pl. 8: fig. 12).

The distribution of this species is as yet only very imperfectly known. It
was originally described from Cayenne, French Guiana, South America.
Budde-Lund* says it is “commonest in the Island of Madeira.” ... “I
have seen some from Cayenne taken by Don Gelski.” . . . “Two specimens
taken by Professor Reinhardt in the Isiand of Nicobar, and determined by t
Cl. Kroyer under the name of Armadillidium purpurascens, are preserved in
the Museum at Copenhagen.” f.

Dollfus § records it from many localities. He states :—“The genus Hluma
is entirely Western, and contains only one species, /. purpurascens, B.-L., a
woodlouse of a purple-red colour, remarkable for its simple eyes. Very
abundant in the Atlantic archipelagos, in the Canaries, the Azores, and
Madeira ; it oceurs again at Cintra (Portugal), and in Western Algeria, but

* Crust. Isop. Terr. 1885, p. 48.

+ Mr. Pack-Beresford (op. cit. p. 257) translates this described by.

{ Dr. kX. Stephenson has very kindly referred to these specimens and sent me notes thereon.
§ 3 Congrés Intern. Zool., Leyden, 1895, p. 357.

106 THE TERRESTRIAL ISOPOD ELUMA CHLATUM.

the most curious fact about it is its range northwards to the French Depart-
ment of Les Charentes, where it is acclimatised from Angouléme to Royan
(very common twenty years ago in this locality, it has become much rarer
lately), and southwards to Cayenne!” (The Cayenne here quoted would
appear to be in France, and must not be confused with Miers’s original
locality.) The same author records it from Constantina in the Sierra Morena
(Spain), and states, with reference to the record for the French Department
of Les Charentes *, “oti il a été certainement introduit,’ but gives no reason
for this supposition. g

Norman f records it from Madeira, where he found it “‘up to heights of
between 2000 and 3000 feet.”

Finally, Mr. Pack-Beresford (op. cit.) has recorded it from the Hill of
‘Howth and Portmarnock, Co. Dublin, Ireland.

The allied species L. helleri, Verhoett, is from the Island of Teneriffe.

EXPLANATION OF PLATE 8.

Illustrating Dr. Walter E. Collinge’s paper “ On the Terrestrial Isopod Elwma
celatum (Miers).”

Eluma celatum (Miers).

Fig. 1. Dorsal view of the cephalon. x10.
2. Anterior view of the cephalon. X7‘5.
3. Antennule. X110. -
4, Right antenna.
5. Terminal portions of the inner and outer lobes of the right first maxilla. x56.
6. ” portion of the left maxillipede. x56.
7. External view of the pleural plate of the first mesosomatic segment.
8. Internal view of the pleural plate of the first mesosomatic segment.
9: ; 2 lateral portion of the second mesosomatic segment.
10, Right uropod, dorsal view. :
1 " » wentral view.

SS)

. Last metasomatic segment and telson. x 8.

The Author desires to thank the Carnegie Trust for the Universities of
Scotland for a grant to defray the artist’s charges.

* Ann, Soc. Espan. dist. Nat. 1892, xxi. p. 164.
+ Ann. & Mag. Nat. Hist. 1899 (s. 7), iii. p. 69.

Journ. Linn. Soc. Zoot, Vol XXXV P18

Aa, J.J, Del. adnat. JT. Rennie Reid Lith Fdint

ELUMA CAELATUM. (MIERS)

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TWO NEW TERRESTRIAL ISOPODS FROM MADAGASCAR. 107

On two new Terrestrial Isopods from Madagascar. By Waurer H. CoLiincE,
D.Se., F.L.S., Keeper of the Yorkshire Museum, York.

(PLATE 9.)

| Read 15th June, 1922. |

CONTENTS,
Page
Nigel ntroduchronppraawrier teary cect rare aiaie perce Percent 107
If. The Isopodean Fauna of Madagascar...................+..00-- 107
III. Description of .Al/oniscws macrews, sp. NOV. .. 1... eee eee eee 108
IV. Description of the Genus Culmanesia, gen. nov. ..........-+00-- 109
Calmenescaymethuents spel OVep ci tye liersi= i ielelelrfelche el eteysiisiels celts 109
Gs CEMA INT oogocansdoneceseh Goseobudovcnaonoo8 110
Os, Mia. Coane. “sercos oo poncGbaceopap oa cuaaeuboUmoe on 111
1. yes. 2. Antennule. 5. Antenne. 4. Oral Appen-
dages.
G@, MUNG WIOROMN® S506 nondboceopE Loud doonoosedanongoD 111
1. Pleural Plates. 2. Appendages.
dewbhepMetasoniewmerry etic erie reaicteirrrisier triers 112
1. Uropoda. 2. Telson.
Wool ZNO Wo sig olg oo cope enoNen Doom SUCH Hor xododoeoHbonuiads 112
Bibliograp tive mua weiiee te mer iyi eat ieee tversee ra ele te tomscel steneeeeel ck vero vaoreee ere er toe 112
Tq OF WHO IRM Gen aovcogdvoevausoaeodangodbooeNoovuEOE 113

I. InrRoDuctrION.
For the privilege of examining the very interesting specimens of Terrestrial
Isopods here described, I am indebted to the kindness of my friend Dr. W-
T. Calman, of the British Museum (Natural History). The material was
collected in Madagascar in 1911 by the Hon. Paul A. Methuen.

There is a single specimen referable to the genus Adloniscus, Dana, not
hitherto described, remarkable for its great size; and a further very interest-
ing new species, for the reception of which I have constituted a new genus,
to which I have given the name of Calmanesia in honour of Dr. W. T. Calman.

II. Tor Isopopran Fauna or Mapacascar.

Very little attention has been paid to the Terrestrial Isopods of Madagascar.
Budde-Lund (3) has described certain species in the collection of the Berlin
Museum. Dollfus (8) in 1889 described the Porcellio cristatus, placed
by Budde-Lund in the genus Lyprobius and later in the genus Nagara. The
same author in 1895 (9) described a further series of species, amongst which

108 DR. W. E. COLLINGE ON TWO

there were four new species of Alloniscus ; these Budde-Lund placed in a
new genus, Miacara, for reasons which are not altogether clear ; there are
certainly slight differences in the form of the uropods, but these are scarcely
sufficient to warrant generic distinction. In 1908 (4) Budde-Lund published
his memoir on the Isopoda of Madagascar and Hast Africa, in which he
described and partly figured nine further new species, re-describing and
partly figuring other known forms. No figures, however, are given of
Periscyphops preconius and Armadillo horridus. According to this author,
there are known from Madagascar thirty-two species referable to twenty
genera,

If, as is generally supposed, Madagascar represents a part of a great sub-
merged southern continent, then we should expect to find representatives of
South-Hastern and South-Western genera, but whether owing to long isolation
these forms have changed or died out, it is remarkable that the bulk of the
known forms show affinities rather with Northern forms than with Southern
ones, and they further seem to have little relationship with the South African
genera. Hithertono genus has been found peculiar to Madagascar ; the new
genus Calmanesia, here described, is therefore of more than usual interest.
Until, however, we know much more of the fauna of this island, it is futile
to speculate.

III. Drscriprion oF Axzonrscus vACREUS, Sp. NOY.

Aoniscus, Dana.

This genus as yet is only imperfectly understood, and the members are
subject to a wide range of variation. It occurs in North and South America,
India, Siam, the Malay Peninsula, South Africa, Sumatra, Java, and the
Maldive, Hawaiian, Nicobar, and Celebes Islands. Arhina, Collge. (6) is
a closely-allied genus from India.

Of the twenty-one described species of Alloniscus, seven are known to
occur in Madagascar, viz.: A. alluaudi, Dollf., A. elegans, Dollf., A. guttatus,
Dollf., A. tigris, Dollf., A. brevis, Budde-Lund, A. pallidulus, Budde-L.,
and A. pigmentatus, Budde-L.

ALLONISCUS NACREUS, sp. nov. (PI. 9. figs. 1-12.)

Body broadly oval, strongly convex, irregularly pitted, finely sculptured
on each side of the mesosome. Cephalon (figs. 2 & 3) medium size, convex
above, frontal margin distinct, lateral lobes small; epistome slightly de-
pressed, with prominence between the antennze. Eyes oval, dorso-lateral.
Antennule (fig. 4) small, curved, and 3-jointed, the terminal joint with
a number of bristle-like setee on the inner side. Antenne (fig. 5) short,
joints gradually enlarging from the Ist to 4th, 5th joint elongated and more

NEW TERRESTRIAL ISOPODS FROM MADAGASCAR. 109

slender ; flagellum 3-jointed, with short terminal style. First mavxillee
(fig. 6): outer lobe terminating in four stout spines and six smaller ones, with
stout setee on the outer margin of the appendage. Second maxillee (fig. 7)
thin and plate-like, with terminal bifurcation, densely setose. The segments
of the mesosome are strongly convex and finely pitted, with lateral sculptur-
ing; pleural plates with terminal margin more or less truncate, posterior
margin of the 5th, 6th, and 7th slightly produced backward. Maxillipedes
(fig. 8) rather narrow ; outer lobe 3-jointed, with two spines on the first joint,
two tufts of spines on the second, and asingle terminal tuft on the third; the
inner lobe is somewhat conical and surmounted with numerous small setze.
Appendages of mesosome (fig. 9) stout and strongly spinous. Metasome
comparatively small, segments 3-5 with pleural plates directed backward and
inwards. Uropoda (figs. 10 & 11) short, extending beyond the telson, basal
plate sparsely covered with sete; exopodite sickle-shaped; endopodite
slender, terminating in three long sete. Telson (fig. 12) triangular, lateral
margins slightly rounded, apex subacute, with slight depression above.

Length 19 mm.

Colour (in alcohol) creamy white.

Hab. Tamatave, East coast of Madagascar (Herschell §- Chauvin).

This fine species is the largest member of the genus yet described. Apart
from this feature, it conforms with the essential characters of the genus.

IV. Discription oF THE GENUS Cazes.

CALMANESIA, gen. nov.

Body oblong-oval, capable of partly rolling into a ball. Segments of
mesosome and metasome with a series of long-jointed spines. Cephalon
with lateral and median lobes. Antennulse very small. Antenne elongated ;
flagellum 2-jointed. Pleural plates, excepting those of the first segment,
drawn out into a long spinous process. Uropoda small and without endo-
podites. Telson short and obtuse, not extending beyond the uropoda.

CALMANESIA METHUENI, sp. nov.. (Pl. 9. figs. 13-25.)

Body oblong-oval, covered with elongated jointed spines. Cephalon (figs.
14 & 15) short, slightly convex dorsally, frontal margin distinct, with lateral
and median lobes; seven spines on the dorsal surface ; epistome slightly
convex. Hyes situated dorso-laterally, facets few and large. Antennulee
(fig. 16) small, 3-jointed, middle joint very small. Antenne (fig. 17) elon-
gated, 1st joint small, 2nd and 38rd larger, 4th and 5th greatly elongated ;
flagellum 2-jointed, distal joint rather longer than the proximal one, with
terminal bunch of sete. First maxillee (fig. 18): outer lobe with six short

110 DR. W. E. COLLINGE ON TWO

spines, of which the outermost is the largest and has a short, stunted tooth-
like body at its base; on the inner side is a single needle-like spine ; densely
setose on the outer margin; inner lobe small, with two elongated setose
spines terminally. Second maxille thin and plate-like, setose terminally.
Segments of the mesosome strongly arched, the Ist with eleven jointed
spines and the remainder with seven ; pleura] plates, excepting those of the
Ist segment, produced into long spinous processes. Maxillipedes (fig. 19)
somewhat robust ; outer lobe 3-jointed, with two straight spines on the Ist
joint, four on the 2nd, and one small one on the outer border of the 3rd joint,
which latter terminates as a blunt process surmounted by a number of sete ;
inner lobe plate-like, with two small tooth-like spines and a single straight
spine on the ventral face. Appendages of the mesosome (fig. 20) compara-
tively short. Segments of the metasome crowded together, 3rd, 4th, and 5th
each with two jointed spines in the mid-dorsal line ; pleural plates greatly
elongated, directed backward and terminating in a finely drawn out spine.
Uropoda (fig. 24) extending beyond the telson, basal plate oval-shaped, with
a single appendage (exopodite) which terminates in a strong curved spine.
Telson (fig. 25) somewhat triangular in shape, terminally obtusely pointed,
with two 3-jointed spines and a small median papilla in front of these.

Length 10 mm.

Colour (in alcohol) creamy white with dark chocolate-coloured markings.
Younger forms creamy white.

Hab. Forest of Folohy, Hast Madagascar, 1911 (Herschell § Chauvin) ;
Analamazétra, Eastern Forest, June 1911.. Under rotten logs (P. A.
Methuen). The specimens from the latter locality are evidently immature.

a. General Form.—This is undoubtedly one of the most handsome and
peculiar Terrestrial Isopods known, rivalling in both form and colour any
species hitherto described. The long-jointed spines covering the mesosome
and metasome and the drawn-out spinous pleural plates give the animal the
appearance of some Coleopterous larva rather than a Crustacean.

There is, I think, little doubt but that the members of this genus are
capable of rolling themselves up into a ball like the Armadillidiidee. Apart
from the faet that one specimen was found (in alcohol) so rolled up, the
extreme mobility of the segments support such a view.

The remarkable jointed spines on the body would seem to be a hitherto
undescribed character in the Isopoda. I have been unable to find any
reference to such in the literature on these Crustaceans. So far as I can
make out from the material available, the spines first appear as small excres-
cences on the surface of the segments (fig. 23, a); these increase in size and
later appear as ordinary spinous processes (fig. 23, >); a little later they con-
tinue to grow, a distinct joint forming at the base of the first period of
growth (fig. 23, ¢c), and this continues until, with the exception of those

NEW TERRESTRIAL ISOPODS FROM MADAGASCAR. 1G

on the cephalon and telson, where the spines are 3-jointed, they become
_ 4-jointed, the longest measuring 6°5 mm. (fig. 21). All the spines are
covered with minute scales like the rest of the body (fig. 22). There are
seven spines on the cephalon, eleven on the first mesosomatic segment, seven
on the second to seventh segments, and two on-each of the last three meta-
somatic segments and the telson.

b. The Cephalon.—This is extremely narrow from before backwards and
slightly convex. Anteriorly there is a well-marked marginal ridge which
laterally develops into the two lateral lobes, and in the middle into the
median lobe. Posteriorly the margin is deeply excavate, and from the middle
line a strong spine arises ; lateral to this there are a pair of spines on each
side, and a single one on each side, slightly nearer the centre, in front of the
former. All of these spines are 3-jointed. The epistome is slightly convex
dorsally and distinctly so ventrally, whilst laterally the sockets for the
articulation of the antenne stand out as conspicuous bodies.

1. The Eyes are placed dorso-laterally ; they are of medium size and with
few but large facets.

2. The Antennule (fig. 16) are small, and consist of an elongated basal
joint, a tiny ring-like middle joint, and a gradually tapering terminal one.
At the distal end of the third joint and on the inner side are two blunt
papillee.

3. The Antenne (fig. 17) are greatly elongated. The first joint is small,
and the second about two and a half times as long, the third is still longer
and narrower, whilst the fourth and fifth are almost coequal and nearly as
long as the second and third together. All the segments are covered with
minute setee. The flagellum is 2-jointed, the distal joint being rather longer
than the proximal one, and terminates in a bunch of elongate sete.

4. Oral Appendages (figs. 18 & 19).—These afford very little assistance in
placing this genus and species. The 2nd maxillse and the maxillipedes are
of the usual type. The Ist maxillee in the form of the first tooth of the outer
lobe differs somewhat from any other genus. —-

c. The Mesosome.—-The segments of the mesosome are strongly arched.
There are eleven jointed spines on- the first segment and seven on the
remainder.

1. The Pleural Plates of the first segment are broad plate-like bodies with
rounded anterior and posterior angles; anteriorly they bound the postero-
lateral half of the cephalon. Those of segments 2-4 are somewhat triangular
in shape, the apex of the triangle being drawn out in a fine curved spine ;
the whole of the plate is curved, the free spinous end bending upward. On
segments 5-7 the plates terminate more abruptly before the commencement
of the spine, and they are rather flatter. -

112 DR. W. E. COLLINGE ON TWO

2. Appendages.—The walking limbs are comparatively short, and of a less
robust type than in most genera of Oniscidee.

d. The Metasome.—The segments of the metasome are crowded together,
the Ist and 2nd being scarcely visible in a dorsal view and without pleural
plates. Of the remaining three the last is the smallest. The pleural plates
are of a similar type to those of the last three metasomatic segments. There
are a pair of jointed spines on each of the last three segments.

1. Uropoda.—The uropoda extend beyond the telson. The basal plate is
somewhat oval in shape and has a fold on its posterior side. Only a single
appendage appears to be present, which is slightly longer than the width of
the basal plate and terminates in a strong curved spine. I take this appendage
to represent the exopodite.

2. Telson—The telson is comparatively small, somewhat triangular in
shape, and terminating in an obtuse point. It bears a pair of 3-jointed
spines and a median papilla anteriorly and between the bases of the spines.

V. AFFINITIES.
The genus Calmanesia appears to be widely separated from any known
genus, and at present we must leave it without attempting to relegate it to
its proper position in the classification.

BIBLIOGRAPHY.

1. Branpt, J. F.—Conspectus monographize Crustaceorum Oniscodorum
Latreilli. Bull. Soc. Imp. Nat. Moscou, 1833, vol. vi. pp. 71-193.

2. BuppE-Lunp, G@.—Crustacea Isopoda Terrestria. Hauniz, 1885.

3. —— A Revision of “Crustacea Isopoda Terrestria.” 1899-1904,
pts. i.iii., pp. 1-144, t. 1-10.

4, Isopoda von Madagascar und Ostafrika. Voeltzkow, Reise in
den Jahren 1903-1905, Bd. ii. pp. 265-308, t. 12-18. Stuttgart,
1908.

5. Terrestrial Isopoda, particularly considered in relation to the

Distribution of the Southern Indo-Pacific Species. Trans. Linn. Soc.
Lond., Zool. ser. 2, vol. xv. 1912, pp. 367-394, pls. 20-22.

6. Contincr, WALTER E.—Contributions to a Knowledge of the Terrestrial
Isopoda of India. Part I.—Rec. Indian Mus. 1915, vol. xi. pp. 143-
151, pls. 4-12.

Contributions to a Knowledge of the Terrestrial Isopoda of
Natal.—Part I. Ann. Natal Mus. vol. iii. 1917, pp. 567-585,
pls. 40-42.

8. Dotirus, A.—Sur quelques Isopodes du Musée de Leyde. Notes fr.
Leyden Mus. vol. xi. 1889, pp. 91-94, t. 5.

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AGASCAR.

Journ. Linn. Soc. Zoo. Vel XXXV P]. 9.

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NEW TERRESTRIAL ISOPODS FROM MADAGASCAR. 113

9. Douirus, A.—Isopodes terrestres recueillies 4 Diégo-Suarez, 4 Tamatave
et ala Réunion. Mém. Soe. Zool. France, 1895, T. viii. pp. 180-188,
fies. 1-12.

a

10.

Voyage de M. H. Simon dans l’Afrique australe. bid. pp. 345-

352, figs. 1-9.

EXPLANATION OF PLATE 9.

Illustrating Dr. Walter E. Collinge’s paper “On Two new Terrestrial Isopods

Fig. 1.

SOM ND HP to

22,
23,
24,
25.

from Madagascar.” :

Dorsal view of Adloniscus nacreus, sp.nov. X 3.
5 0 the cephalon. x 12.
Anterior view of the cephalon. X 12.

. Left antennule. x 50.

Right antenna. x 25.

. Terminal portion of the inner lobe of the left 1st maxilla, ventral view. x 80.

left 2nd maxilla, ventral view. x 80.
right maxillipede, ventral view. x 80.

” ”

” ”

. Second thoracic appendage, dorsal view. x 25.
. Ventral surface of the terminal portion of the 2nd thoracic appendage, showing

spinous area. X 25.

. Dorsal view of the right uropod. x 25.

D », the telson and last metastomatic seement. x 10,

. Dorsal view of Calmanesia methueni, gen. et sp.nov. xX 6.

5 », thecephalon. x 20.

. Anterior view of the cephalon. x 20.
. Left antennule. x 80.

Right antenna. xX 20.

. Terminal portion of the inner and outer lobes of the right 1st maxilla, ventral

view. x 80.

. Terminal portion of the right maxillipede, ventral view. x 75.
. Second thoracic appendage, dorsal view. x 20.
. Jointed spine from off the mesosome. X 20,

Portion of same, showing the scales. x 80.

a-d. Stages in the development of one of the jointed spines.

Dorsal view of the right uropod. x 50,

Terminal segment of the metasome and telson, seen from the anterior. x 20.

The Author desires to thank the Carnegie Trustees for the Universities of
Scotland for a grant to defray the artist’s charges.

LINN. JOURN.— ZOOLOGY, VOL. XXXYV.

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THE MOUTH-PARTS OF THE SHORE CRAB. 115

~ On the Mouth-parts of the Shore Crab. By L. A. Borrapatnr, Se.D.
Fellow and Tutor of Selwyn College, Cambridge, and Lecturer in
Zoology in the University. (Communicated by Professor H. S.
Goopricu, M.A., F.R.S.)

(PratEs 10, 11.)
Read 6th April, 1922.
I

I,

THE organs which stand about the mouth of a Decapod Crustacean make up
a complex as intricate as any that is to be found in the Animal kingdom.
They are also extremely important to their possessor, for without them the
animal can neither feed nor breathe, and to one of them falls, at least in
many cases, the duty of keeping clean the indispensable organs of special
sense. Yet they are at present but little understood.

In a study upon the Common Prawn, published in 1917 (5), I endeavoured
to solve the problem which the morphology of these organs presents, and
made a beginning with the investigation of their working. This paper
contains an account of some observations upon Carcinus menas, a species at
the other end of the decapod series.

The term “ mouth-parts ” denotes, in the Crab, a number of organs which
stand upon the under side of the body, in the region which is bounded behind
by the anterior edge of the mass of fused postoral sterna, at the sides by the
edge of the inturned carapace where this encloses the exhalent passage of
the gill-chamber, and in front by the fused antennal and mandibular sterna
(epistome). The sterna of the maxillulary to second maxillipedal segments
inclusive are more intimately united with one another than those of the legs;
and they form,a mass, roughly triangular with the apex forwards, that
stands out steeply from an area, in front of and beside it, which is covered
by a thin cuticle supported upon the pieces of the endophragmal skeleton.
It is upon this area that the mouth-parts are inserted. They are: the six
pairs of limbs from the mandibles to the third maxillipeds inclusive, the
upper lip or labrum, the lower lip or metastoma, and the fleshy opening of
the mouth itself. I propose to describe in succession each of these parts and
its movements, and then to discuss the functions of the complex as a whole.

Il.

1. The Third Maxilliped is built wpon the plan of the legs. This statement,
by a phenomenon often seen in serially homologous structures, is true even of
features—such as the fusion of the basis and ischium and the nature of the
articulation of the joints—which cannot be supposed to have existed in the
schizopod ancestor whose thoracic limbs were not differentiated from one

&

116 DR. L. A. BORRADAILE ON THE

another. ‘he limb is a broad structure, flattened in that direction which is
morphologically antero-posterior, and widened from side to side. In what
may be called the normal position it is turned forwards till its wide plane
passes the horizontal and slopes a little upward in front. In this position it
meets its fellow on the middle line, and the two form an operculum which
almost wholly encloses the mouth-field, abutting behind on the sterna, at the
sides upon the edge of the carapace, and in front upon the epistome save in
the middle, where a gap is left through which the gill-stream can flow even
when the operculum is most tightly closed. Hach third maxilliped articu-
lates ventrally with the hinder angle of its sternum, and dorsally (anteriorly)
with the epimerite of its segment.

The cova is of oval transverse section. Its proximal rim bears on the
under side the knob which makes articulation with the sternum, where it is
received by a saddle-shaped hollow between two processes. Adjoining this
is a facet on the median surface of the joint, which works against the
external (antero-lateral) face of the sternum. On the dorsal side the rim
bears a socket which articulates with a correspondingly shaped process of the
epimerite. On the outer side the coxa has a backward-curving flange, to
which, by a flexible suture, is hinged the epipodite. This organ has a stout
base, which bears on the dorsal side the small podobranch, and a long, blade-
like process which, diving under the edge of the branchiostegite, enters the
gill-chamber and passes between the posterior arthrobranch of its own limb
and the anterior arthrobranch of the cheliped, to lie between the gills and
the side of the body. The whole organ, and the flange upon which it is
borne, is spirally twisted, so that, starting in the horizontal plane in which
the main part of the limb lies below the mouth, it ends in a vertical plane
against the flank. The flange and the base of the epipodite stand in that
gap, between the anterior face of the coxa of the cheliped and the branchio-
stegite, which is the anterior inhalent opening of the gill-chamber; and their
twisted shape bears such a relation to the opening that when the maxillipeds
are in the normal position they lie across it and almost but not quite close it,
but when the maxillipeds are divaricated, the epipodites lie in the midst of
the opening, with their flat sides parallel fo the stream, to which they offer
little opposition. The part of the opening which is covered when the
maxillipeds-are approximated is the anterior. The extent to which the
hinder part remains open varies with the position of the cheliped.

The basis and ischium are fused, though traces of their junction usually
remain in the form of a groove. By this it is shown that the basis is a small,
region which in width makes the transition from the stout coxa-to the flat
ischium, and in position fills a triangular gap between them, due to the fact
that the ischium is displaced to the median side of the coxa. The free (median)
edge of the basis abuts upon the edge of the sternum, and continues the
contact made by the facet upon the coxa. On the outer side the basis bears

MOUTH-PARTS OF THE SHORE CRAB. 117

the exopodite. The stem of this is along, narrow, flattened structure, directed
forwards between the endopodite and the branchiostegite, and bearing at its
end a flagellum which is directed inwards above the merus towards the middle
line. The flagellum consists of a basal joint, and a many-jointed lash,
curved with the concave side forwards. The exopodite stands at the junction
of basi-ischium and coxa, and besides being articulated to the former is
attached by membrane to the latter.

The ischium is an oblong, roughly rectangular, and very flat joint, which
makes nearly half the total surface of the limb. It is succeeded by the merws,
a flat, subquadrate structure, stouter than the ischium and tilted a little
outwards upon the latter. Owing to its stouter form it has a median face in
place of an edge, and upon this face, which is hollowed, is set the carpus.
That joint, with the two which succeed it, forms a subcylindrical, tapering
palp, which can be stretched out so as to be roughly in line with the axis of
the basi-ischium and merus, but in the normal position is folded back against
the hollow face of the merus. In this position the palps fill the gap which
would otherwise be formed by the outward tilting of the merus, and thus
complete the operculum.

As has been mentioned, the mode of articulation of the joints is that which
is found in the legs. At the proximal end of each, the rim of the hard
cuticle is raised, at two points on opposite sides, into articular processes
which, bridging over the arthrodial membrane, play against corresponding
surfaces on the distal rim of the preceding joint; and these articular
surfaces are so shaped as to limit the movement of the distal of the two joints
toa particular direction. Since, however, this plane is a different one in each
joint, the limb as a whole has a good deal of mobility, though owing to its
shape and position the third maxilliped is less freely movable than the legs.
I have not thought it necessary to give details of the articulations of the:
several joints, but particulars of the movements they permit will be found
below. They are modifications of those of the segments of the legs.

A remarkable feature of the third maxilliped is the way in which it is
completely outlined with hairs, set along the edges of its flat surfaces. Most
of these hairs are feathered in one way or another. They are short where
the edge is, in the normal position, apposed to some other structure, but long
on that part of the anterior (morphologically external) edge of the merus and
carpus over which the gill-stream flows. Along the median edge of the
ischium there run on the lower (ventrally-facing) side two narrow parallel
bands of hairs with a naked, convex strip between them. These hairs are
stout, with a close, stiff feathering on all sides. The actual edge is finely but
bluntly toothed. On the inner or dorsal (morphologically anterior) side of
this joint there is a row of hair-tufts near the median edge, and a rather
sparse band of hairs near the outer edge. The dorsal side of each of the last
three joints is covered with long, and stiff, serrated bristles. The epipodite

118 DR. L. A. BORRADAILE ON THE

is fringed on both edges with long hairs, and the ventral (hinder) surface of
the flange of the coxa upon which it stands is very hairy. The flagellum of
the exopodite bears a fringe of long, feathered hairs, many of them jointed.
A more detailed description of the hair-system of this and other limbs of the
Shore Crab is given hy McIntosh (11). With its functions I will deal later.

The cuticle of the third maxilliped, except the flagellum and the blade of
the epipodite, is hard and pigmented, like that of the rest of the exposed
surfaces of the body. In this respect it contrasts strongly with that of the
other mouth-parts.

The muscular system of the third maxilliped closely resembles that
described by Pearson (18) for the same limb of the Edible Crab, and I have
not examined it further than was necessary tq assure me of this fact. Hach
of the joints of the limb has a flexor and an extensor muscle, those of the
coxa and basi-ischium arising from the endophragmal skeleton and being
inserted by conspicuous tendons, those of the remaining joints arising each
from the joint preceding that upon which it is inserted. The exopodile has
flexor and extensor muscles for the stem, which they enter from the coxa;
but the Hagellum has an extensor only, and must be brought back into the
flexed position by its elasticity. The epipodite possesses muscles, but its
most important movements are probably those which it carries out passively,
under the action of the powerful muscles of the coxa.

The action of the flecors and extensors of the coxa is to draw the whole
limb to and from the normal position, in which its median edge meets that
of its fellow on the middle line. In moving outward it passes a little
ventrally, so that it clears the sub-branchial region of the carapace. This
movement at the same time uncovers both the mouth-field and the inhalent
opening of the gill-chamber, and sweeps the epipodite under the gills, as
IT will explain later. The movements of the basi-ischium are more
complex. They may be analysed into: (a) a movement to and from the
middle line independently of the movement of the coxa, (b) a dorso-ventral
rotation, like the letting down of a flap, (c) a rotation about a longitudinal
axis on the outer side of the limb, like the opening of a door, (d) a divarica-
tion of endopodite from exopodite. This latter movement must be due to a
contraction of the extensor muscle of the exopodite simultaneously with that
of the flexors of the basi-ischium, for this bears the exopodite and tends to
earry it in its own direction. The other movements of this joint are more
difficult to understand, since they seem to take place with a good deal of
independence, and the muscular apparatus does not appear to be adequate
for this. Actually, they are performed together, combined into a single
sweeping movement whose direction is decided by the form of the articula-
tion between basi-ischium and coxa, but a compensating movement of the coxee
can keep the edges of the two limbs together, so that they are let down as a
flap without at the same time necessarily opening as a pair of doors, though

MOUTH-PARTS OF THE SHORE CRAB. 119

their planes are now directed obliquely and meet ventrally at an angle. The
articulation is, moreover, not absolutely rigid, and the movements may there-
fore perhaps be modified by the small additional flexor muscle described by
Pearson. The merus, of course, shares the movements with the basi-ischium,
but it is capable of a moderate amount of rotation upon its articulation with
the latter. The rotation is oblique, from within outwards and at the same
time dorso-ventrally. It is not conspicuous in the living crab, but it is used
sometimes to lower only the distal half of the operculum, and sometimes to
give free play to the movements of the palp. The joints of that organ can
moye upon one another, but it is generally flexed or extended rather stiffly
asa whole. It is so articulated that in extension it moves upwards as well
as forwards. The flagellum of the exopodite is flicked outwards and inwards
with very great rapidity, so that at times it cannot be followed by the eye.
I have already alluded to the movements of the epipodite.

2. The Second Mawilliped resembles the third in general plan, but differs
from it greatly in appearance on account of the following features:—It is
stnaller, and its cuticle is thinner. In the endopodite, the basi-ischiopodite is
short and its components are easily recognizable, the merus is very long, and
both, though flat, like the rest of the limb, are narrow. Actually they are
narrower than the last three joints, which are more flattened than those of
the third maxilliped. The ewvopodite is as large as that of the third maxil-
liped, and much longer than its own endopodite. It is grooved to fit against
the edge of the branchiostegite. The epipodite is not hinged to the coxa,
and a supporting rod of chitin runs from that joint along its anterior edge.
It bears a long podobranch, and its distal part enters the gill-chamber,
passes between its own arthrobranch and the anterior arthrobranch of the
third maxilliped, and lies, like the epipodite of that limb, within the gills.
The second maxilliped has the same relations to the sternum and epimeral
region as the third, but is not connected with them by articulations like
those of the latter limb.

The second maxilliped is not outlined with hairs like the third, but a good
deal of long hair is developed upon it, especially on the hinder face of the
basal joints, on the epipodite, along the outer and towards the end of the
inner edge of the merus, at the end of each of the last three joints, and on
the outer edge of the exopodite. The end of the last joint bears about eight
very strong spines. The inner edge of the merus is not toothed. On the
basal joints, and on the exopodite, the hairs are feathered.

The musculature of the second maxilliped resembles that of the third, but
the limb is more mobile.

3. The First Maailliped is a thin, flat limb, covered for the most part with
delicate cuticle, from which the tajority of the articulations have dis-
appeared. The cowa is a short, broad region, whose entity is established by

120 DR. L. A. BORRADAILE ON THE

the presence, on the anterior side, of a rod-like sclerite, which runs across it
and at its median end expands into the stout cuticle of a large, subpyramidal
endite. The apex of this ondite is directed dorsally, towards the mouth, and
that side of it which is opposed to the sternum forms a facet, above the
membranous attachment of the limb. At its outer end, the transverse sclerite
articulates with the stout cuticle at the base of the epipodite, and that in turn
makes articulation by means of a hard process with the epimerite. I have
net been able to find a true articulation on the inner side of the limb. The
epipodite, which lies outside the gills, is much expanded at its proximal end,
especially in a triangular forward lobe. A strong tendon runs along it.
The basipodite is represented by a soft region distal to the coxa. It bears on
the median side a large, flat, forwardly projecting endite or lacinia, which
has a curved outer and a straight’ median edge, and is a little concave
dorsally (anteriorly), so as to fit over the surface of the mandible. Distally
the basis bears the exopodite, which resembles that of the second maxilliped,
and the endopodite, which consists of an unjointed shaft, flattened in a plane
which faces obliquely outward and inward, with a triangular distalexpansion,
flattened in an almost horizontal plane and separated from the shaft by a not
very flexible suture. This expansion completes in front the exhalent channel
of the gill-chamber. Its median edge is turned ventrally and lies against
the anterior side of the mandible.

The limb is outlined with stout cuticle, which also passes along the sutures
between the protopodite and the exopodite. endopodite, and endites. Its
relation to the sternal and epimeral regions resembles that of the second
maxilliped, but at the base of the epipodite there lies, as has been men-
tioned, a rather ill-developed articulation, which is not present in the limb
behind it.

A row of long, flexible hairs follows the outer edge of the exopodite, the
distal and inner edges of the endopodite, and the outer edge of the large
endite. On the inner edge of this endite the hairs are more numerous,
shorter, and rather stouter. The proximal endite is covered with long hairs,
all directed towards the mouth. On the exopodite, the inner edge of the
endopodite, and the outer edge of the large endite the hairs are feathered.

Flexor and extensor muscles of the coxa move the limb asa whole. The
exopodite has a musculature like that of the third maxilliped, and the
endites have muscles of their own. The epipodite has a system of three
powerful muscles.

The limb can be moved to and from the middle line of the body, doubtless
by means of the muscles of the coxa. In moving outwards it also travels a
little backwards. The large endite makes independent movements, rotating
forward-outwards and inward-backwards and thus with its edge describing
an ellipse. Usually it moves inward-backwards towards the mouth when
the rest of the limb is rotating outward-backwards, and thus the smaller

MOUTH-PARTS OF THE SHORE CRAB. 121

endite, which projects towards the mouth, is being drawn away from the
latter. When the smaller endite comes forward into the mouth the larger
endite moves away from it.

4. The Mavilla (Second Mavilla) is a very broad limb, flattened in its
distal parts, but with a swollen nucleus representing the main part of the
protopodite. This region bears on the outer side the large, quadrant-shaped,
slanting exopodite or scaphognathite, distally the roughly-ogival, peaked
endopodite, and internally the two cleft lobes, representing four endites,
which characterise the maxilla of the Decapoda. I have already (5) sug-
gested that the first of these endites belongs to a precoxal segment of the limb,
the seeond to the coxa, and the third and fourth to the basis. The first cleft
lobe is in Carcinus reduced to a pair of narrow ribbons. The limb is covered
with a delicate cuticle, thickened in places to form certain sclerites. Across
the protopodite there runs on the ventral (morphologically posterior) face,
from the junction between the cleft lobes to the base of the limb at its outer
side, a raised ridge, strengthened by a sclerite which probably represents the
coxa. At its outer end this abuts upon a longitudinal piece which caps the
side of the swollen nucleus of the limb, and which is sutured to, but not
definitely articulated with, the epimerite. More distally, the protopodite is
crossed by an irregular M-shaped, transverse ridge, strengthened by two
sclerites, each of the form of an inverted V, which are hinged to one another
where they meet at the apex of the M. This appears to represent the distal
limit of the basis. On the dorsal (anterior) face of the limb a longitudinal
ridge, strengthened by a sclerite which roughly corresponds in position to
the outer member of the M on the opposite face of the appendage, partly
separates exopodite from protopodite. On the anterior face of the scapho-
gnathite are two sheets of stout cuticle, with thickened edges, which support
it and provide for the insertion of the accessory muscles. The maxilla is
seated upon the membranous body-wall, in front of the sternum, upon which
it does not abut. It is attached by membrane only, except at its outer end,
where, as mentioned above, one of its sclerites is joined by a flexible suture
to the epimeral region.

The hairs which in a prawn fringe the whole edge of the scapbognathite
are here found at its anterior and posterior returning edges only. As in the
prawn, they are feathered. The endopodite is fringed with long, silky hairs,
but these are feathered only at the base of the outer edge. The ends of the
cleft lobes bear relatively short, and, for the most part, simple hairs.

The swollen base of the maxilla harbours a complicated and relatively
powerful musculature. According to Pearson (13), whose account is, I think,
applicable to Carcinus, there are in Cancer two extensors and two flexors of
the coxa, and four “extensors” and four “flexors” of the scaphognathite.
It would perhaps be preferable to call these muscles abductors and adductors
respectively, since they draw the limb downwards from the side of

.

122 DR. L. A. BORRADAILE ON THE

the body and upwards towards it. They arise from the endophragmal
skeleton, and are inserted on the base of the scaphognathite. In its anterior
part, the scaphognathite is crossed, at right angles to its long axis, by half-a-
dozen bands of muscle, which, pulling downwards, curve its surface. These
are the “accessory muscles.” They arise from the sclerites of the basal
portion of the limb, and are inserted on the anterior of those which support
the scaphognathite. A feeble strand of muscle runs across the base of the
endopodite to the second cleft lobe, but independent movements of these
structures, if they take place at all, are insignificant.

The movement of the maxilla consists in a flap ee of the scaphognathite
to and ‘from the roof of the exhalent passage of the gill-chamber. It is
carried out in two ways, according as the current is being directed backward
or forward. To drive the water forward, the posterior and outer end of the
scaphognathite is smartly applied (presumably by the ‘outer flexors”) to
the roof. Then the accessory muscles, which have been keeping the organ
curved, allow the flexors to bring the remainder of it into the same position,
with the undulating movement described by Garstang (8). Finally, it is
drawn downwards by the extensors, being at the same time curved by the
accessory muscles, so that it becomes concave towards the roof. To drive
the water backward this procedure is reversed. The bending of the scapho-
gnathite which it involves is facilitated by the fact tnat the two supporting
sclerites of that organ are united by a flexible region. It seems possible
that the function of the muscles of the coxa is to hold firm the protopodite
when the accessory muscles contract. The lobes of the median edge of the
limb may be seen to be drawn passively to and fro you each stroke ot the
scaphognathite.

5. The Mawillule (First Maxilla) is a small limb, flattened and ead to
fit against the surface of the mandible, and composed of three inwardly-
directed lobes—the inner and outer lacinize and the endopodite—and an
external basal portion which unites them. It is usual to regard the laciniz as
representing the coxa and basis, but, as I have elsewhere argued (5), the prowi-
mal lacinia is probably the gnathobase, or endite of the true first segment of
the Crustacean limb, “the ‘precoxa’ or ‘pleuropodite,’ which may or may not
have originally existed as a free joint in every biramous limb, but has now
nearly always disappeared, either by fusion with the trunk or with the second
joint (coxa or coxopodite), or perhaps sometimes by excalation.” In the
maxilla it is represented by one of the components of the first cleft lobe.
This lacinia is narrow and strong, and curves backward and dorsally to enter
the mouth, at the hinder end of the mandible. Its base is widened and is
continued across the face of the limb as a ridge, covered partly with thin
cuticle but supported by a sclerite, and projecting anteriorly. Towards its
outer end this ridge turns backwards (posteriorly), with a suture in the
sclerite which supports it, and dies away upon the stout articular sclerite,

MOUTH-PARTS OF THE SHORE CRAB. 123

with which its sclerite makes a narrow articulation. ‘The second segment of
the limb, the true coxa, is represented by the region which connects the two
lacinie. For the most part it is covered with soft cuticle, but its outer side
is strengthened by a stout bar, the articular sclérite just alluded to. This runs
along the outer side of the basal portion of the limb. Proximally it articu-
lates with a hard piece upon the epimeral region; near this it is joined by
the sclerite of the precoxal ridge; distally it has a swelling with which the
sclerite of the basipoditic (outer Jacinial) ridge articulates. On this swel-
ling stands a tuft of very long sete, which may be called the “ coxopoditic
setee.”” The outer lacinia is much larger than the inner, and its end is
expanded and has roughly the same shape as the large distal endite of the
first maxilliped. Like the inner lacinia it is prolonged as a ridge across the
face of the limb, but the ridge has no projection of its anterior edge and its
sclerite makes a better articulation with the articular sclerite. It represents
the basipodite. The endopodite has a wide base and a narrow, strap-like,
~ blunt-ended continuation, separated by a suture. When the limb is in situ,
this strap passes in a remarkable way over the shoulder of the mandible,
lying in a notch, much as the endopodite of the maxillule of the Prawn is
carried in a notch on the edge of the metastoma.

The maxillule is attached in a little depression of the body-wall imme-
diately behind the mandible. Like the maxilla it does not abut upon the
sternum. It has an external but not a median articulation.

Besides the coxopoditic setee, which are very long, strong, and thread-like,
the limb bears on the outside of the endopodite a small patch of feathered
hairs, at the end of the endopodite another such patch, and on the inner edge
of the endopodite a fringe of long silky hairs. The lacinize bear on their
free ends a mass of very strong spines, which on the tip of the inner lacinia
are curved in the same direction as the lacinia itself. The cutécle of the
endopodite and basal region is thin, but on the lacinie and their ridges and
the articular sclerite it is of considerable strength.

Pearson (13) describes in the maxillule of Cancer four muscles—two outer
and two inner, a “flexor” and an “extensor” in each pair, the flexors
arising from the protogastric region of the carapace and the extensors from
the endopleurites of the endophragmal skeleton: In Carcinus six strands
of muscle enter the maxillule. Of these, two are inserted on the articular
sclerite, one at each end of the sclerite of the outer lacinia, and two on the
sclerite of the inner lacinia, at the base and near the middle. ‘his set of
muscles appears to be adapted to move the whole appendage inward and
outward, and to rock each lacinia to and fro in an antero-posterior direction.
There is also a band of muscle running from the sclerite of the outer lacinia
to the middle of the broad part of the endopodite. I believe that this when
it contracts pulls the lacinia forwards towards the endopodite, which is firmly
strapped to the mandible by its narrow end.

124 DR. L. A. BORRADAILE ON THE

The lacinize are capable of independent movement. The outer moves to
and from the middle line and also forwards and backwards in the body, and
it combines these movements in varying degrees. The inner moves in and
out of the mouth.

6. The Mandible is a short limb, whose proximal region or body is widened
athwart the body of the crab and very strongly calcified. This region,
which may represent either the coxa or precoxa, but is more probably both
combined, is divided into two portions which it is convenient to know as the
“head” and the ‘‘apophysis.” The junction between these is marked by
deep notches on the anterior and hinder sides and by an oblique, inflexible
suture which joins them. The apophysis appears to run deep into the body,
but it is in reality not interna] like the pieces of the endophragmal skeleton
but an external structure, the true base of the limb, which pushes in the
membranous body-wall till it comes to lie in a deep, close-fitting pocket. It
presents to this pocket a convex anterior face and to the interior of the body
a deeply concave posterior face. Upon its edges are inserted the muscles
which move the limb. The head is much deeper from before backwards than
the apophysis, and convex ventrally. It expands towards tke middle line,
where it presents to its fellow a sharp cutting or ‘‘incisor” edge, in the
middle of which is an obsolescent tooth. Dorsal to the cutting-edge, in its
concavity, is a low mound, the “ molar process.” Hxternal to this, also on
the dorsal side, is a process with an outwardly facing concavity whiclr
articulates with a knob on the epistome. Hxternal to the articular process,
and still on the dorsal face of the limb, is inserted the palp, an inwardly-
curved structure, which should be composed of three joints but actually has
only two, because the first and second have fused. In the normal position it
is folded back above the limb and almost hidden. The joints of the palp are
flattened, the second more than the first. The limb articulates in front with
the epistome, and behind is flexibly sutured to the sclerite which supports

ce

the metastoma.

External to the base of the palp is a group of feathered hairs. The palp
itself is bordered with rather long hairs, which are feathered at its base -but
stout and simple at its apex. There is also a patch of sparse hairs on the
ventral face of the apophysis.

The musculature of the mandible resembles that described for Cancer by
Pearson. There are four muscles. A large outer adductor, inserted by a
broad tendon upon the outer angle of the apophysis, arises from the subhépatic
region of the carapace, and by its contraction must pull the end of the
apophysis downwards and so bring its cutting-edge upwards and inwards
against that of the other mandible. An inner adductor arising from the
dorsal carapace is inserted by a very long tendon near the inner end of the
mandible, which it must pull directly upwards. Two abductors, inserted
respectively near the outer angle and on the posterior border of the

-MOUTH-PARTS OF THE SHORE CRAB. 125

apophysis, will pull that part of the limb upwards and so rotate the cutting-
edge downwards and outwards.

The sole movement of the head of the mandible is a rotation upon its
articulation which alternately parts the incisor edges, opening them like a
pair of doors, and brings them together. The palp is extended and flexed,
digging its apex into the space between the incisor edge and the flank of the
labrum.

7. The Metastoma isa fleshy structure which lies behind the mouth, between
the mandibles and the maxillules. It has two forwardly-directed lobes,
the paragnatha, which stand wide apart against the mandibles, covering the
notches on the hinder side of the latter between the head and the apophysis.
The paragnatha are joined by a low transverse cushion, a little raised in the
middle, on the border of the mouth. The whole is supported on each side
by a selerite which follows its base and at the end is sutured to the mandibles.
The paragnatha contain some glandular tissue, but are not muscular and
appear to be moved only passively.

8. The Labrum is a large, fleshy lobe which forms the anterior border of
the mouth. Its base is rounded and swollen in front but narrows behind,
where it projects into the mouth. Distally it is produced behind into a
nose-like process which overhangs the mouth. Its exposed (ventral) wall is
strengthened by a triangular sclerite. Its sides are moulded to fit closely
against the dorsal faces of the mandibles when ithe palps are flexed against
them. It contains, besides'a good deal of glandular tissue, much muscle,
notably in two longitudinal bands. Its movements are hard to observe, since
it has not so sharp an outline as the limbs and is only exposed when the
mandibles part, but they appear to consist chiefly in a tucking of the nose
into the mouth and its withdrawal.

9. The Mouth lies above the hinder end of the mandibles. It is a longi-
tudinal slit, which forks in front owing to the projection into it of the base
of the labrum, and behind owing to the low median prominence of the meta-
stoma. To it converge all the surrounding structures—the nose of the
labrum, the mandibular palps, the inflected hinder angle of the incisor edge
of the mandible, the inner lacinia of the maxillule, and the pyramidal endite
of the first maxilliped. All these are so formed as to make entry to the
mouth easy but egress from it very difficult.

III.

1. The functions of the mouth-parts are threefold. They subserve respira-
tion by keeping a stream of water flowing through the gill-chamber and
hindering particles from lodging upon the gills ; they subserve alimentation
by tearing up the food and thrusting it into the mouth; and one of them,
the third maxillipeds, cleans the eyes, antennules, and antenne.

2. The branchial chamber of the crab is no less complex and specialized

126 DR. L. A. BORRADAILE ON THE

than the rest of the organisation of that animal. In its widest sense, the
term may be applied to the whole of the very large space that lies between
the flank of the body and the over-arching fold of the carapace known as the
branchiostegite, which encloses the chamber above, and without, and partly
below. The flank itself constitutes the inner wall of the chamber. Of the
two layers which compose the branchiostegal fold, the outer is hard and
calcified, that which is towards the chamber is membranous. In their hinder
region the two layers are not widely separated, and there are between them
only blood-vessels and connective tissue; but anteriorly certain viscera
intrude into the upper part of the fold. The hard layer of the branchio-
stegite lies at first almost horizontal, so as to form a roof above the chamber
and the intruding viscera. It is then turned downwards at an angle, which
in the hinder region is obtuse but becomes more and more acute as it is
followed forwards, to form a wall which curves inwards till it reaches the
flank of the body above the bases of the limbs. As the angle at which this
wall joins the roof becomes more acute, the wall faces more downwards, till
in the fore part of the body a portion of it becomes horizontal; there it forms
a floor to the chamber. The shape of the chamber is of course determined
not by the contour of the outer layer of the branchiostegite, but by that of the
inner. This layer makes in the hinder part of the thorax a continuous curve
‘from above downwards, constituting an arched inner roof to the chamber,
which has here no floor properly so called. In front, however, where the
outer wall becomes horizontal, the inner wall turns inward at an angle to line
the floor of the chamber. In the thoracic part of the latter, the floor under-
lies the overhanging part of certain gills which project considerably above
the bases of the limbs. In the head, it closes from below the exhalent
chamber shortly to be described. A number of “ dorso-ventral muscles ”
enable the membranous roof to be raised or lowered.

The chamber is sharply divided into two parts—a true
the thoracic region, and an “exhalent passage” or “ prebranchial chamber”
in the cephalic region. Of these divisions the gill-chamber is much the larger
in every dimension. Its inner wall is battered back so as to face upwards
as well as outwards, and is brittle, though thin, and composed of broad ribs
—the so-called “ epimera ”—one to each limb from the last leg to the second
maxilliped inclusive. That which lies above the cheliped is larger than the
rest and prominent. Those behind it face a little backwards and those in
front face forwards, so that the whole wall is convex outwards, forming a
low, roughly half-conical mound. In front, the chamber narrows rapidly,
its membranous roof at the same time falling steeply to the hinder opening
of the exhalent passage, where the roof is upheld, and the opening maintained,
by an arching, calcified sclerite. Here the roof turns forward as that of the
exhalent passage. his isa shallow chamber which diminishes in width as

“

gill-chamber ” in

the carapace narrows forwards. Its inner wall has become merged in the

MOUTH-PARTS OF THE SHORE CRAB. 127

roof, so that it has only roof and floor. Shallow though this channel is, the
effective depth of its entrance is lessened by a double barrier. One com-
ponent of this is the epipodite of the first maxilliped, which, sweeping round
on its spiral course, roughly parallel with the epipodite of the third maxilliped
but starting in front of the gills and lying upon their outer surface, crosses
the portal of the exhalent passage in such a way as to bar it from behind and
from outside and leave access to it only from above and from within. Since
the wide base of the epipodite does not merely touch the floor with its edge
but lies flat against it, the movements by which the tail is caused to travel
over the gills probably never separate the base from the wall of the passage.
The other component of the barrier is the ‘‘ branchial ridge” of Pearson,
a fold of the membranous layer of the floor, parallel with the anterior side of
the principal inhalent opening but at a short distance from it, which fits
between the podobranch of the second maxilliped and the epipodite of the
first and helps to enable the latter to make effective contact with the floor.
On the inner side of the exhalent passage, the edge of the branchiostegite
does not meet the body closely at the bases of the limbs, but leaves there a
long, narrow gap. This is normally filled by the endopodite and exopodite
of the first maxilliped and the exopodite of the second, which are moulded
longitudinally to fit together and against the mandible and to receive the
branchiostegite. Further forward, in front of the mandible, the body-wall,
which is here the hinder part (endostome) of the epistome, is concave, and
so falls away dorsalwards from the branchiostegite and widens the gap.
Thus is formed the ewhalent opening. It is bordered behind, and contracted,
by the expanded end of the endopodite of the first maxilliped, and discharges
forwards and towards the middle line, so as to direct the current of its side
to the antenna and eye ot the opposite side.

The gills of Carcinus are nine on each side. The second maxilliped has a
podobranch and an arthrobranch, the third a podobranch and two arthro-
branchs, the cheliped two arthrobranchs, and the first and second walking-
legs each a pleurobranch. WNach gill is a tapering structure. Hxcept the
podobranch of the second maxilliped, which lies horizontally and is directed
backwards at the bases of the maxillipeds in front of the principal inhalent
opening, the gills are turned upwards and inwards, and lie against the inner
walls of the chamber, converging to its highest point, just before the roof
falls in front. They are closely applied to one another and separate a
shallow ‘‘hypobranchial space” against the inner wall from an ‘“ epi-
branchial space” under the branchiostegites. The gills are phyllobranchs.
Each of them, save the arthrobranch of the second maxilliped, is heart-
shaped in transverse section, owing to the fact that the leaflets project as
lobes above the axis but die into it below. Thus it comes about that where
two gills tie side by side there is between them on the under side a “ hypo-
branchial channel,” and above each of them is an “ epibranchial channe] ”

128 DR. L. A. BORRADAILE ON THE

along the axis, between the leaflets. The arthrobranch of the second
maxilliped has leaflets on the posterior side only, the anterior side, which is
applied to the wall of the chamber, being flat; but along its edge an
epibranchial channel runs as a gutter. The hypobranchial channels are part
of the hypobranchial space and are in communication with one another
under the gills, especially at the bases of the latter, which are there a little
arched, leaving a longitudinal corridor in which lies normally the epipodite
of the third maxilliped. The epibranchial channels are part of the epi-
branchial space, and will communicate over the gills unless the roof be
lowered on to the latter. At the hinder end of the chamber there is a space
which contains no gills. This space is very shallow and in it lies the
enigmatical fold of the body-wall known as the “ pericardial lobe.” It is
shielded from the current entering over the last leg by a ridge which, as I
shall presently show, directs the water forwards and downwards, and it
probably plays no important part in the circulation of the water about the
gills. The branchiostegite fits closely against the bases of the gills, which
are flattened back to receive it. Above most of the rest of the surface of
the gills the epibranchial space is probably asa rule deep enough to allow
the epipodite of the first maxilliped to play freely, but by the action of the
dorso-ventral muscles varies in depth from time to time owing to circum-
stances of which nothing is known, and which may be related to other
functions than respiration. At the anterior end of the chamber, where the
arthrobranchs of the third maxilliped and clieliped face partly forwards
towards the roof as it falls to the opening of the exhalent channel, there is
a deeper part of tle cavity. Since its roof is flexible like that of the rest of
the epibranchial space, it is possible that this does not always exist, Lut I
have always found it, and I believe that it is kept in being, in its lower part
at least, by the attachment of the roof to the arched sclerite that I have
mentioned. It slants downwards, forwards, aud outwards to the opening of
the exhalent channel. From it water must be drawn by the action of the
scaphognathite into the exhalent channel, which, as I have shown, cannot
receive water from the region directly behind itself. Thus the water from
all parts of the chamber must pass through this space, and, though it is not
sharply defined from the rest of the epibranchial space, it may be dis-
tinguished as the “collecting space.”

The gill stream*. The water which bathes the gills normally makes entry
under the edge of the branchiostegite in the thoracic region and leaves in
the same way in the preoral region. According to the classical account of
this process, given by Milne-dwards (12), the entry of the water takes place
only by an opening which lies in front of the coxa of the cheliped and

* It was not till I had written the paragraphs on this subject that Isaw the paper of
Mr. R. K. S. Lim, published in 1918 (10). As my work extends as well as confirms
that of My. Lim, I have left unaltered what I had written. '

MOUTH-PARTS OF THE SHORE CRAB. . 129

external to that of the third maxilliped. Other authors (Bell (1), Giard (7),
Bohn (2, 4)) have stated that the water enters above the legs along the whole
length of the carapace behind the third maxilliped. Pearson, however (18),
finds that in Cancer entry is made principally by an anterior opening, which
is that of Milne-Hdwards, and secondarily by a posterior opening above the
last leg, but not between these apertures. An examination of the structures
which adjoin the edge of the carapace in Carcinus shows (1) a relatively
large opening in the position indicated by M'lIne-Edwards, (2) a smaller
opening above each leg, the cheliped included. The openings above the first
three walking-legs are separated from one another by meetings between the
prominences upon the flank with which the legs articulate and corresponding
prominences of the branchiostegal edge. They are longitudinal, and slit-like
but well formed. That above the last leg is very narrow, and imperfectly
separated from the one in front of it *, and that above the cheliped is separated
from the opening of Milne-Hdwards only by a close fitting of the branchio-
stegite to the coxopodite when the limbis turned forwards. Access to each of
the openings above the legs is obtaiaed principally between its coxopodite and
that behind it, and by moving the leg backward and upward the crab can
almost close it, though at the same time the approach to the orifice above the
leg next in front is more widely opened. A large hairy tract on the under
face of the branchiostegite has no doubt the function of filtering the water
which is drawn through it towards the inhalent openings, and especially that
of Milne-Edwards. It rises into a long fringe around a bare patch where the
cheliped lies against the branchiostegite, and is there met by fringes on the
borders of the cheliped so as to form what is probably a very efficient guard
for Milne-Hdwards’s opening. Along the edge of the branchiostegite another
long fringe forms a similar protection for the openings above the legs. By
placing with a pipette against each opening a little carmine suspended in
sea-water, it may be seen that water enters at all of them, even when the
third maxillipeds are opposed so as partly to close with their epipodites
Milne-Hdwards’s opening, and that the water from any opening takes a little
Jonger to reach the exhalent orifice than that from the openings in front of
it. When the third maxilliped is divaricated from its fellow, so as to open
widely the aperture of Milne-Hdwards, water enters there more freely,
passing forwards as well as backwards, since the anterior side of the opening
is now uncovered, and bathing very copiously the podobranch of the third
maxilliped, which has been drawn backward into the full stream. hat

* The crevice above the last leg is continuous with that between the hinder edge of the
carapace and the first abdominal segment. Narrow though this opening is, the state of the
hairs which line it shows that a little water entersit. No doubt this water flows both
ways towards the two gill-chambers. The occasional presence of alittle mud in the middle
chamber which is connected with the hinder crevice of the carapace proves that water must
enter it also, though probably it does not there perform any important function.

LINN, JOURN.—ZOOLOGY, VOL. XXXV. i)

130 DR. L. A. BORRADAILE ON THE

podobranch may thus have a greater physiological importance than its size
would suggest, though it seems doubtful whether in its more isolated position
the water would be drawn between its leaflets. The divarication of the third
maxilliped also exposes an opening between it and the limb in front of it
which helps to admit water to the fore part of the chamber.

The course of the water within the gill-chamber is more difficult to follow,
but there can be no doubt that on entering, it passes wnder the gills, that is
into the hypobranchial space, then comes outwards between the gill leaflets
into the epibranchial space, and finally flows by way of the collecting-space
into the exhalent passage. That practically all the water which enters the
gill-chamber takes this course, and does not pass directly from the exterior
to the epibranchial space, I am convinced by the following considerations :-—
(1) It is exceedingly unlikely that the blood in the leaflets of the gills is
exposed to the stream of water only on their edges, and that their flat
surfaces are dependent on eddies or diffusion for the renewal of the water in
contact with them. (2) When carmine is caused to enter through any of
the openings it is found principally, and the large particles are always found,
underneath the gills. (3) The disposition of the parts is such as to suggest
that the current flows in the direction that I have described. From each
of the openings behind that of Milne-Edwards the shortest route to the
exhalent passage leads under and through the gills. If there were a wide
space between the gills and the border of the branchiostegite, the resistance
due to friction with the leaflets might cause the water to pass over the gills,
but actually the branchiostegite fits, as has been said, close against the gill-
bases. In front of the cheliped the relations of the parts are different.
Here, when the third maxilliped is in its normal position, its coxa and
epipodite bar the passage of the water forwards and direct it inwards under
the gills; but when the maxilliped is divaricated, the shortest route to the
inhalent channel would be, were it not for the barrier formed by the
epipodite of the first maxilliped, directly forward. It would then pass partly
through the podobranch of the second maxilliped, but in great part between
that structure and the floor of the chamber or the bases of the other gills, and
would not be distributed to the latter organs, which lie above the direct
course. Actually, however, the existence of the barrier mentioned must
prevent the water from being chiefly drawn in this direction and cause it to
circulate through the upper gills. That the water does actually take the
route through the gills can easily be seen by cutting a window in the floor
of the exhalent passage and placing carmine in Milne-Hdwards’s opening.
The carmine will be found always to pass under the gills and to reappear
above the scaphognathite, not to take the direct route.

The water which enters above the first and second walking-legs flows
forwards and inwards along a shallow gutter hollowed on the epimerite tld it
reaches the entry of a hypobranchial channel, between the pleurobranch of its

MOUTH-PARTS OF THE SHORE CRAB. 131

leg and the gill which adjoins it in front ; and through this it passes into the
hypobranchial space. The water which enters the opening behind the
cheliped flows similarly forward over the arthrodial membrane, which is
shaped to provide a kind of duct for it, to the entry of the hypobranchial
channel between the, anterior arthrobranch of the cheliped and the posterior
arthrobranch of the third maxilliped*. At this very large portal of the
hypobranchial space it meets the water from Milne-Edwards’s opening.
From. that aperture the current passes, when the third maxillipeds are
opposed, obliquely backwards through the portal just mentioned, though a
little may stray forwards to pass between the arthrobranchs of the third
maxilliped. When, ‘however, the maxilliped is divaricated and the opening
thus fully uncovered, water passes also to the channels between the arthro-
branchs of the third and second maxillipeds, and to the podobranchs of those
limbs. The water which enters the opening behind the last leg is prevented
from flowing directly forwards by a ridge on the epimeral region, passes over
the articulation of the last leg, which is not prominent like the others, joins the
stream which enters between the last two legs, and reaches the hypobranchial
space by passing forwards through a definite entrance formed by the shaping
of the hinder leaflets of the last gill at a spot near its base and at the end of
the longitudinal corridor mentioned above. The arrangements may be
summed up by the statement that the water which enters over each limb
passes into the hypobranchial space by an opening between its gill and the
gill of the limb in front of it, except that most of the water entering above
the third maxilliped passes behind its gills.

In each hypobranchial channel, the water flows, I believe, upwards, mingling
to some extent under the gills with that in the adjacent channels, and
diminishing as it goes by loss between the leaflets of the gills to the
epibranchial space. Since this loss is hindered by friction against the
leaflets, the water does not all escape till the top of the gills is reached, and
is thus distributed over as wide a gill-surface as possible.

The course of the water over the outer surface of the gills presents a very
difficult problem. The shape of the chamber gives no convincing indication
of the direction in which the stream gets through it, and its size, asa whole
and in each part, depends upon the changing form of its roof. I can only
offer some suggestions on this subject.

Normally, the water will pass direct from the point at which it issues from
between the leaflets of the gills to the collecting-space. If, however, the
roof be lowered on to the surface of any part of the gills, the water which
is passing through them is probably got away by the epibranchial channels.
The appearance of these structures strongly suggests that a current flows

* The entry to the hypobranchial channel between the arthrobranchs of the cheliped is

very small, and that channel probably receives its water from the longitudinal corridor.
Ox

132 DR. L. A. BORRADAILE ON THE

along them. ‘They widen from above downward, and it would at first seem
as if the current must flow in that direction. But (1) as each channel widens
it grows shallower, so that its capacity is no greater, but rather less ; (2) its
lower end is probably closed by the edge of the branchiostegite, and in any
case leads to a part of the chamber that does not directly communicate with
the exhalent passage; and (3) the direction of the leaflets is such as at the
lower end to cast the water into the channel in an upward direction. It is
therefore probable that the current flows upwards. On the other hand, near
the upper end the channel narrows so much that its capacity is greatly
diminished, and here the water must be overflowing from it into the space
over the gills, which is near the top of the collecting-space. The grooves
between the convex surfaces cf adjoining gills, which may be known as
‘““interbranchial channels,” form a similar system.

All the water enters the exhalent passage from above and within—that is,
from the collecting-space. This is due partly to the fact that, as has already
been shown, access to the passage from behind is barred, and partly to the
fact that the stroke of the scaphognathite is made from below upwards, against:
the roof ; and, as has also been shown, it has the important effect of causing
the water to flow through instead of over the gills.

The working of the scaphognathite has already been described. The
effective stroke is the upward one, but the downstroke must act upon the
water which has entered the lower part of the passage during the upstroke.
Probably the bulk of this is driven into the collecting-space and a small
portion sent forward to the exhalent orifice.

The stream that issues from each branchial chamber is Tnacied. as has been
said, obliquely across the epistome to the opposite side of the front, and often
it does in fact take that course. But normally it meets in the middle line the
current of the other chamber, and the two deflect one another so that they flow

‘forwards under the antennules. A further modification of the direction of
the current is brought about by the activity of the flagella of the maxillipeds.
Flicking to and fro extremely rapidly, these exert their foree more on
the outward stroke when they are drawn by their extensors in the direction
of their concavity as a cilium moves than when they are returning by their
own elasticity in the direction of their convex sides. They not only reinforce
the current very notably, but also turn it outward, and in particular, I think,
by means of the hairs with which they are fringed they draw away particles
which might otherwise lodge upon the organs of special sense.

‘Lhe regulation of the gill-stream is brought about in two ways—by
alterations in the size of the openings, and by changes in the beat of the
scaphognathite and the exopodites of the maxillipeds. I have already shown
how the flow through the anterior inhalent opening is regulated by the third
maxilliped, and how the legs can close the openings which lie behind them.
The size and form of the exhalent opening must also haye an important

MOUTH-PARTS OF THE SHORE CRAB. 133

influence on the current. ‘The aperture is smallest when the third maxillipeds
are closely opposed. At such times a steady stream issues from the opening
which they leave in the middle of the epistome and flows forwards under the
antennules. When a wider opening is necessary, either to provide for a
greater flow or to allow the current to be directed to the sides of the body,
the operculum may be opened in varying degrees by lowering either the meri
only or also the basi-ischia. At such times the form of the exhalent opening
proper may be modified by alterations in the position of the expanded end of
the endopodite of the first maxilliped. |

The function of the epipodites* is the cleaning of the gills. Moving to and
fro over the gill-surface they brush it, drag over it their long, flexible, barbed
hairs, and thus prevent particles that are brought in by the gill-stream from
settling there and closing the minute passages between the leaflets. The
importance of this function is shown by an observation of Pearson, who
found that ina crab in which the epipodite of one of the first maxillipeds
had been destroyed, the outer surface of the gills of its side was covered with
a layer of fine mud. In one of my crabs there-was a similar deposit of fine
sandy particles. In this case the epipodite was intact, but I have no doubt
that it was in some way paralyzed f.

By an admirable mechanism the three epipodites, between them, reach
almost every part of the gills. The epipodite of the first maxilliped lies
above the gills and sweeps their outer surface. It is probably moved more
by the action of its own powerful muscles than by the excursions of the
maxilliped as a whole, which are not extensive, and if they were so would
interfere with the other functions of the limb. This epipodite is very flexible,
and, doubtless by the action of its muscles, it is kept closely applied to the
rounded surface of the gill-mound while it swings upwards and downwards
over the gills, describing an are with its tip. In an almost vertical position it
stands against the forward face of the mound. The epipodites of the second
and third maxillipeds lie below the gills and sweep the inner surface of the
latter. Both are stiffer than that of the first maxilliped. ‘The principal
movements of each are probably those which it undergoes passively with the
coxa of its limb. These movements can easily be imitated upon a dead
specimen, and they must occur with each of the excursions which the limbs
are constantly making in life. The epipodite of the second maxilliped lies
on the anterior, forwardly-facing region of the inner wall of the gill-chamber.
When the maxilliped swings outwards, pivoting on its attachment, the
epipodite makes a corresponding inward movement over the face of the
thoracic wall, sweeping the inner surfaces of the gills that stand there, with

* The epipodites of the first and third maxillipeds have also the passive function of
directing with their bases the course of the currents in the way that I have already described,

+ Since the foreion matter was above the gills, ithad presumably been brought in during
reversals of the current.

134 DR. L. A. BORRADAILE ON THE

which it is almost or quite in contact. As the maxilliped returns, the
epipodite is drawn back to its former position. The epipodite of the third
maxilliped reaches over the epimeron of the cheliped to lie against the
posterior region of the inner wall. When the maxilliped is in the normal
position the epipodite lies in the longitudinal corridor at the base of the gills.
If the movement of the limb were simply outward, and if the epipodite were
not hinged to it, the effect of its divarication from its fellow would be merely
to press the epipodite more firmly inward against the wall above the bases
of the gills, with which it would therefore tend not to be in close contact.
Actually, however, the maxilliped, as I have stated above, rotates downwards
as well as outwards, and thus moves its epipodite upwards over the flank of
the body, while the movement presently brings the stout base of the epipodite
against the articulation of the cheliped, and this opposition flexes it at its
hinge and directs it outward as well as upward. hus when the maxilliped
is divaricated, the blade of its epipodite moves upwards and presses outwards
against the under surface of the gills, which it sweeps and lifts a little from
the thoracic wall, thereby flushing these parts with water. As the maxilliped
returns to the normal position, the base of the epipodite is pressed against the
arthrobranchs of the cheliped, and thus bent back into the longitudinal
direction. ‘The position of the epipodite of the second maxilliped upon the
forwardly-facing epimera makes unnecessary any such special mechanism to
bring it against the inner surface of the gills.

No doubt the movements of the epipodites have the effect of mingling and
distributing the water in the gill-chamber while they clean the gills, but it is
not clear that this has any such physiological importance as has been
attributed to it.

The variations of the gill-stream are a very striking feature of the
physiology of the crab. They may be studied either by the carmine method
or by watching the movements of the flagella of the exopodites of the
maxillipeds, and, after making a window in the branchiostegite, those of the
scaphognathite. Since the latter method studies directly the working of the
principal agent, it is the most instructive, though it is open to the obvious
objection that the action of the scaphognathite is affected by the operation.
But when the shock of the latter las passed off, its effects are less serious
than might be expected. The most remarkable feature of the action of the
scaphognathite is its extreme sensitiveness. Any rough or sudden handling
of the crab is liable to cause it to stop—in the face of danger the creature
holds its breath. Yet this does not always happen when it is expected.
Other variations are brought about by less obvious causes. The scapho-
gnathites work independently, and either of them may cease working while
ihe other continues. Their beating changes its rate and force from time to
time in the same individual, and differs in different individuals in the same
vessel at the same time. Clearly the causes are sometimes internal: probably

MOUTH-PARTS OF THE SHORE CRAB. 135

they are often external. From some observations of Bohn (4) on the reversal
of the stroke it would appear that the crab is sensitive to changes both in the
oxygen-content and in the carbon-dioxide-content of the water. Whether
these circumstances act by altering the composition of the blood, or through
the sense-organs, or in both ways is not clear. I would hazard a surmise
that the third of these alternatives is correct, and from some casual
observations I suspect that the antennules are sensitive to oxygen. Changes
in the activities of the scaphognathite are accompanied, and its working
facilitated, by actions of the subsidiary parts of the apparatus. Probably in
the normal, quiet breathing of a resting crab, when the water is clear and
perfectly oxygenated, the third maxillipeds are opposed and the flagella of
the exopodites at rest. Most commonly, however, at least in experimental
conditions, the crab sits with its maxillipeds lowered to some extent. This
must have the advantage of lessening the work of the scaphognathite, when
a considerable volume of water is being dealt with, by allowing it to get
away with less friction ; and it also enables the flagella to be brought into
play, both to help the scaphognathite, and to drive the current and its
contained particles outward, away from the sense-organs. Like the seapho-
gnathite, the flagella may work on one side only or on both at once. One,
two, or three of them may work at the same time on each side, and they may
give a single stroke or work continuously for long periods. The flagella of
one side are complemental to the scaphognathite of the other, driving the
current in the same direction, and I have not seen the flagella and scapho-
gnathite of one side both at work while on the other side both are at rest.
Further, the maxillipeds may be divaricated. This admits more water to the
gill-chamber, admits it where it has a relatively short distance to travel and
hence causes relatively little increase in work, and also supplies a set of gills
which are out of the main stream when the maxillipeds are apposed.
Changes in the posture of the crab have also, as Bohn has pointed out, an
effect upon the work of the scaphognathite. When the water is clean and
well oxygenated, a horizontal position is possible, though there is always some
upcast, at least when the maxillipeds are opposed. But when the water is
muddy or foul, the vertical position is necessary to enable reversal of the
current to obtain more oxygen, as will presently be shown. The exhalent
opening is then often above the surface, and work is heavier while the
flow is forward. From time to time the third and second maxillipeds make
violent excursions outwards. This happens more frequently when the water
is not clear, and it has the effect of sweeping the gills with the epipodites.
From time to time also, the secaphognathite reverses its action, and for a
shorter or longer period draws water in at the exhalent opening and drives it
backward through the gills and out at the inhalent opening. ‘This is done
when foreign particles or distasteful substances are being drawn in, and it is
noticeable in carmine experiments when the particles are too coarse. Its

136 DR. L. A. BORRADAILE ON THE

function in this case is obvious. But it also happens at intervals when the
water is quite clear and pure. It probably then drives out particles which
have gradually accumulated under the gills while the current was flowing in
the forward direction and which would in course of time prove harmful.
The reversal of the current, which was first studied by Garstang (8, 9) in
Corystes and other genera, has been examined in Carcinus by Bohn (28, 4),
who has shown that it always takes place, but varies in frequency and
duration with the foulness of and amount of matter in suspension in the
water, and that when the water is chemically foul the crab will raise the
normally exhalent opening to the surface so that it draws in either the better
oxygenated water of the surface or air, which passes out in bubbles at the
opening of Milne-Hdwards after oxygenating the water in the chamber.
These results I can confirm from my own observation. The crabs will live in
exceedingly foul water if they be permitted to raise the front of the body
out of it, but may be asphyxiated by preventing this. Bohn (4) thinks that
the reversal of the scaphognathite has an advantage in resting its muscles.
I am not clear that it is necessary to suppose that it has any other function
than that of cleaning the chamber and enabling the animal to obtain a
better supply of oxygen.

3. In feeding, the crab severs the food into morsels which it swallows
without finely dividing them. It always seizes the food with the chele.
The third maxillipeds then part, and the food is placed by the chelee between
the mandibles. If the mass of it be very bulky, one or more of the legs
may be brought into action to assist in lifting it and thrusting it towards the
mouth. The mandibles part to receive the food, and then close upon it.
They do not cut it by a slicing action or chew it, but unless it be soft enough
for them at once to sever a morsel of it by pressure, they hold it firm while it
is being divided by the action of other organs. If the food be very soft,
they may even sometimes be seen to be held wide apart while it is thrust into
the mouth by the action of their palps and of the nose of the labrum.
Usually, however, the food (I have fed my crabs upon various parts of the
body of fishes and upon meat) requires the assistance of other organs for its
severance. In this case it is torn by being pulled outwards from the
mandibles, much as a crust may be torn by being held with the teeth while
the hand wrenches a part of it away. The outward pull is sometimes given
by the chelee, especially if the mass of food be tough, and they often keep a
hold upon it when they do not appear actually to be pulling ; but most often
the work is done by the third maxillipeds, which grasp the food by pressing
upon it with the toothed inner edges of their ischiopods and at the same time
press downwards upon it with their palps, somewhat as the digit of a sub-
chela is closed. These limbs do not cut or chew the food, and rarely pass
it towards the mouth, but by pulling it outwards tear from it the portion
which is held by the mandibles. The second maxillipeds work in various
ways, and their action, like that of the organs in front of them, is often hard

a
wo
~_

MOUTH-PARTS OF THE SHORE CRAB,

to follow because they are hidden by the third maxillipeds. After the
removal of the latter, movements of the inner jaws may be observed, but I
-have not been able to induce crabs which have been so operated upon to
feed*, and in any ease their feeding would not be normal. ‘The second
maxillipeds apply to the food their last joints, armed with the strong spines
which have been mentioned. With these they sometimes thrust outwards,
aiding the third maxillipeds or, if the morsel be small, even taking their
place. At other times they appear to be thrusting the food inwards, towards
the gape of the mandibles, at others to work up and“down upon it, strained
as it is between the third maxillipeds and the mandibles, so as to cut through
it. It is difficult to discover what is the work of the first maxilliped. The
action of the great lacinize would suggest that they are cutting the strand of
food, but their armature of bristies is so much feebler than the arrangement
of spines upon the dactylopodites of the second maxillipeds and on the lacinize_
of the maxillule as to throw doubt upon the correctness of this impression.
In view of the alternation of their action with that of the pyramidal endites,
it seems possible that they are really brushing the frayed portions and
fragments of the food backwards towards the opening of the mouth, into
which the pyramidal endites help to thrust it. The shape of these endites
and of the hairs upon them are such as to make it easy for the food to pass
between them into the mouth, but difficult for it to emerge. The maxille
are, | think, quite useless as Jaws. Their distal lobes, at least, are feeble,
inefficient structures, drawn passively to and fro with the strokes of the
seaphognathite. J am less certain about the proximal lobes, but if these have
a function I am unable to suggest what it may be. The maxillules are
important and relatively powerful structures. Lt have described the
movements of their laciniz. The outer of these cuts at the food with its
powerful bristles and the inner, I think, thrusts it into the mouth. Probably
in doing so it is cutting or tearing it from the main mass. The paragnatha
appear to have the funetion of closing the notches on the hinder edge of the
mandibles, through which the food might work outwards, and of pouring
upon it the secretions of their glands, whatever be the functions of these.
I have already described the working of the mandibles. I am unable to see
that their “molar ” faces have any grinding action, but they appear to help
to wall in the food which is being thrust into the mouth by the palps, and
especially by the action and powerful nose of the labrum. I have been a
little surprised, in feeding to the crabs portions of fish and meat containing
bone which I should have expected them to be able to crush, to observe that
they were rejected, sometimes after being stripped of the softer tissues. It
is quite possible, however, that in other circumstances these would have been
devoured. After the meal a flurry of the mouth-parts is usually to be seen.

* Possibly reflexes from these limbs are needed to co-ordinate the movements of the
other mouth-parts. A prawn will as readily take food from a pair of forceps as from its
own chele. A crab will not.

138 DR. L. A. BORRADAILE ON THE

By brushing their various hairy surfaces against one another they detach
the small particles of food that are clinging to them, and these are swept
away by strong currents set up by the activity of the exopodites of the
maxillipeds.

The opercular function of the third mauillipeds deserves mention here. The
operculum is not of importance to the crab on account of any part that it
plays in feeding. It is freely open during that process, and is not closed
again till the mouth-parts have cleaned themselves. It undoubtedly protects
the more delicate organs within it, as may be seen by the way in which it is
closed when they are approached by any implement. But the closeness of
its fitting, and its fringe of hairs, indicate that it is also a part of the
respiratory apparatus. The current set up by the scaphognathite is a
wonderfully strong one, partly because it is working in a closed system of
passages, and there are several places in the neighbourhood of the inner
mouth-parts where leakage is probably liable to take place, and throw
unnecessary work upon the scaphognathite in keeping up a current of the
swiftness which is needed. The closing of the operculum prevents this.

4. The hairs which are arranged in so definite a manner upon the limbs
play no insignificant part in the events which have just been described.
They are of an immense variety. A few particulars about them have been
given above, in the course of the descriptions of the limbs, but it is quite
impossible to do justice to this subject without devoting to it a special inves-
tigation. McIntosh (11) has examined their structure in’ detail, but their
functions still remain to be elucidated. The great majority of the hairs are
in some way feathered or toothed, and the lateral members which they then
bear may be of every relative dimension and shape, from the finest filaments
to the coarsest serration, may vary in different parts of the same hair, be
directed at any angle to the axes, or recurved, as on some of the hairs of the
epipodites, and be set on one side of it, on two, or on all sides. Most of the
hairs are placed on edges or surfaces where they meet the water, either
actively in the moyement of the limb or passively by the flow of the gill-
stream. Fyrom their form and position it is clear that they must serve more
uses than one, though whether any of them have a double function is less
obvious. Some of them undoubtedly serve to jilter the water entering
various orifices. his is clear from the particles with which they may often
be seen to be laden. It is notably the function of those which outline the
parts of the operculum formed by the third maxilliped, and on its coxa help
to guard the opening of Milne-Hdwards, as all the openings of the gill-
chamber are guarded by hairs on the carapace and on the coxee of the legs.

Others are sensory. Most if not all of these are probably tactile, that is
give information as to the pressure which is exerted against them, but their
sensibility is used in various ways. It is clear from its behaviour that the
crab is able to detect the presence of particles in any part of its gill-stream

MOUTH-PARITS OF THE SHORE CRAB. 139

and accordingly to stop, reverse, or alter its direction. This can only be due
to tactile organs at various points in the system. Another use of the tactile
sense is involved in the appreciation of the strength of the current, which
the animal must possess. Conceivably this might be due to a muscular sense
of the power used in producing the current, but, as Doflein (6) has shown, it
is very probable that in Crustacea the muscular sense is supplemented, if not
largely replaced, by the information giyen by tactile hairs brought against
the water by the movement of the limbs. In any case, it is likely that this
information is possessed by the crab and guides it in the use of its limbs.
Probably the more delicately feathered hairs are those which are exposed té
and detect finer changes in the pressure of the water. From these there isa
gradation to the coarser ones which are used in filtering. Whether any or
all of the latter are sensitive does not appear.

Whether, again, any of the pairs subserve a chemical sense is doubtful.
Certainly the behaviour of the crab shows that it possesses such a sense, aid
by it is informed of the quality of the water and of the nature of the food.
But it is quite possible that this is due to the antennules. Over these the water
passes on leaving the gill-chamber, and the juices and débris of the food are
rejected towards them, and probably to some extent reach them in spite of or
by permission of the exopodites of the maxillipeds. The crab appears,
though less clearly than a prawn, to taste as well as to smell its food, but
this. may be due to the sensibility of the antennule ; much as in Man, the
aromas of food are appreciated by the olfactory epithelium.

Others of the hairs have a cleaning function. This is exercised in very
different ways by the long threads on the epipodites of the maxillipeds (and
possibly by the coxopoditiec sete of the maxillule, for which I can suggest no
other function), and by the serrated bristles of the palps of the third maxilli-
peds. Those organs are continually busy cleaning the various structures in
front of them. They brush the antennules, sometimes acting singly, some-
times combing an antennule between them. Hach of them brushes the eye
and antenna of its side, and reaches across to clean the mouth-parts of the
opposite side, attending, for instance, to the delicate and probably sensory
hairs of the expanded end of the endopodite of the first maxilliped. In
these activities, different parts of the palp are brought each against a different
organ, and very possibly the difference in the serration of the bristles in
various parts is in correspondence with the structures to be cleaned in the
organs to which they are applied.

The function of current-making must perhaps be attributed to the hairs
on the flagella of the exopodites.

Lastly, the bristles which are used in manipulating the food are special
members of the series of hairs.

140

12.

13.

DR. L. A. BORRADAILE ON THE

BIBLIOGRAPHY.

Bert, I.—A History of the British Stalk-eyed Crustacea. London,
1853.

. Bony, G.—Sur la respiration du Carcinus menas Leach. C. R. Acad.

Sci. exxv. p. 441 (1897).

. Bonn, G.—Sur le renversement du courant respiratoire chez les Orus-

tacés Décapodes. bed. p. 539 (1897).

. Boxy, G.—Des mécanisines respiratoires chez les Crustacés Décapodes.

Bull. Sci. France et Belgique, xxxvi. (1907).
Borrapaite, L. A.—On the Structure and Function of the Mouth-parts
of the Paleemonid Prawns. Proc. Zool. Soc. Lond. 1917, p. 37.

. Doriery, F.—Lebensgewohnheiten u. Anpassungen bei dekapoden

Krebsen. Fest. 60ten Geburtstag R. Hertwigs, iii. (1910).

. Grarp, A., & J. Bonnrer.—Contribution a Pétude des Bopyriens. Trav.

Inst. Lille, v. (1887).

. Garstanc, W.—The Habits and Respiratory Mechanism of Corystes

cassivelanum. Journ. Mar. Biol. Assoc. (n. s.), iv. (1896).

. Garstane, W.—The Systematic Features, Habits, and Respiratory

Phenomena of Portwmnus nasutus Latr. bid. (1897).

. Lim, R. K. §8.—Experiments on the Respiratory Mechanism of the

Shore Crab (Caretnus menas). Proc. Roy Soc. Edinburgh, 1918.

. McIyrosu, W. C.—On the Hairs of Carcinus menas. Trans. Linn. Soc.

London, xxiv. (1863), pp. 79-100, t. 19, 20.
Miitne-Howarps, H.—Recherches sur le mécanisme de la respiration
chez les Crustacés. Ann. Sci. Nat. Zool. (2) ii. p. 129 (1839).
Parson, J.—Cancer. London, 1908.

ig. 15.

ig. 19.

. 16.
palidfe

» Allis),

MOUTH-PARTS OF THE SHORE CRAB. 141

EXPLANATION OF THE PLATES.

PuateE 10.

. A ventral view of a female of the Shore Crab (Carcinus menas). On the left

side of the animal the first and second walking-legs haye been parted, to
expose the approach to the inhalent opening between them, and on the right
side the cheliped has been turned back to reveal as much as possible of the
flange on the coxa of the third maxilliped, by which the inhalent opening of
Milne-Edwards is guarded.

. The forepart of a similar view aftef the third maxillipeds have been divaricated,

one of them ent short beyond its basis, and the chelipeds removed with the
exception of their coxee.

. A ventral (posterior) view of the third maxilliped of the right side, removed from

the body and flattened. The dotted line; show the normal position of the
‘epipodite. The small drawing annexed shows a portion of the dorsal (anterior)
surface of the ischium of the yellow appendage, enlarged.

. A similar view of the second maxilliped of the right side.

. A similar view of the first maxilliped of the right side.

. A similar view of the maxilla (second maxilla) of the right side.

. A similar view of the maxillule (first maxilla) of the right side.

. A similar view of the mandible of the right side.

. A view of the mouth and of the structures around it, after removal of all the paired

appendages save the maxillule and mandible of the right side.

. A diagram of a transverse section through the branchiostegite and the structures

which underlie it at the level of the articulation of the first walking-leg.

. A diagram of a longitudinal section through the branchial chamber of the left side.

The podobranch of the second maxilliped is shown in perspective, and the
position of the arthrobranchs of the third maxillipeds is indicated in dotted lines.

_ A view of the Crab from the left side after removal of the branchiostegite. Part

of the inner roof of the gill-chamber remains, and is held back by a hook above
the third walking-leg. :

. A view from within of a portion of the removed branchiostegite of the same side.
, A view from below of the inhalent openings above the first and second walking-

legs of the same side.

A view from above of the gill-chamber of the same side. The portions of the inner
roof which have not been removed are held back by hooks.

Portions of hairs from the third maxilliped:—A, from the median band of the
ischium; B, from the palp of the same limb; C, from the flagellum of the
exopodite. : :

* Puate 11.

A view similar to that in Fig. 15, after the removal of the greater part of the gills.

A similar view after complete removal of all the gills, save those of the second
maxilliped, the third maxilliped being in the normal position.

A similar view with the third maxilliped divaricated from its fellow.

142 THE MOUTH-PARTS OF THE SHORE CRAB.

Explanation of the lettering of the figures.

a. antenna.
a. antennule.
ab. abdomen,
ac. accessory muscles.
ap. apophysis.
ar.ch, arthrobranchs of cheliped, or
stumps of same.
ar.mp. (2, 8). avthrobranchs ef maxillipeds.
art. articulation.
b, basis.
br. branchial ridge.
¢c. carpus,
ch. cheliped.
col. collecting space.
cCa@, COXA.
cv.s, coxopoditic sete,
d. dactylus.
e. eye.
eb.c, epibranchial channel.
eb.sp. epibranchial space.
en. endopodites.
en.mp. (1,2). endopodites of maxillipeds.
ep.(1, 2,8). epipodites.
epist. epistome.
et. (1,2). endites of first maxilla,
ev, exopodite.
ex.mp. (1,2). exopodites of maxillipeds.
evh.c. exhalent canal or passage.
evh.o, exhalent opening.
Jl. flagella.
Jy. flange.
h. hinge.
hy. hypobranchial space.
openings in front of gills,

hy. ar. ch. \ :
leading to hypobranchial

hy, plb. (1, 2).
space,

7. ischium,
2. fl. inner floor.

2.2. inner lacinia,
ir. inner roof.
in.o. approach to iifthalent opening.
1. (1, 2). cleft lobes of maxilla.
im. labrum.
m. merus. —
md. mandible.
mp. (1, 2,3). maxillipeds.
mth. mouth.
o.l. outer lacinia.
ofl. outer floor.
or. outer roof.
o.mp. opening leading to gill-chamber,
in front of third maxilliped.
o,M.Z. opening of Milne-Edwards.
p- propus.
p.hy. posterior opening of hypo-
branchial chamber.
po. (1, 2). podobranchs.
pe.l. pericardial lobe.
pgn. paragnathum.
plb. (1, 2). pleurobranchs.
plp. palp of mandible.
pr. (1,2,8). processes upon rim of branchio-
stegite which meet corre-
sponding processes above
articulations of legs.
pr. processes above articulation of
legs.
se. scaphognathite.
sel. sclerite.
st. sternum of third maxilliped.
st’. fused sterna of mouth-parts in
front of third maxillipeds.
t.i.ad. tendon of inner adductor,
t.o.ab. tendon of outer abductor.
t.o.ad. tendon of outer adductor.
w.l. (1, 2,3, 4). walking-legs.

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THE JOURNAL

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Vou. XXXV. ZOOLOGY. No. 233.
CONTENTS.
Page
I. The Wing-Venation of the Order Plectoptera or Mayflies. By
R. J. Trstyarp, M.A., Se.D. (Cantab.), D.Sc.(Sydney), F.L.8.,
C.M.Z.S., F.E.S8., Entomologist and Chief of the Biological
Department, Cawthron Institute, Nelson, N.Z. (With 10 Text-
CSOT OS) Ramee AR Mikes ee eee allan en ee ter Arie Me ar aoe na a Ct 143
II. The Structure of Certain Paleozoic Dipnoi. By D. M. 8.
Warson, F.R.S., and E. L. Grit, M.Sc. (Communicated by
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LINNEAN SOCIETY OF LONDON.

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LIST OF THE OFFICERS ANB COUNCIL.
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ON THE WING-VENATION OF THE PLECTOPTERA. 143

The Wing-Venation of the Order Plectoptera or Mayflies. By R. J.
TrntyarpD, M.A., Se.D.(Cantab.), D.Sc. (Sydney), F.L.S8., C.M.Z.8.,
F.E.S., Entomologist and Chief of the Biological Department,
Cawthron Institute, Nelson, N.Z.

(With 10 Text-figures. )
[Read 30th November, 1922.]

THE present paper is intended to be read in conjunction with the series of
papers being prepared by me on the Wing-Venation of the Order Odonata
or Dragonflies, the first of which has already been published (Tillyard, 1922),
the remainder having been kept back to allow of the present results being
considered first.

At the present time, considerable doubt appears to exist as to the true
interpretation of the homologies of the wing-veins in Mayflies. This is not
to be wondered at when we consider that Comstock and Needham originally
offered one interpretation (1899), which became widely accepted, but that,
following on Needham’s discovery (1903) of the crossing-over of the supposed
Rs in Anisopterous Dragonflies, Miss Anna Morgan (1912), working clearly
under the influence of this discovery, offered a new interpretation bringing
the venation of the Mayflies into line with that of the Dragonflies in this
very important respect, although the evidence brought forward in her paper
to support this view is admittedly of the slenderest kind. Space will not
allow us to give here a full critical discussion of Miss Morgan’s paper. It
will only be necessary to make a single quotation from it, as follows
(J. ¢. p. 98) :—

“ An actual connection between R and Rs trachea (in Mayflies) cannot be
shown by constant structures. Moreover, Mayflies and Dragonflies are
closely-allied groups, and their general tracheation is similar in many points.
Furthermore, this condition of the radial sector trachea is exactly the same
as that just described in the Damsel-flies, where there can be no doubt that
such a crossing has taken place. It is, therefore, highly probable that the
radial sector is present in Mayflies, and that both the sector trachea and the
vein Rs have been stranded on M,, and have left no positive trace of their
origin.”

On these very slender and debatable grounds, Miss Morgan proceeded to
alter the homologies of the wing-veins in the Order. While recognising
and admiring the painstaking work shown in her study of the tracheation
and venation of so many little-known types, we have to confess that, like
Prof. MacGillivray, under whom the work was done (J.c. p. 89), we
“disagree with some of the interpretations . . . . presented.”

Now that I have shown the strong improbability that the supposed Rs
of Needham in the Anisoptera is really not that vein (Tillyard, 1922), and
definite proof is forthcoming that, in the Zygoptera, the supposed Rs never

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 10

144 DR. R. J. TILLYARD ON THE

had any connection with R at all, and is most certainty not that vein, any
system of homologies based on these results of Needham in the Odonata
becomes so highly speculative in character as to cease to attract even those
who may have originally accepted it. It is, therefore, necessary to enquire
anew into the whole question of the homologies of the wing-veins of the Order.
This I have attempted to do during the last three years. The results obtained
appear to me to show, not only that the system proposed by Miss Morgan
was incorrect, but that the original system proposed by Comstock and
Needham (1899) was also wrong, and that the true interpretation of the
homologies is one hitherto unsuspected in any quarter. These results, again,
also support some startling new results obtained for the Odonata ; and it is
for that reason that the publication of the present paper is interpolated
between the first and second papers of the series on that Order.

For the purpose of obtaining a really trustworthy result in an admittedly
difficult problem, I have attempted to combine three methods of study, viz. :
(1) the study of the Paleozoic Mayflies; (2) the study of the nymphal
tracheation in archaic existing types ; and (3) the application of the principles
of convex and concave veins, both to fossil and recent forms. Good fortune
favoured this plan. In the first place, when visiting New Zealand in
1919-20, Prof. ©. Chilton, of Canterbury College, Christchurch, very kindly
accompanied me on a visit to the Cass Biological Station, which is under his
charge. Here we found abundant material of the larvee of the archaic types
of Siphluridee for which New Zealand is famous. As all the necessary
apparatus for dissections and microscopic study were at hand in the Station,
I was able to make a thorough study of these larvee on the spot. This
produced a surprise, in that the tracheation was found to be almost complete
in the various instars, and fairly easy of interpretation ; whereas, in the
Mayflies studied by Miss Morgan, the tracheation was both incomplete and
irregular. Following on this discovery, during my visit to America in 1920,
through the kindness of Prof. C. Schuchert, I studied the Lower Permian
fossils left behind by Dr. Sellards in Yale University, and was surprised to
find among them a very fine specimen of Protereisma, so beautifully preserved
that it at once gave me the key to the whole venation of the Order, and,
incidentally, supported the evidence given by the study of the larval
tracheation. It then became clear to me that, if the significance of the
alternation of convex and concave veins had been fully understood, and
applied to the solution of this problem, the homologies would at once have
become perfectly clear. In his paper on the Lower Permian Mayflies,
Sellards (1907) entirely ignored this important point. Consequently, no
help was to be obtained from his work by Miss Morgan, and she merely
contents herself with copying his figures (1912, pl. 9. figs. 62-66), and
attaching thereto the results obtained by her own researches.

Text-fig. la shows the hind-wing of a species of Protereisma with the
homologies of the veins obtained by this new method of study. For

WING-VENATION OF THE PLECTOPTERA. 145

purposes of comparison the fore-wing of Ameletus ornatus is placed on p. 146
(text-fig. 1b). The corresponding notations of Comstock-Needham and
Morgan are given in the accompanying table. In the second column the
alternate convexity and concavity of these veins is indicated by the signs
+ and —.

Text-ria. | a.

apis DSS S0SSCRSSs5
= (joeoegeS nas S Ss =
Ssaaese POEs Tie ts

eae fae Goeeess ea

Seen
CIRRQ
RR
AERA
Hind-wing of Protereisma sp,, Lower Permian of Kansas; the fore-wing is closely similar, but a little

narrower at base, as shown in text-fig. 3.

TasiE OF WING-VENATION OF MAYFLIES.

New Notation. Sign. Cometerts Needham Morgan’s Notation.
Cc + C C
Se — Se Se
nt Te R, R,

Roa = Ry M,
2Roa + (omitted) (omitted)
8Roa = (omitted) (omitted)
4Roa + (omitted ) (omitted)

5 5Roa = 1p Rs
Rep + 183 Interpolated vein 1.
Rs; = R; M,
Raa + M, M;
Rup — M:, (omitted)
R; + My M,
M, _ Cu, Cu,
M, = Interpolated vein. Interpolated vein.
Ms+4 — Cu Cuz
Cu, + 1A 1A
Cuz — 2A QIN
1A + 3A 3A
2A + 3A 3A
3A ar 3A 3A

146 DR. R. J. TILLYARD ON THE

Tt will be noticed that no less than five branches of Re, are indicated in
the new notation. An explanation of this will be found in the argument on
page 148 concerning the nature of triads.

The results obtained by the three methods of study, already mentioned,
support one another, except that in the case of the study of the larval
tracheation a certain amount of variation is normally present. I have there-
fore presented the argument based on the Convexity and Coneavity of the
veins first, followed by that deduced from the study of these conditions as
noted in the fossil Proterezsma, and have kept for the last place the results
obtained from the larval tracheze. In dealing with these last, the alternative
interpretations given in the table on p. 145 will be considered, and reasons
given for rejecting them.

Trxt-Fig. 16.

Fore-wine of Ameletus ornatus (Eaton), Recent, for comparison with text-fig. 1@. tn, tornus,
Lettering as on p. 162.

Convex AND Concave Wrnas.

In all generalised insects, we are able to distinguish the presence of two
kinds of veins on the wings, viz. those which occupy the summits of ridges,
commonly called convex veins (indicated by a plus sign), and those which lie
in the bottoms of grooves or hollows, commonly called concave veins (indicated
by a miuus sign). In the ideal archetypic wing, convex and concave veins
follow one another alternately across the main portion of the wing,
Se being —, R, and its sector +, M —, the vein commonly called Cu, 254
and Cu, —. This last is always to be distinguished by lying either in or
very closely anterior to the deep anal furrow, which separates off the clavus
or anal area from the rest of the wing. The clavus itself is a wholly convex
area, and carries only convex veins, viz. the three anal veins 1A, 2A, and 3A.
Tn an ideal forked vein, both branches of the fork keep the same condition
of convexity or concavity as the main stem; but the allernation of ridge and

WING-VENATION OF THE PLECTOPTERA. 147

hollow is preserved, in the case of areas broad enough, by the development of
intermediate branches which take the opposite condition to the two branches
which they separate. These have been considered as interpolated veins in
the Mayflies ; ¢.e., they have been supposed to develop from the margin
of the wings inwards, and not to be true branches of the main veins at
all. A study of the fossil Mayflies shows, however, that originally there
‘were no interpolated veins in this Order at all, and that the discontinuity
of these veins has been brought about actually by absorption of the basal
connections with the veins of which they are true branches.

The primitive system of branching which is shown almost to perfection in
the Permian Mayflies is the type which we may conveniently term the triad.
This can be understood from text-fig. 2. V is a convex vein which divides

TExT-FIG. 2,

Va, *

Diagram of the triadic system of branching of main veins. For explanation, see text.

at its primary fork vf into two equally convex branches V; and V,. In the
diverging angle between these two, a concave vein arises from either
V, or V2, its point of origin being not far distad from vf; this secondary
fork is termed vf’. In the illustration before us, the concave vein is made to
arise from V,, so that the two branches of this vein become Vj, (+),
Vip(—), and V,(+). Hf, instead of being a convex vein, V were a concave
vein, then the two branches VY, and V, would be concave, and the inter-
mediate vein Vy, would be convex. The two cases can be distinguished by
calling the triad developed from an originally convex vein a positive triad,
that from an originally concave vein a negative triad.

Further branching may take place from any one of the three veins of a
triad, and usually consists of the development of further triads. In text-fig. 2,
a further triad is seen developed from Vo, viz. the positive triad Vo.(+),
Vou (—), and Vo,(+). If a triad had been developed from Vy, it would
have been a negative triad, and the notation used would have been 1V;,(—),
2Vi,(+), and 3V),(—); this notation was chosen to avoid duplication of
suffixes.

We can now see what has happened to the radius in Protereisma. This
vein is convex, and divides first of all into a positive triad, R, (+), Ro,3(—),
and R,y,5(+). The negative member of this group, R23, next develops a

148 DR. R. J. TILLYARD ON THE

negative triad, viz. Rea(—), Ra,(+), and R;(—). The second and third
members of this triad remain unbranched to the margin; but the first,
R.,, develops another negative triad, from the third branch of which a
further negative triad is also developed. To save complications in the
notation, I have named the five veins developed on the wing-margin by
these two latter triads 1Ro,, 2Ro, 3Ro, 4Ro, and 5R», ; it will be noticed
that they are alternately concave and convex.

The triad system, as will be clearly seen, always results in the formation
of alternately convex and concave veins along the wing-margin. There is a
large body of evidence to show that, apart from the Mayflies, this system
was the original system of branching of the veins in the insect wing.
Consideration of this evidence is beyond the scope of this paper, but attention
is drawn to it in the hope that students of other Orders will attempt to
recognise the remains of triads in the wings before them, As examples
of archaic triads, we ‘may take the universally recognised primitive set of
three branches of R, viz., Ry, Ro,3, Ry,;, which is formed from an original
positive triad with the intermediate vein developed from the lower branch Rs.
A similar archaic triad appears also to have developed from M, viz. M,(—),
M,(+), and M3,,(—), as shown in Protereisma, though this triad often
appears in other insects as M,,.(—), M;(+), and M,y(—), owing to a
difference in the position of the origin of the intermediate vein. It will
readily be seen how easily.a pectinate series can be developed from the
triadie system ; an example of this can be seen in the vein Cu, in Protereisma.
Simple dichotomies, on the other hand, may be explained by either the non-
development or suppression of the middle member of a triad, as in the case
of the secondary branches of Rs in most insects.

We now pass from the consideration of the triad to the application of the
rule of alternate convexity and concavity of veins as it affects the problem of
the elucidation of the homologies of the wing-veins in an archaic Mayfly such
as Protereisma (text-figs. 1, 3). The general rules applicable to all except
very highly-specialised wing-types may be stated as follows :—

(1) Two strongly convex veins can always be recognised lying between
the costal margin and the anal furrow, viz., Ry and Cu.

(2) The concave vein lying between R, and the costal margin is Se.

(3) M is a weakly concave vein lying between two ridges of R, and Cu.
In many archaic types it gives off a posterior branch close to the base, which
joins with Cu,, thus forming the cubito-median Y-vein, the main stem of
which is properly denoted by M;+Cu,, though usually called simply Ca,.
If this Y-vein can be recognised, any doubts as to the limits of M and Rs
should be finally removed.

(4) The vein lying either in or slightly anterior to the anal furrow is the
vena dividens, Cus.

WING-VENATION OF THE PLECTOPTERA. 149

(5) The clavus or anal area is a wholly convex area, and may carry from
one to three convex anal veins, any of which may be branched or simple.

Applying these rules to a well-preserved specimen of Protereisma, such as
the one figured in text-fig. 3, which Prof. Schuchert kindly allowed me
to keep for study during the last two years, the archaic condition of the
wing-venation is clearly shown in that the application of all five of the above
rules can be made without any doubt or hesitation. The high ridge of R,
ean be found at once with the concave vein Se lying anterior to it. At the
base of the wing there is developed a short convex vein between Se and the
costal border. This must be considered as a true primitive costal vein, C,
not a basal branch of Se, since it is convex and not concave. Sellards (1907)
has already shown how it forms a brace for the costal border, and has its
bomologue in existing Mayflies.

- T pxt-ria. 38.

Basal portion of fore-wing of Protereisma sp., Lower Permian of Kansas, considerably
enlarged to show details of venation. Note the cubito-median Y-vein haying M,
as upper arm, Cu, as lower arm, and M;+Cu as main stem. For lettering,
see p. 162.

We should next set about determining Cu,, the anal furrow and the clavus,
keeping the consideration of M and Rs until the last, as it is the limits of
these veins which have been the chief trouble in the past. Again returning
to our specimen of Protereisma, we find that the only vein which satisfies the
conditions for Cu, is that which has previously been called 1A, and that its
coneave branch Cu, can be clearly seen running in the anal furrow with a
characteristic curve basally, and separating off from the rest of the wing
a small, but quite distinct, convex clavus on which only convex veins are

150 DR. R. J. TILLYARD ON THE

present. In order to test this conclusion, we next look for the cubito-median
Y-vein, and find that it is present quite close to the base of the wing in the
very characteristic form shown in text-fig. 8. This definitely determines the
correct homology of the vein M;+Cuy, and enables us to pick up at once the
true median above it, viz. the concave vein which had previously been called
Cu, in spite of the fact that it is not convex. We thus find that M is a
three-branched vein, excluding M,, of triadic type, just as it is in so many
of the Paleeodictyoptera themselves.

We now have only the limits of Rs to consider. The whole of the many-
branched vein which had previously been considered to be formed of Rs and
M, combined by basal fusion, is now seen to belong to Rs alone. It arises
from R, near the base, and its short basa] piece is convex, It then branches
into a concave upper branch (R,,3) and a convex lower one (R,,;); and thus,
as already indicated above, it makes with R, a true positive triad. ‘The
numerous branchings of Rs shown in Protereisma need not be a cause of
wonder, since they are paralleled in quite a number of Palseodictyopterous
types.

In text-figs. 1 and 3, it should be noted that, excluding the clavus, on
which all the veins are convex, the whole of the veins reaching the wing-
margin are pliced alternately convex and concave. The complete notation
for the wing of Protereisma has been already given in the table on p. 145.

CoMPARISON OF THE VENATIONS OF FossIL AND
Recent MAYFLIES.

The next step isto make a careful comparison between the venation of fossil
Mayflies and the more archaic representatives of existing types. Tor this
purpose, I have chosen the wings of Ameletus ornatus, family Siphluridee ;
not because I believe it to be the most archaic of existing types, but because
it is archaic enough for our purpose, and happens to be the species on which
most of the studies of larval wing-tracheation were carried out.

Comparing the wings of Protereisma (text-figs. la, $) with the fore-wing
of Ameletus (text-figs. 16, 4), the first thing we notice is the change ‘in the
general shape of the wing. That of Protercisma is of the archaic elongate
oval form seen in most of the Palsodictyoptera and in the fore-wings of
Perlaria, in which no definite tornus and termen are developed, but only a
single continuously curved posterior margin from base to apex. In most
recent Mayflies, Ameletus included, a definite tornus (tr) is to be recognised ;
the angle of the tornus divides the original posterior margin into a shorter
basal part, to which the term posterior margin is still applied (called the
dorsum in Lepidoptera, but the term is inapplicable to Mayflies, which do
not fold their wings over their body in a roof-like manner), and a much
longer distal part, called the termen or distal margin. The result of this is
that the wing is now no longer of the primitive elongate oval shape, but

WING-VENATION OF THE PLECTOPTERA. 151

definitely triangular, the three angles being the base, the apex, and the tornus.
In the evolution of this shape of wing there is a definite reduction of the
areas served by the cubitus and anal veins, and a definite increase in the
area served by the media and radial sector. Consequently, if Comstock and
Needham’s theory of interpolated veins is justified, we should certainly
expect to find a number of them developed along the termen in Ameletus.
At first sight, this appears to have happened, since we can find at once no
less than five long veins, viz. 2Roa, 4Ro,, Re, Ry, and M., which are not
connected basally with their adjacent main veins. On the evidence of
recent Mayflies alone, Needham’s conclusions were perhaps warranted. But
when we come to compare the venation of <Ameletus with the fossil
Protereisma, we find that every one of these veins corresponds exactly with
a true branch of a main vein present in the fossil; and, further, the total
number of veins present in Ameletus, and their alternate convexity and con-
cavity, agrees exactly with the conditions in Protereisma. I have therefore
no hesitation in assigning to the veins in the fore-wing of Ameletus the same
names as I have already given to their homologues in Protereisma.

In the section dealing with larval wing-tracheation, it will be shown that
disconnection of the base of any branch vein is correlated with loss of its
precedent trachea in the larval wing, and may well be a direct result of this
larval condition.

By comparing the wings of Ameletus and Protereisma, we are now able to
see that the lengthening of the termen has taken placé chiefly in the region
served by Ryy; and M, the branches of these veins standing considerably
further apart in Ameletus than in the fossil, whereas the area seryed by
Ry, 3 has altered very little.

Turning our attention to Cu,, we note that this remains a strongly convex
vein, but its original: triadic system of branching has been changed to a
simple pectinate series, though the number of branches, seven in all, remains
the same, the branches still being alternately convex and concave. Though
superficially striking, this change is really a very slight one, consisting only
of a movement basad, on to the main stem, of the triad called Cu,, in
text-figs. la, 3.

The greatest changes that have taken place in the evolution of the existing
type of Mayfly wing are those in the basal region of the wing. For these,
we must refer to the enlarged basal area of Ameletus shown in text-fig. 4,
and this must be compared with the basal portion of the wing of Protereisma
in text-fig. 3. Here we note the following changes :—

(1) The separate costal vein C of the fossil has become fused with the
costal margin, forming a thickened basal margin; only its posterior distal
branch, hm, remains free, and forms the strong oblique brace, hm, in Ameletus
and all recent Mayflies. The intermediate stage in this evolutionary
process is well seen in the Jurassic Mayflies, the condition in Mesephemera
cellulosa (Hagen) being shown in text-fig. 4, upper figure.

iy DR. R. J. TILLYARD ON THE

(2) M has become fused basally for a short distance with Rs, the basal
stem of the two veins becoming obsolescent, but actually showing a weak
connection with M only, while the lost origin of Rs is shown by a very
definite stump projecting from R, close to the base. In some recent
May flies, however, Rs remains separate from M, and ‘is attached tangentially
to Ry just beneath hm. Both conditions may be regarded as specialisations
involving the abortion of the original basal piece of Rs.

(3) The basal portions of M and Cu, before the first forkings of these
veins, run close together as in Protereisma, but both of them turn upwards
almost transversely across the wing. At the upper end of this bent portion
is the formation which I have called the cubito-median Y-vein, so greatly

Text -FiG. 4.

Basal portion of fore-wing of Amedetus ornatus (Katon), Recent, considerably enlarged
to show details of venation. /, stump of Rs. Above, costal vein and humeral
brace (Am) in Mesephemera cellulosa (Hagen), Upper Jurassic of Solenhofen.
For lettering, see p. 162.

.

reduced from its original conditions (as seen in Proteresima) that it can
only just be made out in text-fig. 4. At this point, Cu forks, and its two
branches turn at right angles to its former course and run outwards about
parallel to the posterior border of the wing. Cu, remains, as in the
fossil, a strongly-curved vein concave to the posterior border; but the
shortening of its length has brought with it a definite increase of curvature,
so that the weak sigmoid curve which this vein shows in Protereisma
becomes replaced by a single strong arch, concave to the posterior border.
Other recent Mayflies, such as Hphemera, have the original sigmoid curve
intensified, but the vein itself greatly reduced in length, and the clavus
almost completely obliterated.

WING-VENATION OF THE PLECTOPTERA. 153

(4) The anal area, or clavus, in greatly reduced size, being about one-fifth
of the total wing-length in Ameletus, as compared with about one-third in
Protereisma. The number of branches of the anal veins are correspondingly
reduced, those of 1A to two definite branches, while 2A ne 3 A are only
indicated by short, weakly-formed veins, as shown in text-fig.

(5) The articulation of the wing, situated at the origin - Se and R,
becomes greatly strengthened by the formation of a hme ovoid callus
(text-fig. 4). It appears to be the growth of this callus which has forced
the basal portions of M and Cu below it into their peculiar curved positions.
In some recent Mayflies of higher type than Ameletws, these curved basal
parts of M and Cu become sinonyadnened and partially fused together ;
uniting with the bases of the anal vein also, they form an upwardly-
curving arch conyex to the posterior border, from which all the branches
of Cu and A take their rise.

We thus see that, in the archaic family Siphluride, as represented by
Ameletus, the correspondence with the Lower Permian fossil Mayflies is
exceedingly close, and the homologies of the veins perfectly clear. This, of
course, refers to the fore-wing wally The hind-wing, which has undergone
great reduction since Jurassic times, will be dealt crt after the uctention
of the larval fore-wing has been considered.

TRACHEATION OF THE LARVAL WINGS.

The study of the tracheation of the larval wings in New Zealand genera of
the family Siphluridz was carried out chiefly, as already stated, in the Cass
Biological Station. Some material was also obtained around Wellington
during my stay with Mr. Harold Hamilton, of the Dominion Museum, and
the éiccontions of these were caried out at his home in Karori. I wish here
to thank Prof. Chilton !and Mr. Hamilton for the facilities they afforded me
for carrying out this work.

The material studied consisted of numerous larvee of the following
species :—Oniscigaster distans Haton, Coloburiscus humeralis (Walk.),
Ameletus ornatus (Eaton), and Ameletus perscitus Eaton. The larvee of the
first and last of these were very abundant at Cass, and provided the bulk
of the dissections. The number of instars is not known for any of these
larvee, but the wing-sheaths can be dissected out for at least five instars,
showing comparative stages of growth from a tiny triangular bud up to a
consideribly elongated flap. For comparative purposes, I have designated the
youngest of these last five instars as the nth instar, viz. that in which the
wing-buds are very small flaps, capable of being dissected off with very fine
scissors if great care is exercised. The following instars will be termed the
(n+1)th, (n+2)th, (n+8)th, (n+4)th respectively, the (n+4)th being
the last larval instar. In order to avoid too many figures, I shall give

154 DR. R. J. TILLYARD ON THE

here only a series of drawings which I made from the wing-sheaths of
Ameletus ornatus, a larva which has fairly transparent wings, is easy to
dissect, and in which the trachese can be followed without any trouble.
Characters in which the other larvee differed from this will be noted
below.

The first point to be noticed is that the larval wing, in this Order,
is soldered to the thorax over a considerable portion of its basal area.

The free distal portion of the wing is that which lies to the right of the
dotted line in text-figs. 5-9. Great care is necessary, in dissection, to
remove sufficient of the soldered basal part to ensure that the bases of the
wing-tracheze, and their connections, are not severed or damaged in the
operation.

Just distad of the middle of the line marking the junction of the flap
of the wing with the thorax, there is always present a somewhat triangular
patch of hard chitin, shown shaded in all the figures. It at first separates
the subcosto-radial group of trachez, including M, from the cubito-anal
group, as shown in text-fig. 5 for the nth instar. But, in the next instar, it

Text-riag, 5,

Ameletus ornatus (Maton), Tracheation of fore-wing in 7th larval instar.
For lettering, see p. 162. (x97.)

moves forward a little, so as to overlie part of the base of M ; and finally, in
the last two instars, it comes to overlie the wholly or partially fused bases of
R and M. From its final position in the last instar, it seems to me that
this hard piece of chitin represents the large callus of the imaginal wing ;
its position in relation to the thorax is probably constant, its apparent
movement forward being probably due to changes in the actual positions of
the wing-trachee. The chief point of interest appears to me to be that it
certainly does cause trachea M to run closer up to R than it naturally
would if this callus were absent. Consequently, it may well be one of the
factors which helped to bring about the fusion of M with Rs basally, as we -
find it in the majority of larval wings in the last four instars (text-fios. 6-8).

WING-VENATION OF THE PLECTOPTERA. 155

Text-fig. 5 shows a typical fore-wing of Ameletus ornatus in the nth
larval instar. There is no costal trachea visible. The alar trunk connecting
the subcosto-radial with the cubito-anal group of trache is complete,
though of small calibre, and the two groups are widely separated. Instead
of coming off separately from the alar trunk, the tracheze Sc, R, and M arise
as separate branches from a single trachea of large calibre. Rs branches, as
a simple triad, into Ry, R;, and R,,5. Trachea M arches gently in front
of the callus already mentioned, and thus comes very close up to Rs just
where it comes off from R,. Most of the larvee examined in this instar
show exactly the condition given in text-fig. 5; but [ found one larva in
which trachea M had already captured trachea Rs at its base. This
condition becomes the usual one in the following instars. M is only two-
branched, as in the Order Perlaria. The trachez of the cubito-anal group
are very fine in calibre; but { could distinctly make out Cu, and 1A in all
the larvee examined, and could see Cu, in most of them. In following
instars, no trachea is usually visible in Cu,, though I found it quite well
developed in a fine larval wing of the penultimate or (n+3)th instar shown
in text-fig. 8.

TRXT-FIG, 6,

Ameletus ornatus (Haton).- Tracheation of fore-wing in (n+)th larval instar.
For lettering, see p. 162. ( x87.)

Passing on to the next, or (n+ 7)th instar (text-fig. 6), we note that the
callus has now come to overlie the base of M, and in the great majority of
wings examined, trachea M has captured trachea Rs at its point of nearest
approach basally. I have notes of a few exceptions, chiefly in the genus
Oniscigaster, in which, both in this and the following instars, any of the
following conditions may happen to occur :—

(1) Trachez Se, R, Rs, and M all come off in a bunch as separate trachesa -
from a single point on the alar trunk.

(2) RB arises from Se, but Rs and M arise close together from a point on
the alar trunk a little below them.

156 DR. R. J. TILLYARD ON THE

(3) Rs and M are slightly separated basally, but come together so as to be
almost indistinguishable while passing the callus, and then separate distally.

(4) Se arises by itself from the alar trunk ; considerably further down, a
group of three separate trachez, R,, Rs, and M, arise close together.

All the above conditions are occasional variants only from the dominant
condition, which is that shown in text-fig. 6. It should be noted that
trachea M, in this instar, usually arises quite separately from trachea
Se +R, and its course basally from alar trunk to callus diverges widely from
that of the latter. The number of trachez developed from Rs is increased
to four by the forking of R,,;, and there is an indication of the addition of
an anterior branch to R, in the form of a pale band, which never, at any
time, as far as I could see, develops more than the merest basal rudiment of
a trachea, and usually possesses none at all. The number of branches of M

TEXT-FI4e, 7.

is

Ameletus ornatus (Katon). Tracheation of fore-wing in (x +2)th or antepenultimate
larval instar. For lettering, see p. 162. (x41.)

is also increased to three by formation of the triad M,, M,, M3,,4; but the
tracheal supply of the middle branch My is quite irregular, and varies
greatly in the different larvee examined. A separate costal trachea appears as
a short basal anterior branch from Se. The alar trunk continues complete,
and of fair calibre.

Passing next to the (n+2)th or antepenultimate instar (text-fig. 7), we
find the conditions not very different from those indicated for the previous
instar, except for the presence of additional branches of R,, which now
number five in all, and the establishment of the triad M,, M,, M;,4 on a firm
basis, with a single trachea supplying M, as shown. In text-fig. 7, I show
a curious condition of trachea Cu, near the base. The course of the vein Cu,
is indicated by the dotted line, but the trachea arches up out of its normal
course, and passes close under M. I have scen this condition several times,

WING-VENATION OF THE PLECTOPTERA, 157

not only with Cu,, but in other parts of the larval wing. It seems to
indicate a certain amount of instability in the tracheation, and may possibly
be the stage precedent to a more complete breaking down of the tracheal
supply, as 5, exdavibitdl § in many of the forms studied by Miss Morgan.
Text-fig. 8 shows the tracheation of the fore-wing in ihe. (n+3)th or
penultimate instar. In this stage, R, Rs, and M generally appear to arise
as a single thickened trachea from the alar trunk, at a point somewhat
removed from the origin of Se, and beneath the callus. By careful
manipulation, this apparently single trachea can be separated into its
component parts, when it will be seen that it is in reality three separate
trachese arising from almost the same point. In some specimens, however,
R, may still be found arising asa branch of Se, while Rs and M arise as a
single trachea well below it. I think the varying conditions seen in the later

TrxtT-FiG. 8,

Ameletus ornatus (Katon). Tracheation of fore-wing in (n+3)th or penultimate
larval instar, For lettering, see p. 162. (x48.)

instars are almost wholly due to the presence of the callus, one or more of the
original trachez taking a devious course to avoid passing under it, or possibly
becoming displaced at ecdysis by the pressure of the harder mass of chitin.
Jn this instar, the number of branches of Rg, is still five, and the most
anterior of these never has any tracheal supply except a mere rudiment.
Various irregularities may be noted. In the specimen figured, the trachea
which usually supplies the most posterior branch of R, arises alongside the
trachea belonging to the branch above, and runs with it to about half its
length, when it turns downwards and supplies the course of the most
posterior branch for its distal half only. The trachea supplying M,
sometimes arises from M;,,. Cu, possesses a complete trachea in this
specimen ; but in the majority of wings examined, it either has no trachea or
a mere rudiment. These are all indiy Fava variations in the tracheation, and

158 DR. R. J. TILLYARD ON THE
are of no significance except to indicate the essential variability of the
tracheation in many of its details.

In this instar, the line of fusion of the wing-sheath with the thorax ceases
to be straight, and shows a definite convex bulging between the subcosto-
radial and cubito-anal groups of tracheze. The alar trunk is still complete,
but thrown into a strong loop. The large anterior callus already noted in
previous instars now appears in close contact wiih the axillary area of the
thorax, this position corresponding with that in the imaginal wing, where the
eallus articulates with the axillary process of the thorax. A new structure
is apparent in the cubito-anal region, viz. a smaller, somewhat reniform,
posterior thickened patch, shown shaded in text-fig. 8. This corresponds
with the opaque and somewhat thickened anal area at the base of the imaginal
wing ; but it never becomes a true callus, 7.e. it does not articulate with an
axillary process from the thorax.

The last or (n+4)th instar differs from the previous one in the much larger’
size of the wing-sheaths, the strong development of the anterior callus, the
very clear indication of the whole of the imaginal venation, including all
the cross-veins and veinlets, and the presence of all six branches of R,.
In this instar, the tracheation is not easy to follow, owing to the strong
pigment bands of the imaginal venation overlying it. As the general
tracheation scheme is closely similar to that of the previous instar, and the
same individual variations are equally apparent, it has not appeared necessary
to me to figure it. One point of interest needs to be mentioned, viz. that I
only succeeded in finding the complete alar trunk in one specimen of
Oniscigaster distans. In all the others examined, the basal portion cu-a was
missing, and the single trachea which gives origin to Cu and the anal veins
appeared to arise from the same trunk as the other tracheze above it. his is
exactly the condition shown in Miss Morgan’s figures (1912, pls. 5-7).
Thus it will be seen that, if she had studied other instars as well as the last,
she would not have fallen into the error of failing to homologise correctly
the cubital tracheze and veins, which it is quite evident she has done.

The above discussion applies to the fore-wing only. As the hind-wing is
greatly reduced in size in all recent Mayflies, and differs very greatly from
that of the Permian Mayflies, as well as from the fore-wing of recent forms,
I have thought it best to summarise the evidence for its venational homologies
in a separate section.

Tar Hinp-wine.

The dissection of the small hind wing-sheath of a Mayfly nymph is not
easy, and I only succeeded in getting good preparations from the last two
instars. Text-fig. 9 shows the tracheation in the hind-wing of Ameletus
ornatus, penultimate instar. The alar trunk is complete, but there are no
signs of a callus. The line of fusion with the thorax lies well out beyond the
origins of the wing-trachese, as shown by the dotted line in the figure.

WING-VENATION OF THE PLECTOPTERA. 159

There is a separate costal trachea OC, lying far above Sc ; the latter trachea
lies far above the group of R and M, which arise close together well down
towards the cubito-anal group. Rs arises from R very close to the point

TEXT-FIG. 9.

Ameletus ornatus (Eaton). Tracheation of hind-wing in (x+3)th or penultimate
larval instar. For lettering, see p. 162. (X87.)

where M curves slightly upwards, so as to come very close toit. Rs divides
into two branches, R,,3 and Ry,;, each of which gives rise to a simple triad
with incomplete tracheation. The middle branch of the triad of R.,3 is
supplied by the trachea R;, but R, and the distal part of R3 have no trachee.

The two outer branches of the triad of R,,; have their tracheze present, but

Text-Fie. 10.

Ameletus ornatus (Eaton). Venation of hind-wing. For lettering, see p. 162. (x16.)

the middle branch has none. M gives rise toa simple triad in which the
trachea supplying M;,, switches up into the middle branch distally, leaving
the distal half of M, without a trachea. Cuy has no trachea.

LINN. JOURN.—ZOOLOGY, VOL. XXXV. ital

160 DR. R. J. LILLYARD ON THE

In interpreting the above scheme, we must compare the imaginal venation
shown in text-fig. 10 with the tracheation shown in text-fig. 9. There can
then be no doubt as to the correct homologies of the main veins. Two points
only are open to doubt. Firstly, what is the correct notation for the middle
branch of the triad of R.,;? This appears to be a branch of R;. But in the
penultimate instar of Oniscigaster distans, I find the trachea R, supplying
this branch. I therefore conclude that the tracheation*is variable, and give
preference to the notation Ry, for this vein. This is done because, in the
fore-wings of recent Mayflies, and also in both wings of the fossil Protereisma,
R; isan unbranched vein. Secondly, what is the correct notation for the
middle branch of the triad of M? In this case, both in the larva of
Oniscigaster distans and in the one figured, the trachea supplying it comes
from M;,4. In addition to this, the vein itself, in the imaginal wing, arises
from M;,4 as a definite branch. All the evidence, then, points to the correct
notation for this vein being M3, the other members of the triad being M,,,
and M,. In this respect, we must reckon that the hind-wings of Ameletus
and Oniscigaster differ from their fore-wings, and also from both wings of
Protereisma. The explanation probably lies ina shifting, at some fairly early
point in the evolution of the hind-wing, of the point of attachment of the
middle member of the triad, followed at a slightly later time by the
development of a corresponding new tracheal supply.

ALTERNATIVE INTERPRETATIONS.

Having now considered all the evidence available under the three headings
already indicated, we may ask whether any of them offer any basis for an
alternative interpretation. The answer to this must be, that, in the case of
the evidence from Convex and Concave Veins, and from the Fossil Record,
there can be no doubt whatever as to the true homologies of the veins, and
no alternatives to the solution here given can be offered. In the case of the
larval tracheation, we have noted a considerable amount of variation both in
the branch tracheal supply and in the mode of origin of the main trachez
from the alar trunk. To anyone, therefore, who pins his faith upon this kind
of evidence alone, and who refuses to admit that offered under the two other
headings, alternative solutions are possible. The two chief alternatives, of
course, are those given in the table on p. 145 as the Comstock-Needham and
Morgan: notations respectively. We may ask whether the evidence from
the tracheze alone does really support either of these two schemes rather than
the one we have given in this paper.

With regard to the original interpretation given by Comstock and Needham,
this clearly assumed that there had been a basal fusion between Rs, M, and:

ju. The evidence of the tracheation certainly supports the idea of a basal
fusion between Rs and M; but it shows clearly enough that Cu remains
quite distinct. It does not seem necessary to say more about this alternative

WING-VENATION OF THE PLECTOPTERA. 161

notation, for the simple reason that its authors themselves admitted that it
was unsatisfactory ; and, later on, they accepted Miss Morgan’s interpretation
instead (see Comstock, 1918, chap. x.). We have, then, only to consider
Miss Morgan’s notation. With respect to this, we may note the following
points :--—

(1) The research was carried out on wings of the last larval instar only.
In this instar the tracheation is the most variable of all, and the posterior »
continuation of the alar trunk is usually absent. In spite of this, Miss
Morgan correctly interpreted the homologies of the cubital trachez, but
failed, for some reason, to connect them with their true venational homologues
in the imago, Had she examined the tracheation in this region in earlier
instars, she could not have failed to note the close correspondence between
the strongly bent cubitus in the larva and imago; and her results, as far as
the limits of Cu were concerned, would then have agreed with those given in
this paper.

(2) The evidence for her chief innovation; viz. the crossing of a simple Rs
over two branches of M, as in Anisopterous Dragonflies, is not forthcoming
from the great majority of the larval wings examined, but only from the
wings of a single species, in which, as she admits (J. c. p. 98), “ half of
the wing-pads showed the radial branching just described and half of them
gave no sign of it.” The species which gave, in only one-half of its
representatives, an indication of Rs crossing over M, was a species of
Heptagenia, a genus which is by no means the most archaic of the fifteen
genera studied by Miss Morgan, Yet she accepts this evidence, against the
weight of evidence supplied by all the other fifteen genera and the other
‘species of Heptagenia included !

As I have already pointed out, new light on the venation of Dragonflies
has made it certain that Rs does not cross over M in the Zygoptera, and has
also rendered it extremely improbable that the trachea which does this in the
Anisoptera is Rs at all. The evidence for a crossing of Rs over M in
Mayflies to be found within the Order itself is, as we have seen, practically
negligible. Thus Miss Morgan’s whole case totters to the ground, and must
be replaced by a more lasting structure.

To sum up, we haye seen how the evidence from the Convexity and
‘Coneavity of Veins agrees absolutely with that offered by a study of the
Permian fossil Protereisma, and the combined evidence of these two lines of
study is supported fo a very great extent by that of the tracheation of larval
wings, the only doubtful point in this latter evidence being its undoubted
tendency to vary. Taken all together, the new system of. homologies appears
to me to be supported by so great a weight of evidence as to be at once
acceptable in place of the hypotheses previously offered in the Comstock-
Needham and Morgan notations respectively. At the same time it offers a

new and clear basis on which to build up a system of classification within the

162 ON THE WING-VENATION OF THE PLECTOPTERA.

Order, which will present, to some extent at any rate, the real lines of
evolution of the Mayflies through the long period of time from the Lower
Permian to the present day. That problem may well be left toa later paper.
IT also hope to continue the series already begun on the venation of Odonata,
and to show how, in that Order, the behaviour of the radial sector has
followed the lines set out in this paper for the Mayflies, though very high
‘specialisations of the Odonate venation have masked the original close
similarity between it and that of Mayflies.
Cawthron Institute, Nelson,
31 March, 1922.

Literature Quoted.

Comstock, J. H., 1918.—‘ The Wings of Insects.”” Ithaca, N.Y. (Chap. x.).

Comstock, J. H., and Nuepxaam, J. G., 1899.—‘‘The Wings of Insects :
chap. iv. contd.” Amer. Nat. xxxili. pp. 118-126.

Morean, Anna H., 1912.—‘* Homologies in the Wing-Veins of Mayflies.”
Ann. Ent. Soc. Amer. vol. ii. pp. 89-106, pls. 5-9.
Neepuaw, J. G., 1903.—“ A Genealogic Study of Dragonfly Wing- Venation.”
Proc. U.S. Nat. Mus. xxvi. pp. 703-761, 24 plates (No. 1331).
Sutiarps, W. H., 1907.—* Types of Permian Insects: Part II. Plectoptera.”’
Amer. Journ. Sci. xxiii. pp. 345-355.

TrttyarD, R. J., 1922.—“ New Researches upon the Problem of the Wing-
Venation of Odonata. I. A Study of the Tracheation in the Larval
Wings in the genus Uropetala from New Zealand.” Ent. News, xxxii'.

Venational Notation used in the Text-sigures.

LA, first analis; 1A,, 1 A,, its two branches. 2A, second analis. 3A,
third analis. C, costal vein or trachea, separate from the costal margin.
Cu, cubitus. Cuy, first cubitus; Cuj,, Cu, Cu,., its principal triad in
Protereisma. Cup, second cubitus or vena dividens. cu-a, cubito-anal trunk
trachea. cuf, primary cubital fork. hm, humeral brace, formed from the
descending posterior branch of the costal vein. &, stump of Rs. M, media ;
M,, M2, M;,4, the three branches of its triad. M;, its posterior branch,
forming the upper arm of the cubito-median Y-vein, the lower arm being
Cu; M;+Cu,, in Protereisma, the main stem of the cubito-median Y-vein,
commonly called Cu,. mf, the primary median fork. Pe, precostal margin.
R, radius ; Rj, its main stem or anterior branch. Rs, radial sector; Ry,3,.
R,,5, its two main branches ; further branchings are indicated, first by the
four veins R,, R3, Ry, R;, then by the suffixes a (anterior) or b (posterior)
attached to any of these four veins, and finally by numerical prefixes 1 to 5,
to avoid further suffixes. Sc, subcosta. sc-7, subcosto-radial trunk trachea.
tn, tornus.

THE STRUCTURE Of CERTAIN PALMHOZOIC DIPNOI. 163

The Structure of Certain Paleozoic Dipnoi. By D. M. 8. Warson, F.R.S
and EH. L. Girt, M.Se. (Communicated by i Idig Se ehsonmen
F.R.S., Sec. L.8.)

(With 34 Text-ficures.)

[Read 30th November, 1922. |

Tats work began by an attempt to write an account of the structure of the
Coal-Measure fish Sagenodus and Ctenodus on the basis of the splendid
material in Newcastle. This collection when worked over by us left certain
problems unsolved which have in part been cleared up by fossils in Hdinburgh
and elsewhere. During our search for this information we saw the splendid
specimens of Uronemus in Edinburgh, and the wish for further light on
certain morphological problems led us to extend the investigation to several
other paleeozoic Dipnoi.

The resulting paper now gives an account of the structure of these fish,
which certainly includes the chief variants known to have occurred in the
group, although some forms which might prove of great interest are still
known to us only by the original descriptions or by. unsatisfactory specimens.

With the possible exception of Jipterus, the most completely known of
the fossil Dipnoi is now Sagenodus, and we therefore deal with that fish first,
following on with an account of the related genus Ctenodus. For these two
genera, the chief source of our material has been the Atthey Collection in the
Hancock Museum at Neweastle-on-Tyne, but additional information of great
importance has been derived from fine specimens in the Royal Seoiish
Museum and British Museum (Natural History).

The material is from the following horizons and localities :—

Lower Carboniferous (Oil Shale group) : Broxburn, Loanhead, and
other localities near Hdinburgh—(Sagenodus and Ctenodus).

Upper Carboniferous (Westphalian): Newsham, Northumberland;
Longton, Staffs.; Ardwick, Lanes.; Newarthill, Lanarkshire ;
Linton, Ohio ; Kansas—(Sagenodus and Ctenodus).

Upper Carboniferous (Stephanian): Nyran, Bohemia—(Sageno-
dus).

Lower Permian (Artinskian): Wichita, Texas; Kounova, Bohemia—
(Sagenodus).

SAGENODUS, Owen.
Until the publication of Dr. Smith Woodward’s second volume of the
“ Catalogue of Fossil Fishes” in 1891, the generic distinctness of Sagenodus
from Ctenodus was not generally recognized. Nearly all the published work
of any importance on the structure of these fishes appeared before that date,

164 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

and consequently in that work Sagenodus was usually referred to under the
name of Ctenodus obliquus. The specific names imbricatus, ellipticus, elegans,
and others, many of them probably synonyms of obliquus and of Owen’s species
S. inequalis, were also applied by the earlier investigators to remains of
Sagenodus; and more recently a large number of names of very doubtful
value have been put forward, founded asa rule on isolated examples of the:
abundant and very variable tooth-plates.

Fie. 1.

tab.h. oer

Sagenodus. Dorsal aspect of cranial roof, x 2. Mainly from a specimen from the Low
Main Seam of Newsham, in the Royal Scottish Museum. Jr., “frontal” (=frontal
+ post-frontal ?); 7.fr., interfrontal ; ¢.tem., intertemporal ; na., nasal ; oce.f., occipital
flange ; p.na., prenasal ossicles; par., “parietal” (=parietal + dermo-supraoccipital);
sq., ““squamosal”; zab., “tabular” (=tabular+supratemporal) ; tab.h., tabular horn.

What has hitherto been known of the structure of Sagenodus was due
mainly to the investigations of Thos. Atthey and Albany Hancock (1868-
1875). Important additions were made by L. C. Miall (1881). Fritsch, in

STRUCTURE OF CERTAIN PALMOZOIC DIPNOI. 165

his ‘Fauna der Gaskohle’ (vol. ii. 1889), published numerous excellent
figures of detached bones of Sagenodus, but a large proportion of them were
unidentified, and his letterpress added little to what was already known.
More recent contributions will be mentioned incidentally and in the hist of
Literature.

The available material has been re-examined by us, and the results are
here stated, largely in the form of figures.

Fie. 2.

Sagenodus. Cranial roof of six individuals, to illustrate the extent of variation. A-D,
Atthey Coll.,* from the Low Main Seam, Newsham; E, ¥, Royal Seottish Museum,
from the Virtuewell Seam, Newarthill.

In its general structure the skull of Sagyenodus resembles that of Ceratodus
in having an extensive cartilaginous neural cranium, of which no traces are
ever to be seen, surrounded and supported by well-developed membrane
bones. The bones which covered the top of the head frequently remain
connected together in the fossils, but no complete articulated skulls are
known.

* Except where otherwise stated, the originals of all figures are in the Atthey Collection.

166 PROF. D. M. 8S. WATSON AND MR. E. L. GILL ON THE

The Roof of the Skull.

It has been customary to dismiss the cranial roof of Sagenodus and
Ctenodus as being composed of numerous small bones of rather indeterminate
arrangement and doubtful homology, though bearing a considerable resem-
blance to those of the roof of the skull in Dipterus. Incomplete skull-roofs
of both genera were described, but not figured, by Hancock and Atthey
(1872, p. 401), who clearly stated some of the points distinguishing skulls of
Ctenodus tuberculatus (i.e. the genus Ctenodus as now understood) from those
of C. obliquus (i.e. Sagenodus). Rough figures of the posterior roof-bones of
the Sagenodus skull were given by T. P. Barkas (1873, figs. 244-246) and
Miall (1881, fig. 1). Miall’s figure was good as far as it went, except in
the region of the bones that we call the intertemporals, where it is probable
that he was misled by false suggestions of sutures. In a large number of
examples, at any rate, we have met with no similar case, though monstrosities
do occasionally occur (cf. fig. 6, A and U, p. 170).*

Of Barkas’s three figures, 245 evidently represents the same specimen as
was used by Miall for the right side of his figure, while fig. 246 represents a
fragment seen from the underside and showing the shape regularly assumed
underneath by the suture at the front of the “parietal.” His fig. 244, that
of the hinder ~wo-thirds of a skull-roof, is correct on the whole, but the
posterior corners are missing; the original is now in the British Museum
(No. 45852).

The length of an average skull-roof is about 53 inches (14 em.), but some
were rather larger. The figure here given (fig. 1, p. 164), together with the
variations represented in fig. 2, renders unnecessary a detailed description of
the form of the bones. The bones are named in accordance with the sug-
gestions of Watson and Day (1916, p. 42). Their degree of variability may
be estimated by comparing the outline of any one bone through the series of
figures, each drawn from an actual specimen, given in fig. 2, p. 165. It will
be seen that though there is wide variation in details of shape, there is never
any difficulty (apart from rare monstrosities) in recognizing single bones
detached from the skull.

In the state in which they occur in the shales the bones of the skull-roof
are about 4 mm. thick in full-sized heads. Neighbouring bones are locked
together by means of thin lamine, radially ribbed, springing from almost the
lowest level of the thickness of the bones and fitting into the adjacent bone
at the same level. Mr. H. Fletcher, late of the Zoological Museum, Uni-
versity College, Reading, tells us that the bones in the skull-roof of Ceratodus
forstert are articulated in the same way. On the inner surface of the roof
some bones encroach upon others further than they do outside, so that though
the pattern on the two surfaces is nearly the same it is never identical.

* In fig. 6, A, the right frontal (f.7.) is quite abnormal in shape, and in fig. 6, C, the left
intertemporal has entirely vanished, the region in which it should be being occupied by an
enlargement of the bones which normally surround it.

STRUCTURE OF CERTAIN PALHOZOIC DIPNOI. 167

The inner face of the roof-bones has a polished surface and usually bears
radiating ridges. ‘The outer face varies a good deal in surface character, but
it is generally granulated, with a varying degree of gloss, and shows fine lines
radiating on each bone from a central pit or rosette. Bundles of these lines
often run from the rosette of one bone to that of the next. Rosettes are
always more strongly developed on the frontal and squamosal than on the
other bones. The whole appearance of the bones suggests that the head was

Fie. 3. *

Sagenodus. Nasal and prenasal bones. Q and R after Fritsch, all the rest from specimens
in the Atthey Collection from Newsham, x about 2.
A-M, prenasal ossicles: F, G, H, J, under surface, the rest dorsal aspect.
N-S, nasals: P, under surface of O; S, a remarkable form which would not fit into any
skull yet seen.

covered only by a thin skin, and not, as in Ceratodus, by scales. This
supposition is confirmed by the occasional appearance of the grooves of
sensory canals on the skull-roof. A well-marked lateral-line groove is
seen to run from the rosette of the “parietal,” first to the rosette of the

168 PROF. D. M. 8S. WATSON AND MR. I. L. GILL ON THE

intertemporal and then to that of the squamosal. This groove is for a line
of pit organs, found also in Ceratodus. Such grooves are independent of
the bundles of strize already referred to. The corresponding bones of very
young individuals show these grooves much more plainly (see fig 4, A, B, C,
below). The course of the grooves, when they appear at all, is precisely that
of the sensory grooves marked on the skull of Dipterus.

One common type of skull-roof is not represented in fig. 2, p. 165, because
it apparently always came to pieces in fossilization, and is hence only known
by detached bones. It is characterized by the high polish of the external (as
well as of the internal) surface of the bones, by their simplified outline, and
by the conspicuous zonal bands near their margins. A fish of this type was
evidently the predominant Sagenodus in Fritsch’s collection from the

Sagenodus. Parietals, x about 8. A and B, of very young individuals, show well-marked
lateral-line grooves and illustrate the scale-like form of the young bones; C is of a
rather older individual; D isa bone of the zonal and polished type; though small, it
has the adult form.

Gaskohle; in the Atthey Collection it is comparatively scarce and usually
of very small size, but a few large “parietals” prove that in the Northum-
brian coalfield, as in the Bohemian, it reached the same size as the other
forms. A very small “parietal” of this zoned and polished type is
represented in fig. 4, D, above.

The posterior margin of the skull-roof presents as a rule a concave out-
line, sometimes pronouncedly so (fig. 2, H & F, p. 165; fig. 18, A, p. 183).
In a few cases it is nearly straight or even slightly convex. A pair of
tabular horns for the attachment of the shoulder-girdle are conspicuous at
the back of a well-preserved skull. At the level of the inner face of these
horns there is a thin frill of bone (fig. 1, p. 164, oce.f.) stretching across the

STRUCTURE OF CERTAIN PALHOZOIC DIPNOI. 169

whole hinder border of the skull. It springs from the three hindermost
roof-bones, not at their actual edge, but from a little way forward on the
under surface, and along this line the extreme hind border of the bones is
frequently cracked and bent down in fossilization. It is possible that in life
this thin flange stood vertically, more or less at right angles to the skull-roof,
and overlapped the hinder surface of the neural cranium.

From the outer hind corner of the tabular forward to the outermost point
of the squamosal the edge of the skull-roof is rounded off and usually lobed
and corrugated, except for a smooth and very definite notch in the border of
the squamosal. It isalong this part of the roof that ihe operculum is hinged

Fra. 5.

Sagenodus. Cireumorbitals, x about $.
Outer faces on the left, inner on the right.

(see fig. 18, p. 183). The notch in the squamosal lodges the anterior knob of
the operculum, and the two bones forming the hinge-line behind the notch
bear on their under surface a groove, bounded internally by a ridge, which
engages a flange borne by the operculum along its upper border. The
posterior knob of the operculum, more or less strongly developed, fits against
the hinder end of the smaller corrugated bone close to the corner of the
tabular.

From the outer point of the squamosal forwards the edge of the skull-roof
everywhere shows articular faces, with articular laminee where the bones
are well preserved. It is here that the orbitals and other marginal bones
were attached. Asa rule, however, the marginal bones, and the nasals as
well, have been lost, even where the main part of the skull-roof has held
together. Probably this main part (represented in the outlines in fig. 2,
p- 165), was flat or nearly so, whereas the bones bordering it would be

170 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

articulated with it on a downward curve, and would be subjected to much
greater strains during the maceration undergone by the skull before its burial.
The nasals are shown in their natural position in several skulls in the Royal
Scottish Museum; one skull from Newsham, in the same museum (No.
1378.45.7), shows several prenasal ossicles in position as well. They are
introduced in fig. 1, p. 164. In the Atthey Collection there are a number of
detached nasals and prenasals. Both are much more variable in shape than
the other bones of the skull-roof behind them (see fig. 3, p. 167). All the
prenasal ossicles seem to be crossed on the underside by a ridge and
groove (fig. 3, F,G, H, J, p. 167), which may mark the position of a lateral-
line canal.
Fie. 6.

Sagenodus. Outlines of specimens, showing marginal ossicles and circumorbitals.
A. From the Virtuewell Seam, Newarthill, Lanarkshire. Royal Scot. Mus. (1897/110/81).
ptr. & ptl., right and left pterygoids with their teeth t.7. & ¢.1.; man., mandible.
B. From the same horizon and locality as fig. A. R.S.M. (1897/112/5).
C. Coal Measures, Linton, Ohio, U.S.A. British Museum (N.H.) (P. 7778).

Circumorbitals and Marginal Ossicles.

These bones frequently occur scattered among the other bones in dis-
articulated examples of Sagenodus on slabs of shale, and in the Atthey
Collection there are large numbers of them in this sort of association or as
isolated bones freed from the matrix. Many of them were certainly orbitals,
for they have one of their edges thinned out and rounded, and leading on the

STRUCTURE OF CERTAIN PALHOZOIC DIPNOI. ileedale

underside to a hollowed and polished surface. Four of the commonest
patterns (of each of which there are several duplicates) are represented in
fig. 5, p. 169. It will be noticed that the radius of curvature of the orbital
notches is far from uniform, apart from questions of size of bones, which
seems to indicate that the orbit cannot have been circular. One skull in the
Atthey Collection (fig. 18, A, p. 183), several in the Royal Scottish Museum
(e.g. fig. 6, A, B, p. 170), and one fragment from Ohio in the British-
Museum (fig. 6, C, p. 170), show some of the orbital and other marginal
bones in top view in articulation with the skull-roof, but no specimen we have
yet seen gives much help in reconstructing the orbit itself. The hypothetical
arrangement which we suggest in fig. 20, p. 186, is chiefly based on an
examination of the bones themselves, taking account of the edges bearing
orbital notches, the edges with articular faces, and the edges which were
evidently free. It accounts for all the recognizable types of circumorbitals
which occur in the Atthey Collection. Some of the skulls referred to above
show that the smaller marginal bones articulating with the nasals and with
the fore-part of the frontals did not enter into the orbit, and we have
accordingly represented them as intermediate rceofing ossicles.

The Palate.

Neither we nor previous workers have found indications of any kind of
ossification of the brain-case, which must have been as completely cartila-
ginous as it is in Ceratodus. Where the skull-roof and the palate are
undisturbed they lie directly one upon the other, usually with hardly a trace
even of the matrix between them.

The bones of the palate bave been well known for many years, and the
additional observations that we have to make refer mainly to certain details.
of their relations to one another. Like all the other bones of Sagenodus, the
parasphenoid is very variable in form within definite limits, and especially
so in regard to the development of a median ridge on the buccal face of its:
main “lozenge” and in the shape and prominence of the lozenge at its
hinder angle. Common types are represented in fig. 9, p. 174. The para-
sphenoids collected at Newsham were supposed by Hancock and Atthey to
belong to about half-a-dozen distinct species, a view which Miall dissented
from, though. it is not without some measure of support.

Even before the existing Ceratodus became known, it was seen in a general
way how the pterygoids with their tooth-plates must have fitted round the
lozenge of the parasphenoid. Attempts to depict the bones in their natural
position have not, however, been altogether happy. Miall’s restoration
(1874, pl. 47), which is based on the similar palate of Ctenodus, is the most. |
successful, but he shows the hinder ends of the pterygoids projecting as loose:
frills beyond the lozenge. Williston’s figure (1899, pl. 37. fig. 2) has the
same defect, and in addition his pterygoids lack most of their hinder wing,,

172 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

and he shows the parasphenoid with its cranial face turned to the mouth.
‘The cranial and buceal faces of the parasphenoid had been correctly identified
by Miall (1881, fig. 3).

The way in which the pterygoids actually fitted on to the parasphenoid is
perfectly shown by two specimens in the Royal Scottish Museum (Nos. 1902.
73 and 1894.168.2). These actually belong to an early form of Ctenodus
(the form to which Traquair applied Barkas’s name Ctenodus interruptus) ;
but as this form had the palate of a Sagenodus, and as the differences between
typical palates of the two genera are in any case slight, it is appropriate to
refer to them here. The more perfect of the two specimens is depicted,
from a rough sketch, in fig. 24, p. 193. It shows that the lateral corners
of the parasphenoid lozenge, looked at from the buceal aspect, overlay the
edges of the pterygoid, but that towards the front of the lozenge these edges
passed on to its buccal face and met each other across it by a square corner
at the hinder end of their symphysis. Well-preserved examples of the

Fig. 7.

Sagenodus. Sub-operculum, x $. A, outer surface ; B, inner surface of another example.

pterygoid (fig. 8, A, p. 173), as a matter of fact, always show a roughened
surface corresponding to the corner of the parasphenoid to which they were
articulated. (It is of the same character as the corresponding narrower
surface on the pterygoids of Ceratodus, and has been figured by Atthey and
by Williston.) They also show the corner at the hinder end of the symphysis
by which they met their fellow across the face of the parasphenoid, and the
imprint of these corners is occasionally visible on a detached parasphenoid.
It is only in “ Ctenodus interruptus” that the corner is square. In other
forms both of Ctenodus and Sagenodus it is obtuse and meets its fellow in a
forwardly-pointing angle. ‘The arrangement of the elements of the palate in
a typical Sagenodus in seen in fig. 10, p. 176.

The meeting of the pterygoids across the front of the parasphenoid may
in part explain the singularly weak pterygoid symphysis. In Ceratodus the
pterygoid symphysis is very much stronger than that of the lower jaw ; in
Sagenodus it is altogether weaker, for the bones meet only in a long, thin

I

STRUCTURE OF CERTAIN PALHOZOIC DIPNOI. 173

edge. At the back, however, this weak symphysis is supported on the
cranial face for some part of its length by the anterior process of the
parasphenoid, though that still leaves the forward part and the long anterior
ridges of the tooth-plates to all appearance very insufficiently provided
against the strains of mastication. This region, however, no doubt received
further support from the overlying cartilaginous cranium.

Fie. 8.

A, Sagenodus; B, Ctenodus. Left pterygoid and tooth-plate, x 3.

As they occur in the shales, the pterygoids and their tooth-plates are
nearly always crushed into one plane. There can be no doubt, however,
that in life the wing of the pterygoid would be bent downward along its
outer edge to support the quadrate. It may not have been bent so sharply
as in Ceratodus, where it stands at right-angles to the tooth-plate, but that
it was considerably bent is proved by the abundant cracks that break the

174 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

surface of these flattened bones, as well as by the curve actually shown by the
least crushed examples—among them some very small pterygoids which by
their minuteness have escaped the worst of the crushing.

The facet on the j“toe” of the pterygoid was remarked upon by Miall
(1881, p. 292). It is precisely like the corresponding facet in Ceratodus,
except that in Ceratodus the facet itself is composed of cartilage; and
it doubtless had the same function, namely to support the quadrate and
strenethen the articulation of the lower jaw.

The Quadrate.
The bone which we take to be the quadrate has long been known as a
bone of Sagenodus, but has been assigned to various positions. Fritsch

lia. 9.

Sagenodus. Parasphenoids, x about 3.
A and B, palatal surface (B, the commoner type) ; C, dorsal surface.

usually identified it as a scapula (“Schulterblatt”), as in his Taf. 77,
figs. 1 and 4 (1889), but another example, Taf. 77, fig. 12, is labelled
‘““Femur?”. The same bone was well figured by Williston (1899, pl. 28,
fiz. 3), who suggested that it might be the ceratohyal. It is a large.
bone in proportion to the size of the head, but in a Dipnoan with a
head deep enough to accommodate such an operculum as that of Sagenodus
the quadrate would necessarily be large. Lvyen as it is, there is evidence
that its upper end was incompletely ossified. The ossification of the rest
of the bone, too, was more or less superficial; though carried much further
than in the quadrate of Ceratodus. As they are found in the shales, the
bones have always collapsed under pressure as the core of cartilage decayed.

STRUCTURE OF CERTAIN PALMOZOIC DIPNOI. 175

Fig. 11, C and D, p. 177, show good examples from the Low Main shale,
Newsham, in the Atthey Collection. Both ends of the bone were expanded,
the lower more so than the upper. One side of the shaft bore a pronounced
longitudinal ridge or wing, better preserved in Williston’s and some of
Fritsch’s examples than in ours. The aspect shown in fig. 11, C, p. 177,
we take to be the inside of a right quadrate, and we suggest that the facet
on the toe of the pterygoid was applied to the prominence there shown at the
lower end of the bone on the right, possibly also engaging with part of the
longitudinal wing.

We have been unable to identify any further bones as belonging to the
skull proper. Vomerine teeth were described and figured by Atthey (1877),
and we see no reason to doubt his identification ; but no trace of bone is
attached to these teeth, and it is probable that the vomers were represented
solely by the bases of the teeth, as in Ceratodus.

We have not been able, either, to identify any of the hyoid bones with
certainty. Many large rib-like bones occur on slabs of shale among the
other remains of Sagenodus, and there can be little doubt that some of them
are hyoids, others probably cranial ribs. Fritsch was confident that he had
identified both ; the bone he named “first cranial rib ” (1889, Taf. 77, fig. 5)
may well be such, but some of his “hyoids,” especially that figured at
Taf. 71, fig. 5, are probably not bones of Sagenodus at all *.

The Lower Jaw.
The main features of the structure of the lower jaw were made out many

years ago. Hancock and Atthey described and figured the “ splenial” in
1872; and in 1877 Atthey announced the discovery of the angular
(“articular”), which he had been able to identify by comparison with the
newly-discovered Ceratodus forsteri. To complete the resemblance of the
jaw to that of Ceratodus, a “dentary” f element was needed, and its
existence in Sagenodus was first demonstrated by Watson and Day (1916),
who found it in the form of a broken cross-section fitting on to the
front of the angulars in a head in the Manchester Museum (L. 10904).
‘In the Atthey Collection we have found an abundance of examples of the
bone itself, some lying among the other remains of crushed heads and
many others freed from the matrix. In the Dinning Collection, also in

* Fritsch believed (p. 67) that he could recognize the bones of “ Ctenodus obliquus” with
eertainty by their lustre and colour, and in a few instances this criterion possibly led him
astray.

+ As will appear from a later discussion of the jaw of Dzpterus, there is good ground
for thinking that the bones here referred to as “splenial” and “ dentary ’ are not properly
so named. But since, following Huxley (P. Z. 8. 1876, p. 34), the corresponding bones in
Cerdtodus have long been known by those names, we retain them in this description, only
placing them in quotation-marks. ;

LINN. JOURN.—ZOOLOGY, VOL. XXXY. 12

176 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

the Haneock Museum, there is a small angular with the “dentary” still
in natural articulation with it. This specimen bears a label in Atthey’s
handwriting, and as he called it simply “ Ctenodus, articular bone,” he
presumably thought it merely a damaged or aberrant angular. It is
represented in fig. 14, p 180.

The figures here given will make it unnecessary to describe the jaw at
length. Restorations in different aspects are given in fig. 12, A, B, C, D,
p. 178, and corresponding views of the lower jaw of Ceratodus, after removal
of the cartilage, are given in fig. 13, p.179. It will be seen that apart
from the difference in the tooth-plates the two jaws are remarkably alike.
The bones in Sagenodus are very much stouter, but that is a difference that
obtains in nearly all parts of the skeleton.

Sagenodus. Reconstruction of palate, showing pterygoids in natural
articulation with the parasphenoid. The postero-lateral wings
(quadrate rami) of the pterygoids are shown flattened into
the same plane as the parasphenoid.

The bones that differ most in the two forms are the “dentaries,” for
though they are much alike as seen from below, the “dentaries” in
Sagenodus take a more essential share in forming the jaw than they do in
Ceratodus. In the latter fish they are papery bones feebly connected with
the angulars by a very slight groove near the edge. In Sagenodus (fig. 15,
p. 180) they are strong bones firmly united with the angulars, which have a
pronounced groove for their reception along the anterior half of their lower
edge. But the most striking difference in the “dentaries” of the two fishes

STRUCTURE OF CERTAIN PALAOZOIC DIPNOT. WAC

lies in the form of the hind border. In the “ dentary ” of Ceratodus this
border is thin and papery like the rest of the bone. In the Sagenodus
“dentary ”’ it is widened abruptly, and presents a deep, polished face toward
the gular space (fig. 15, D, p. 180). This part of the jaw is, in fact,
finished off in almost~ precisely the same way as the corresponding part in
Dipterus, and, as we show later, for the same purpose : to forma bearing for
gular plates.

All the elements of the jaw are as variable as the teeth have long been
known to be. Fig. 12 E, p. 178, shows that some jaws are deeper than that
figured above it at C, and certain angulars in the Atthey Collection would fit

cS >)

Supposed Quadrates, x about 2.
A, B. Supposed to be of Ctenodus. C,D. Sagenodus. (A and B are opposite sides
of the same specimen: C and D are opposite sides of two different specimens.)

ajaw of this deeper form. Other angulars suggest an even shallower jaw
than C. Well-preserved angulars show that both the upper and lower borders
of the bone were rolled inwards very considerably ; the lower part towards
the middle of its length formed a sort of flopr inside (fig. 12, F, p. 178),
and the upper edge bears a bracket for the support of the back of the tooth-
plate, as was noticed by Atthey (1877, p. 228). Very small angulars, of
3 or 4 em. in length, not having been seriously crushed, exhibit this inward
iQ

178 PROF. D. M.S. WATSON AND MR. E. L. GILL ON THE

rolling of the upper and lower borders very markedly. The ‘“‘ dentaries ” are
even more variable than the other bones of the jaw, both as regards shape
and surface character. Three different patterns are shown in fig. 15, p. 180.
The ‘dentary ” there marked C is one of the shortest in relative length and
deepest at the symphysis ; some are even narrower than the one figured at A.
The symphysis itself is commonly of the character shown in D, much like
that of the “ splenial” in fact ; but a quite different type occurs in which

the “ dentaries” were united by interlocking spines, as shown at A. The

Fie. 12.

Sagenodus. Reconstruction of. lower jaw, x 2. A, dorsal, B, ventral, ©, lateral
D, medial aspects; I, “splenial,” from the outer side; FP, angular, eT Tne eral
side;-ang., angular; “ den.,” “ dentary ” (=splenial); sp., splenial (Spresanticuien)
(Compare with figs. 13 of Ceratudus and 27 of Ctenodus.) Ei is &. imbricatus ; aie

rest of the figures S. obliquus. The deep symphysis of EH can be matched in the
latter species.

line of sensory pits in the angular is often continued on the “

dentary,” and
a . . . <3 s ? .
where this is the case one of the pits seems regularly to lie

on the suture
(fig. 14, p. 180). Near the posterior border of the “ dentary ”’ there is

often a row of three or four much smaller pits. In some pairs of
‘c Pter eta. epT yO , q ;
dentaries” the posterior borders would apparently meet across the middle
line in an even curve; in others thes rders
; In others these borders are strongly hooked at the

STRUCTURE OF CERTAIN PAL#OZOIC DIPNOJ. 179

symphysis, sometimes considerably more so than in fig. 15, C, p. 180, and
together they must have produced a small backward procvess at thie middle
line (fig. 16, B, p. 181), a difference that would be reflected in the shape of
the gular plates which adjoined the “ dentaries ” behind.

Fre. 13.

Ceratodus (Neoceratodus) forstert. The bones and tooth-plates of the lower jaw. A, dorsal,
B, ventral, C, lateral, D, mesial aspects; HW, ‘*splenial,” from the outer side;
F, angular, from the mesial side. (Reference letters as in fig. 12.)

The Gular Plates.

The absence of gular plates has been given as one of the diagnostic points
separating the Ctenodontidee from the Dipteridee. The similarity of the
hinder border of the “ dentaries ” in Sagenodus and Inpterus led us, however,
to look for something in Sagenodus to correspond with the gulars which are
applied to this border in Dipterus, and this we believe we have found in the
bone represented in fig. 16, A, p. 181. There are three examples of this
bone in the Atthey Collection, One of them occurs on a slab of shale

180 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

among the scattered but practically complete remains of a head of Sagenodus ;
the other two are detached. It is a bone of graceful form, delicately striated
on one face in lines converging towards a small tubercle near the blunt end.
The other face is smooth. Fig. 16, B, p. 181, shows the size of the bone in
relation to the lower jaw and the manner in which we suggest that it fitted

Fie. 14.

Sagenodus sp. Right angular and “ dentary ” in natural association, X 14.

in the gular space. Compared with the extensive gular apparatus of
Dipterus, described later, it is meagre, but it is much what might be looked
for in a fish which in so many respects is a transitional form between the
early Dipnoi and Ceratodus.

Fre. 15.

Sagenodus. A,B, ©, outer surface of three ‘“ dentaries,” x about 3; D, inner
surface of a left “dentary,” oblique view showing the character
of the symphysis and of the hinder border.

The Opercular Bones.

Next to the teeth, the operculum is probably the best-known of the remains
of Sagenodus. In proportion to the head as a whole, it is remarkable for
its size and massiveness. Its manner of attachment to the skull has been
described already (p. 169). In common with the other bones, it varies greatly

STRUCTURE OF CERTAIN PALHOZOIC DIPNOT. 181

in outline, but the form shown in fig. 20, p. 186, is a good average. The
variation consists for the most part in the greater or less development of the
hinder lobe of the bone. There is every gradation, independently of size,
from a form in which the body of the bone is fairly symmetrically disposed
under the centre of the hinge-line, to an extreme but quite common pattern
in which the hinder border is so prominent that the centre of the bone lies
below the posterior knob, behind the hinge-line altogether. In this latter
pattern of operculum the posterior knob is ‘always greatly developed and
the anterior one almost obsolete. Ovpercula of this type have been figured
by Miall (1881, fig. 7, where they are shown upside down), and by Williston
(1899, pls. 35 & 36) in-his deseription of S. copeanus.

A small bone, hitherto undescribed, which we take to be the Sud-
operculum, occurs in several of the crushed heads of Sagenodus which we

Fria. 16.

Sagenodus. A, left? gular, inner surface; B, diagram, showing the
probable position of the gulars.

have examined; it is shown, for example, on the slab, part of which is
represented in fig. 18, p. 183, and in the head in the Manchester Museum
(L. 10904) described and figured by Watson and Day (1916). It is a
bone of an inch to an inch-and-a-half in length in an average skull, and is
recognizable by its triangular point (see fig. 18, A, p. 183). Completely
preserved examples, such as those represented in fig. 7, p. 172, show that
half the bone was fairly thick and strengthened by a rounded ridge, while
the other half consisted of a thin flange which might easily be lost. Its
disappearance would give the rest of the bone a much more pointed shape,
such as is seen in fig. 18, p. 183. The sub-operculum of Ceratodus (inter-
operculum, Huxley), to judge by a dried skull, would much resemble this
bone if its cartilaginous fringes were ossified. No specimen that we have
seen shows this bone in its natural position, but on two out of the three slabs
on which we have found it, it lies close to the squamosal, which may indicate
that it was applied to the front rather than to the hind border of the

Sagenodus. Bones of the shoulder-girdle. A, right post-temporal, outer surface; B, the
same, inner surface; C, right cleithrum, outer surface; D, the same, inner surface ;
Ii, right clavicle, upper (chiefly internal) surface; F, the same, lower surface ;
G, right clavicle, imperfect, to show the cracks which result from the flattening of
the bone. In E and I the bones are represented as flattened, but without the
cracks. X 3.
(The elements of the shoulder-girdle are seen in natural association in fig, 19, p. 185.)

STRUCTURE OF CERTAIN PALMOZOIC DIPNOT. 183

Fie, 18.

|

| ) Pe

oN \\\
Ope ae AS \

‘
‘

pt. tem.

B

Sagenodus. Anterior portion of a specimen in the Atthey Collection, x about 3. A, dorsal
B, ventral surface. clav./., left clavicle ; clav.., right clavicle; clei.r., right cleithrum
~ cor, cixcumorhital; op.l., left operculum ; op.r., right operculum; par.sp., parasphenoid ;
pt., pterygoid; pt.tem., post-temporals; s.op. sub-operculum; tad.h., left tabular horn
covered by a scale.

184 PROF. D. M. 8. WATSON AND MR. E. L. GILL ON THE

operculum. In the restoration (fig. 20, p. 186) we have assumed that the
edge of the operculum overlapped the thin flange of the sub-operculum.

A bone labelled “ subopercular ?” was figured by Williston (1899, pl. 36,
fig. 3). Though it is not much like our bone in shape, it may possibly be
another form of it.

The Shoulder-Girdle.

The first bone of the shoulder-girdle to be recognized was the clavicle,
which was correctly described by Hancock and Atthey (1872). Since then
the only investigator who has contributed usefully to what is known of this
part of the skeleton is Miall, who not only published figures (rough and
imperfect it is true) of the clavicle (1881, fig. 11, “ coracoid”), but also
figured the cleithrum (fig. 10, “ scapula”), till then unknown, and made
(pp. 296-7) some suggestions, now in the main confirmed, as to the way
in which these bones were arranged in the girdle. Miall added another
bone (fig. 9) as the ‘“‘ supra-scapula,” but we can find nothing like it, and
it certainly forms no part of the shoulder-girdle of Sagenodus. Fritsch
discussed the shoulder-girdle at length, but the upshot, in the restoration
which he gave in text-ligures 158 and 160, p. 81, was anything but a happy
application of his perfectly sound principle that Ceratodus was the best
guide to the structure of “ Ctenodus.”

In the case of the shoulder-girdle, again, the figures here given (figs. 17,
p. 182, and 19, p. 185) may largely take the place of a detailed description ;
and here again, as with the lower jaw, there is such a strong similarity to
the corresponding parts in Ceratodus that the comparative outlines given
in fig. 19 are practically self-explanatory.

The history of the discovery of the clavicle and eleithrum has been
indicated above. The third bone, the post-temporal, has also long been
known. It was fully discussed by both Miall and Fritsch, and on the
strength of a certain resemblance to the combined squamosal and quadrate
of Ceratodus they both figured it as the squamosal (Miall, fig. 6 ; Fritsch,
text-fig. 153, p. 75). The clue to its real nature, as well as to the arranye-
ment of the shoulder-girdle in general, was given by a fine specimen in the
Atthey Collection, the pertinent portion of which is represented in fig. 18,
p. 183. On its upper surface this specimen is chiefly noteworthy as being
the only example we have met with which shows the operculum in its natural
relation with the skull. It also shows the upper end of the right cleithrum
from its outer aspect. On the under surface, as it was left by Atthey, the
most prominent objects were the two clavicles, lying in nearly their natural
position ; the removal of masses of overlying scales has disclosed other
structures which appear in our figure (fig. 18, B, p. 183). The right
cleithrum and clavicle are in natural articulation. Underlying the left
clavicle are the two post-temporals, the left one showing its strong anterior
process projecting beyond the clavicle in front. Underlying the post-

STRUCTURE OF CERTAIN PALAZOZOIC DIPNOT. 185

temporals, again, is the parasphenoid, the backward prolongation of which
is seen emerging from beneath them. The only other specimen we have
seen showing the connection of cleithrum and clavicle is a detached example
of these two bones in the Royal Scottish Museum.

A few remarks are called for on points of detail. As with other bones of
the skeleton, those of the shoulder-girdle are variable, the cleithrum especially
so. The inner face of the cleithrum is flattened; the outer is convexly
thickened towards the upper end, and is strengthened by ribs of bone running

Fie. 19.

Sagenodus and Ceratodus. Comparison of shoulder-girdles.
A, B. Right side of shoulder-girdle of Sagenodus.
(©), 10); s ‘ % Ceratodus, with the bones supposed
flattened as are the fossil bones of Sagenodus.
E, F. Right side of shoulder-girdle of Ceratodus, with the bones in natural shape.
(Outer views on left, inner on right.)

upwards from the junction with the clavicle. Crushing has often resulted in
these ribs being more or less plainly printed through on to the inner face.
The cleithrum ends below in a transverse edge which is received into the
head of the clavicle, and its hinder border is produced into a flattened and
pointed process which fits into a slot in the back of the clavicle. The
expanded upper end of the clavicle, with its deep pit for the attachment of

186 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

muscle, forms the “hatchet-shape ” which Hancock and Atthey attributed
to this bone.

Apart from the much greater strength of the bones in Sagenodus, the
points of difference between its shoulder-girdle and that of Ceratodus are
insignificant (fig. 19, p. 185). The strong forward process of the post-
temporal in Sagenodus is plainly associated with the presence of a tabular
horn, to which it was doubtless attached by its truncated point. In
Ceratodus the post-temporal is merely attached to cartilage. The process of
the cleithrum which serves to stiffen the union with the clavicle is repre-
sented in Ceratodus by a pyramidal point (fig. 19, p.), which if the bones
-were flattened would lie in the middle of the internal face instead of on the
hinder border. In Ceratodus the muscle-pit in the head of the clavicle is
divided by ridges into two or three pockets, but some of the clavicles of
Sagenodus show an approach to the same condition. Finally, there can
be no doubt that the blade of the clavicle was twisted as it is in Ceratodus,
so that what appear in the flattened bones (fig. 19, A, B, p. 185) as its upper

pam

wr

clav. S.Op.

Sagenodus. Restoration of skull and shoulder-girdle, in side view, x 4. c.or., two of the
circumorbitals ; vm.t., yornerine ‘‘ tooth.” Each bone is founded on specimens in the
Atthey Collection, but the association of the cireumorbitals is conjectural.

and lower edges were in life the inner and outer edges respectively ; while
the surface represented in fig. 17, E, p. 182, and fig. 19, A, p. 185, is really
the upper and internal surface, and that represented in fig. 17, F, and
fig. 19, B, the lower and external. This will be understood from a comparison
between the outline drawings in fig. 19, where the middle figures, C and D,
show the shoulder-virdle of Ceratodus as it would appeat if it were flattened
out, like the bones of Sagenodus, by fossilization in shale. The actual
specimens of the clavicle of Sagenodus show abundant evidence of haying
been strongly curved, for the flattening has produced a system of gaping
eracks on the smooth inner face of the bones (fig. 17, G, p. 182). One

1

STRUCTURE OF CERTAIN PALAOZOIC DIPNOI. 187

remarkable specimen occurring on a slab of shale in the Atthey Collection
has, indeed, retained its original shape, though surrounded by other bones of
Sagenodus which have all suffered the usual flattening. It should be added .
that the faces of the clavicle identified above as upper-and-internal and as
lower-and-external have, as the figures show, all the character of internal
and external faces respectively. Miall described and figured an articular
area on the broad lower end of the clavicle, and he suggested that the
two clavicles met each other at an acute angle in this articulation. Well-
preserved examples, however, show no such articular area, nor would
they fit together at all exactly by their ends. In all probability they
were connected, as in Ceratodus, by an interclavicular cartilage.

We have nothing to add to what has already been published regarding the
structure of the body, fins, and scales. There is every reason to suppose
that these parts of Sagenodus are closely reproduced in the existing
Ceratodus.

The Species of Sagenodus.

A glance at the portions of cranial roofs shown in outline in fig. 2, p. 165,
will suggest that they represent more than one species of Sagenodus. At
least one further species is certainly represented by the zoned and polished
skull-bones found so abundantly in the Bohemian Gaskohle and in some
numbers also at Newsham (cf. fig. 4. D, p. 168). But there is rarely anything
distinctive about the teeth in the cases where it is possible to assign them
with certainty to any particular pattern of skull, and there is little ground
for attaching the slightest value to most of the specific names founded so
freely on the teeth. Most parts of the skeleton, and especially perhaps the
parasphenoids, the separate elements of the lower jaw, and the opereula,
show varieties of pattern as important as those of the cranial roof and the
teeth ; and until the different patterns of all these parts have been studied
and correlated there will be little use in applying specific names to Sagenodus
at all. Sucha study would be extremely difficult on account of the frag-
mentary nature of the evidence, and it might very possibly result in the
conclusion that Sagenodus was, in its sphere of life, a dominant form in such
a fluid evolutionary stage that it would be for the most part impossible to
apply to it the ordinary conceptions of a species.

.

Fritsch’s figures of the Cranial Bones of Sagenodus.

Fritsch’s figures are beautifully drawn, and form a valuable atlas of the detached bones of
Sagenodus, but the fact that so many of them are unidentified detracts from their usefulness.
Some of these unidentified bones are named in the following list :—

Plate 71 (Kauna der Gaskohle, vol. ii. part 2, 1899): fig. 10, right intertemporal; fig. 11,
right nasal.

Plate 72, fiz. 10, left “ tabular.”

Plate 74 includes a good selection of different forms of ‘ parietals”’; fig. 5, an extreme
example of the rectangular type.

188 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

Plate 75, figs. 1-11, marginal ossicles, but including some ordinary roof-bones of small
fishes, e.g. fig. 5, squamosal; 3, intertemporal: fig. 14, left intertemporal; 12, 19,
circumorbitals; 20, 1. “tabular ’’; 22, vr. intertemporal; 24, 1. squamosal; 25, 1.
frontal; 27, y. “tabular”; 28, ry. squamosal, underside; 29, 1. frontal, underside ;
32, 1. “tabular,” underside, with one of its lateral bones; 33, nasal, underside ;
34, fragment of squamosal ; 37, marginal ossicles; 39, r. nasal.

Plate 77, fig. 17, ‘ tabular.” .

Plate 78, fig. 6, post-temporal.

Crmnopus, Agassiz.

The genera Ctenodus and Sagenodus were separated primarily on the
characters of the tooth-plates, though when Owen first published the name
Sagenodus he was under a misapprehension as to the nature of the tooth-
section on which he founded it. Since the publication of Dr. Smith
Woodward’s ‘Catalogue of Fossil Fishes” (vol. ii., 1891), the name Ctenodus
has been generally applied to tooth-plates with about 12 or more ridges,
roughly parallel, as contrasted with the others (Sagenodus) having fewer,
usually 6 or 7, ridges, with a strongly-marked radial arrangement. If this
were the only distinction between the two genera it would be difficult to
maintain, since tooth-plates of intermediate character occur in the Lower
Carboniferous of Edinburgh. ‘There are, however, many other distinctive
characters independent of the difference in the teeth; and in spite of a
general resemblance which shows the two genera to be nearly related, the
additions which we are able to make to what was known of each of them
tend still further to justify their separation.

Sagenodus having been dealt with already, Ctenodus can be sufficiently
described, with the aid of figures, on brief and comparative lines.

In most or all of the beds and horizons from which we have had material
for examination, remains of Ctenodus are much less abundant than those of
Sagenodus, and there is consequently more difficulty in ascertaining its
skeletal structure with completeness, In particular there is in the collections,
up to the present, an almost entire absence of specimens such as were of the
greatest help in the case of Sagenodus, namely slabs bearing the scattered
remains of some considerable part of an individual fish. We have never yet,
for example, seen any considerable part of the shoulder-girdle of Ctenodus
in“association with other portions of its skeleton, and the bones which we
take to be those of its shoulder-girdle are assigned to it without absolute
proof, though on strong grounds of probability.

The fishes of the genus Ctenodus were of decidedly larger average size
thaa those comprised in Sagenodus. Nine or ten inches is a usual length for
skulls of Ctenodus, as compared with five or six inches for skulls of
Sagenodus.

STRUCTURE OF CERTAIN PALMOZOIC DIPNOT. 189

The Roof of the Skull.

The skull of Ctenodus was ossified to the same extent as that of Sagenodus,
and similarly shows no trace of ossification of the neural chondrocranium.
A comparison of fig. 1, p. 164, and fie. 21, below, will show that the skull-
roofs of the two genera differ considerably—more, in fact, than any other

Fre. 21.

Ctenodus cristatus. Cranial roof, fr., frontal; 2.fr., interfrontal; ¢.nqa., internasal; ¢.tem.,

~ intertemporal; maz., marginal bones; par., “parietal” (=parietal + dermo-supra-

occipital); p.fr., pre- or post-frontal; p.na. prenasal; sg., “‘squamosal”; tab.,
“tabular” (=tabular + supratemporal). x about 4.

190 PROF. D. M. 8S. WATSON AND MR. E. L. GILL ON THE

part of the skeleton,—but that the general plan isthe same. In the posterior
region the correspondence is very close. ‘he main points of difference may be
broadly stated thus: in Ctenodus the two median bones (par. and i.fr.) are
much smaller and the bones of the nasal region much larger, \ hile there are
two pairs of bones in the frontal region instead of only one. As far as its
skull-roof is concerned, Ctenodus, being nearer to the Devonian Dipterus, is
presumably the more primitive ; and we may usefully think of the skull-roof
of Sagenodus as being derived from that of Ctenodus by a great reduction of
the snout region together with an increase in size and eventual meeting of
the two median bones, both processes together resulting in the compression,
and finally in the fusion, of the frontal and post-frontal on each side.

As long ago as 1872 Hancock and Atthey correctly pointed out, as one of
the main features distinguishing the skull of Ctenodus proper from that of
Sagenodus (“ Ctenodus obliquus”’), that the two median bones of the roof are
separated and that the hinder one consequently has a pointed instead of a
concave anterior margin. Dr. Smith Woodward (1891, p. 250) refers to the
same distinction when he states that there are ‘‘ two median occipital plates ”
in Sagenodus and only one in Ctenodus. He also gives (pp. 252-3) the only
extended description hitherto published of the skull-roof in Ctenodus, but
the specimen on which he founded it (B.M., P. 5031), figured in pl. 4. vol. ii.,
of his catalogue, is not well preserved; it shows for the most part impres-
sions of the under surface of the bones, and their outlines are too indistinct
for accurate representation. The same specimen was roughly figured by
Fritsch (1889, text-fig. 156), but as he failed even to identify the middle line,
his observations on it (p. 98) were not helpful.

Another specimen (now in the British Museum, P. 7300) was also roughly
figured by Fritsch in his text-figure 155, and is represented in outline in
our fig. 23, A, p. 192. ‘It is chiefly remarkable as showing a considerable
ossification in advance of the nasals, in the form of a radially-ribbed fan.
Fragments of a similar fan in the Atthey Collection are shown in fig. 22,
p. 191. Except at their outer edge, the bone composing them is as thick
as that of any other part of the cranial roof.

The presence of an internasal is a further point of distinction from
Sagenodus. The form of internasal shown in fig. 21, p. 189, of which two
specimens are known, is possibly characteristic of Ctenodus cristatus. The
form figured in B, fig. 23, p. 192, may similarly be characteristic of
“ OC. interruptus.”

The bones of the roof proper in Ctenodus seem to be about as variable in
shape as in Sagenodus. The specimens we have examined show that the
most variable elements are the interfrontal, internasal, and post-frontal
(compare fig. 21, p. 189, with the outlines in fig. 23, p. 192). From the
condition of the available specimens it would appear that the elements of the
cranial roof of Ctenodus were more firmly united than those of Sagenodus,

STRUCTURE OF CERTAIN PALHOZOIC DIPNOI. 191

for they very rarely occur as isolated bones, whereas isolated roof-bones of
Sagenodus are abundant.

The cireumorbitals of Ctenodus are shown only in one specimen among
our material, and this is represented in outline in fig. 23, B, p.192. In
number and in general arrangement they appear to differ little from the
circumorbitals of Sagenodus, except that a small bone, partially embraced by
the squamosal and post-frontal, occurs between them and the bones which
actually enter into the orbit. This, however, may be an arrangement
peculiar to ‘* Ctenodus interruptus.”

In addition to the cireumorbitals there are, as in Sagenodus, some bones
fillmg in the space to the outer side of the nasals. They are not completely
shown in any specimen that we have seen, but they obviously vary a good
deal in shape (mar., fig. 21, p. 189, and fig. 23, p. 192). The single one shown
in fig. 21 is represented as an impression of the underside, which is all that
had been seen of it when the figure was drawn.

Fie. 22.

Fraements of the prenasal fan of Ctenodus, x 3.

The Palate.

A comparison of fig. 10, p. 176, and fig. 26, p. 195, will show the close
resemblance between the bones of the palate in Ctenodus and Sagenodus.
The tooth-plate of Ctenodus is larger (fig. 8, p. 173), the pterygoid behind it
is of rather more slender form, and the parasphenoid (fig. 25, p. 194) has
a more prominent median ridge on the buccal face of the lozenge. The
shaft of the parasphenoid is more abruptly expanded, and shows a pair of
pits close behind the apex of the lozenge. On the cranial surface the ridges
and grooves of the shaft are much more numerous than in Sagenodus (cf.
fig. 9, C, p. 174), and are continued forward to the centre of the lozenge,
where they nearly meet the corresponding ridges from the anterior process.

In the larger parasphenoids, some of which are nearly a foot in length,
the median ule of the lozenge is swollen and club- shaped (fig. 26, p. 195),
and it seems to be usual for the right-hand pit of the pair behind it to com-
municate directly with the central groove of the shaft, as shown in fig. 25, C,
p- 194. The appearance is suggestive of the crossing of a pair of longitudinal

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 13

1192 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

muscles. The species “ Ctenodus interruptus” from the Lower Carboniferous,
though proved by its cranial roof (fig. 23, B, p. 192) to be a true Ctenodus,
has a palate and tooth-plates which in all respects much more nearly resemble
those of a Sagenodus (see fig. 24, p. 193). :

The Quadrate.

The bone which we take to be the quadrate of Ctenodus is represented in
fig. 11, A and B, p.177. There are two or three examples of it in the
Atthey Collection, and one of them was labelled by Atthey himself “ Ctenodus,
os quadratum.” Fig. 11, p. 177, shows sufficiently how it differs from the
quadrate of Sagenodus, the most striking difference being the much smaller
development of the longitudinal ridge on the inner side.

CTENODUS. -

A. Cranial roof of a specimen of C. erzstatus from Longton, Stafts. B.M.N.H. 7300.
B. Cranial roof of ‘ Ctenodus interruptus, Bavkas,” Dunnet shale, Straiton, Midlothian.
1895/155/12, R.S.M. Of importance as being the most perfect cranial roof of this

species existing, and showing more connected circumorbitals than any other specimen

of Ctenodus known. The detached circumorbitals are preserved on the counter slab,
and really partially underlie those which are attached to the roof. A palatal tooth,
omitted from the figure for the sale of clearness, establishes the specific identity of
the specimen. X 3.

The Lower Jaw.
Remains of the lower jaw of Ctenodus are scarce, but specimens from

Newsham in the Atthey Collection and in the British Museum make it
possible to reconstruct the jaw as has been done in fig. 27, p. 196. In
general plan, and to a large extent in detail as well, it agrees with the jaws
of Sagenodus (fig. 12, p. 178) and Ceratodus (fig. 13, p. 179). The chief
difference is in the greater breadth of the tooth-plates and, correlated with
it, the greater width of the *‘dentaries.” Instead of coming almost or quite
to a point in front, as do the ‘“ dentaries” of Sagenodus, those of Ctenodus
have a wide anterior edge and carry the front ends of the angulars some

STRUCTURE OF CERTAIN PALMOZOIC DIPNOI. 193

distance outwards from the middle line. Another result of the greater
breadth of the tooth-plate is that it is supported posteriorly, not, as in
Sagenodus, by a bracket on the inner face of the upper border of the angular,
but bya pocket in the same position on the outer face (see fig. 27, H, p. 196).
The “‘splenial” is a much weaker bone than in either Sagenodus or Ceratodus ;
its symphysial end is fairly strong and some part of it is commonly to be
seen projecting in front of the mandibular teeth, but behind the symphysis
itis so thin that it has usually been folded or crumpled beneath the tooth-
plate in fossilization.

The posterior edge of the “ dentaries” is thickened and polished precisely
as in Sagenodus, and no doubt there were gular plates fitting against it, but
so far we have not found any bones that we could identify as gulars of
Ctenodus.

Fie. 24.

“ Ctenodus interruptus, Barkas.”

Pterygoid with teeth and parasphenoid, oral aspect, X 3.
Oil shale of Broxburn, Midlothian (1902-73, R.S.M.).
The left pterygoid is entirely, the right very nearly in natural
articulation with the parasphenoid.

The Operculum seems to have been exactly like that of Sagenodus.
Several specimens in the Atthey Collection are too large for any known
skull of Sagenodus, and these at least presumably belong to Ctenodus ;. but,
apart from maximum size, we can point to no definite character by which
opercula of the two genera may be distinguished. A sub-operculum was
doubtless present, but we have not yet succeeded in identifying it.

The Shoulder-Girdle.
As already stated, we have never yet seen well-preserved bones of

the shoulder-girdle associated with undoubted remains of Ctenodus.

194 PROF. D. M.S. WATSON AND MR. E, L. GILL ON THE

Nevertheless we have little hesitation in regarding the bones shown in
fig. 28, p. 197, as the cleithrum and clavicle. Both are well represented in
the Atthey Collection; from comparison with the corresponding bones of
Sagenodus (fig. 17, p. 182), there can be no doubt that they are the cleithrum
and clavicle of a Dipnoan ; and both reach too large a size to belong to any
known form of Sagenodus. The cleithrum (fig. 28, A, p. 197) is found up
to 7 inches (33 cm.) in length, the clavicle to at least 6 inches (30 cm.).
The cleithrum differs from that of Sagenodus chiefly in having its hinder
half (the upper part in A, fig. 28, p. 197) greatly thickened instead of

Fie. 25,

Ctenodus cristatus. Parasphenoid.
A. Buceal aspect. B. Cranial aspect.
C. Junction of disc and shaft in large examples.

being strengthened by longitudinal ribs. Moreover, on its outer side this
thickened portion has the character of a superficial bone, as though it had
come to the surface under the skin behind the operculum as in Dipterus,
instead of being buried in muscle as in Ceratodus, and as it presumably was
in Sagenodus too.

The clavicle (fig. 28, B, C, p. 197), though it has the same essential
structure as that of Sagenodus, differs from it in having a longer articular
head and a narrower and stouter shaft. We have found no specimens with

STRUCTURE OF CERTAIN PALZOZOIC DIPNOI. 195

perfectly preserved heads, and our figure is therefore less complete in that
part than the corresponding figures for Sagenodus (fig. 17, p. 182). Miall
had apparently seen examples, though still more imperfect ones, of the same
bone (1880, fig. 12), and he evidently suspected that it belonged to the
shoulder-girdle of Ctenodus proper, for he labels it “ Ctenodus cristatus or
tuberculatus? Coracoid.”

We have found one or two bones which may prove to be post-temporals
of Ctenodus, but have not been able yet to identify them with any approach
to certainty.

Fie. 26.

Cienodus cristatus. Pterygoids with their teeth, in natural articulation
with the parasphenoid. Quadrate rami shown flattened into the same
plane as the parasphenoid. x 2.

The Species of Ctenodus.

Dr. Smith Woodward was probably well advised in reducing the specific
names tuberculatus, ovatus, etc., to the position of synonyms of C. cristatus.
The tooth-plates of Ctenodus, on which all the specific names have been
founded, are if anything more variable even than those of Sagenodus. They
are, in fact, likely to be so, since they represent a departure from the standard
Dipnoan type of dentition as established in Dipterus and carried on in
Sagenodus and Ceratodus. A perfectly distinct species, however, is the

196 PROF. D. M.S. WATSON AND MR. E. L. GILL ON THE
extremely interesting early form which we, following Traquair’s usage, haye
referred to as ‘“ Ctenodus interruptus, Barkas.” (If Dr. Smith Woodward’s
diagnosis of this species, ‘Catalogue of Fossil Fishes,’ part ii. p. 254,
accurately represents Barkas’s intention in founding the species, the name-
cannot properly be applied to the specimens in the Royal Scottish Museum
which Traquair referred toit.) The Sagenodus-like teeth of this fish (fig. 24,
p- 193) are in the strongest contrast with those of the latest known species,
C. murchisoni, Ward, from the Upper Coal Measures, which haye about

twenty practically parallel ridges.

Fie. 27.

Ctenodus. Lower jaw, xX +.

A. Reconstruction of lower jaw, from buccal aspect.
B, Ventral aspect of the same.
C. Left ramus, outer aspect.
D. inner aspect.
E, An angular and dentary in natural articulation. From two specimens,
in the Atthey Collection and the British Museum respectively,
. The same, inner surface, From the Atthey specimen.
(Compare figs. 11 and 12, pp. 178, 179.)

by

Pritsch’s Ctenodus tardus from the Brandschiefer (Lower Permian) of
Bohemia, represented in his Taf. 80b, is apparently a Dipnoan ; but if his
drawing of the bones of the skull-roof is at all accurate, it was certainly
neither a Ctenodus nor a Sagenodus.

SL(RUCTURE OF CERTAIN PALAOZOIC DIPNOI. U7

Ctenodus, Sagenodus, and Ceratodus.

' The close correspondence which has been traced in the structure of these
three genera, notably as regards the palate, the lower jaw and the shoulder-
girdle, leaves no doubt as to their near affinity. The structure of the skull-
roof, if followed through the same three fishes, shows less constancy ; on the

Fre. 28)

Ctenodus? A. Cleithrum. B, C. Clavicle, upper and lower surfaces.
Both bones are represented by numerous specimens in the Atthey Collection.

other hand, it shows an interesting progressive change, starting from the
pattern first established in Dipterus platycephalus. The change consists in
the expansion, at the expense of the surrounding bones, of the” median
“parietal” and interfrontal, together with the reduction of the bones of the
sngut region (nasals and prenasals). From Dipterus to Ctenodus the change,
though obvious, is not great. From Ctenodus to Sagenodus it is in both
respects very marked ; in Ceratodus it has been carried to an extreme, for

Fre. 29 A.
Den.

Parnoe L.Crer.

Conchopoma gadiformis, Kner. Drawn from gelatine casts from the two halves of a
specimen in the Royal Scottish Museum, x 1.

A. Dorsal surface of head. ANG., angular; C.Hy., ceratohyal ; Drn., dentary; Fr. + L.T.,
frontal + intertemporal; I.I'r., interfrontal; L.Cxnrr., left cleithrum; Na., nasal ;
Op., operculum; Panr., parietal 4- dermo-supraoccipital; Par.Sp., parasphenoid ;
Pr.FR., prefrontal; Pr., pterygoid; R.Cxie1, right cleithrum; R.P.Arr., right
pre-articular ; S.Op., sub-operculum; Sq., ‘“squamosal”; Tas., tabular + supra-
temporal.

B, Ventral surface of head, Reference letters as before, with L.Pr., left pterygoid ;
L.Op., left operculum; R.Crav., right clavicle; R.Op., right operculum; R Pr.,
right pterygoid. The mass of bone between Panr.Sp. and L.Cuct. is the exoccipital.

STRUCTURE OF CERTAIN PALASOZOIC DIPNOI. 199

the bones of the snout region have almost or quite disappeared, and the
“parietal” and interfrontal cover the whole median tract of the top of the
head.

The compression of the lateral bones of the skull-roof brought about by
this expansion of two of the median bones appears to have resulted in corre-
sponding degrees of disappearance or fusion. Thus in Sagenodus there is
only one pair of bones in the frontal region as compared with two in
Ctenodus and three or more in Dipterus platycephalus. Ceratodus is extreme
in this respect as in the others: all the bones of the frontal, temporal, and
tabular regions on each side seem to be represented as a rule by a single
ossification.

Conchopoma, another Dipnoan appearing late in order of time and de-
scribed below, furnishes an interesting parallel to Ceratodus. Though none
of the processes have gone as far as in Ceratodus forsteri, yet the two median
bones are greatly expanded, the bones of the snout much reduced, and
those of the lateral part of the roof extensively fused.

Several of the minor characters of the bones of the head in Ceratodus
as compared with those of Sagenodus are plainly correlated with one another.
The side-to-side arching of the skull-roof, bringing the squamosal region
far down on the side of the head, is connected with the reduction in size of
the operculum, and also with the shortening of the quadrate.

CoNCHOPOMA GADIFORMIS, Kner.

The rare fish from the Lebach Shales (uppermost Carboniferous) of
Saarbruck, called Conchopoma gadifomis by Kner, has never been at all
adequately described. There is in Edinburgh a very large individual, pre-
served in an ironstone nodule, which makes the structure very nearly com-
pletely known. It was prepared by softening the already rotted bones with
dilute acid and removing the residue by brushing. Casts from the moulds so
left show all surface details extremely well.

The general morphology will be vbvious from fig. 29, p. 198. The
skull has the usual Dipnoan structure of an extensive cartilaginous neural
cranium, which seems to have been considerably ossified in the exoccipital
region. The head is roofed by a continuous shield of membrane bones,
which, although now flat, seems from its cracked condition to have been
originally considerably curved. This shield consists posteriorly of a row of
three bones, of which the median ‘ parietal” is longer than the lateral
tabular. The “ parietal” has a low median ridge on its visceral surface ; the
“tabulars” are concave ventrally and now no signs of attachments.

The “ parietal” articulates with a median interfrontal, and these two bones
have long lateral attachments to the very large bones which include the

200 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

frontals ; these bones articulate with the ‘ tabular” behind, and seem just:
to reach the free anterior edge of the shield.

The lateral edge of the “tabular” and “ frontal”’ is attached to a series of
bones, the posterior two ? of which are narrow and form that margin of the
skull to which the operculum is attached; they are imperfectly shown from:
the dorsal surface on the right, and are absent on the left side. Immedi-
ately in advance of these is a very large element forming an outstanding
wing behind the orbit. This extends forward and just touches a much
smaller bone, which forms the anterior and upper quadrant of the orbital
margin.

The dorsal shield is completed by a pair of bones which articulate with the-
front margins of the frontals and interfrontal.

There are no traces of other cireumorbitals or of premaxille.

The palate presents an interesting modification of the Dipnoan type. The
parasphenoid has a long, slender dorsally channelled stem, which expands.
into a broad, shovel-shaped, flat bony plate extending forward as far as the
symphysis of the lower jaw. This region of the parasphenoid is covered with
teeth, small and very closely packed anteriorly, somewhat larger and more:
scattered posteriorly. ‘These teeth are quite irregular in arrangement, but
little groups of two or three of them are often supported on a common raised
base ; anteriorly where the teeth become inconspicuous the bases may persist
as short curved ridges.

The pterygoid on the left side seems to retain its natural position, standing
up nearly at right-angles to the palate.

The bone is very narrow anteriorly, forming a border to the great para-
sphenoid and having irregularly arranged small teeth. Posteriorly the bone
becomes converted into a deep thin flange, which was formerly applied to
the quadrate and stands nearly at right-angles to the palate.

The operculum has long been known as a concavo-conyex bone with an
umbo at its antero-dorsal corner. This specimen shows a small bone on the
right side which seems to be a sub-operculum, because of its resemblance to
that bone in Dipterus and Sagenodus.

The structure of the lower jaw is not satisfactorily shown. ‘There is a
powerful symphysis formed by bones of the outer surface homologous with
the “dentaries” of Sagenodus, but possibly including true dentaries in
addition.

There is a large angular of whose structure nothing can be said. Although
this lower jaw lies in its natural position and the mouth is closed, no trace of
the bones usually called splenial can be seen in it. There is, however, a
strange tooth-bearing bone with a concave visceral surface lying displaced in
contact with the right ramus, which apparently can only be this element. It
differs from the corresponding bone in all other Dipnoi in lacking a posterior
flange passing back to the articular and an anterior symphysis. The two

STRUCTURE OF CERTAIN PALAOZOIC DIPNOI. 201

ceratohyals are well shown; they exactly resemble the corresponding bones.
in Ceratodus.

The shoulder-girdle is remarkable for the very small relative size of the
clavicle.

The cleithrum is a large bone of much greater width than is shown in the
figures, where part of it is hidden by matrix in each case. The upper end is
comparatively narrow, and is seen to be covered by a very badly preserved
bone, no doubt the supra-cleithrum. The inner surface is concave, the outer
flat with a turned-in anterior margin; it has a depressed area for the
hinder edge of the opereulum. The lower end of the cleithrum turns
forward and is largely covered by the sub-opereulum, appearing behind that
bone only as a narrow strip.

The clavicle is displaced and shown only from its outer surface. It is
extremely short, attached to the cleithrum by a deeply recessed triangular
area on its outer surface, and widening both ways ventrally.

The neural arches are thoroughly ossitied anteriorly, although there are no
ossifications in the notochordal region. The anterior neural spines are
separately ossified from their arches. The ribs are slender, well curved and
with slightly thickened heads,

The structure of the median fins is already familiar; there is a continuous
fin agreeing exactly in structure and distribution with that of Ceratodus.

The pectoral fins are shown by the Edinburgh specimen to be large, typical
biserial archipterygia, the axis being entirely unossified, whilst the radials of
both series have short, hollow ossifications. There are very well-developed
camptotrichia which do not extend in to overlap the ossified radials.

The structure of the pelvic fins is not so clearly shown, but they obviously
agree in general with the pectoral fins, and are nearly as large.

The scales are not very well shown; they are thin, of considerable size, and
marked with very delicate concentric rings of growth. It is impossible to
say how far forward they extended.

Conchopoma must have been extremely like Ceratodus in proportions and
general build, although the head may have been slightly lower posteriorly.

URONEMUS SPLENDENS (Traq.).

The genus Uronemus was founded by Agassiz for some small fish from
the Burdiehouse Limestone, which have a continuous median fin and an
apparently diphycercal tail. ‘To this genus Dr. Traquair referred some
remains from the Lower Carboniferous, No. 2 Ironstone, Loanhead, Mid-
lothian, which show many details of skull structure. Beyond stating that
these specimens show a skull-roof like that of Ctenodus and prove that the
pterygoids and “splenials” bear a single series of large, compressed, low,
conical teeth in addition to a granulation of small denticles, there being no

202 PROF. D. M. S. WATSON AND MR. &. L. GILL ON THE

dental plates of a normal Dipnoan structure, he gave no further account of
them, and never published any figures in illustration of his description. In
1891 Dr. Smith Woodward gave a figure of an isolated “‘splenial,” since
when no further details have been published.

The types are in the Royal Scottish Museum, and between them make the
structure nearly completely known.

Fig. 380A. Fia. 30B.

Uronemus splendens, Traq. Drawn from plasticene squeezes of a specimen in the
Royal Scottish Museum. X 1.

A. Dorsal surface of head. Fr., frontal; I.FrR., interfrontal; I.TEm., intertemporal ;
L.ANG., left angular; L.Duwn., left dentary; L.P.Anr., left pre-articular; L.Pr.,
left pterygoid ; Mx. ?, maxilla; Na., nasal; P.O., post-orbital; P.V., prevomer ;
Pr.FR., prefrontal; P.Mx.?, premaxilla; Panr., parietal; R.ANG., right angular ;
R:Den,, right dentary ; R.P.Art., right pre-articular ; R.Pr., right pterygoid.

B. Ventral surface of head. Reference letters as before, with C.Hy., ceratohyal; CLE1.,
cleithrum ; Op., operculum; Par.Sp., parasphenoid.

The best specimen is a nearly complete small fish, preserved mainly as an

impression in slab and counter slab.
I have prepared it by removing the scanty remains of bone from the
impressions, and studied it in plasticene squeezes, which reproduce all surface

STRUCTURE OF CERTAIN PALMOZOIC DIPNOI. 203

details with great perfection. Other specimens show the cranial roof from
the visceral surface.

The cranial roof consists of a series of bones arranged very nearly as in
Ctenodus, but the three nearly complete examples in Edinburgh differ in-
details. There is a posterior row of three bones, a median “ parietal,”
which supports a pair of frontals, and lateral tabulars which are shorter than
the “parietal” and are continued forward by intertemporals.

The frontals are separated anteriorly by a small interfrontal, but support
a pair of nasals in two specimens and only a single median nasal in the
skeleton. Finally, in this specimen the skull-roof is completed by a small
median prenasal with a notched anterior border. =

Fre, 31.

Uronemus splendens, Traq. Roof of the skull, seen from the ventral surface, x 2.

Fr., frontal; I.Fr., interfrontal; I.TEm., intertemporal; Na., nasal; P.O., post-orbital ;
Par., parietal; Sq., “squamosal’’; Tas., “ tabular.”

The intertemporal and tabular articulate with a large squamosal, there
being no evidence of the presence of the two small elements which in most
Dipnoi appear on the margin of the shield bordering the tabular and
squamosal.

The intertemporal is continued forward by a large post-frontal, which
itself supported another element, which although now lost in every specimen
cannot have entered the orbital margin.

Four cireumorbital bones are preserved in the skeleton ; probably one more
was originally present. The three specimens which show that region differ
considerably in the details of these circumorbitals, which articulate with the
squamosal and post-frontal. All the bones of the skull-roof bear a crisply-
marked fine ornament of ridges and pits, which in general radiate from the

204 PROF. D. M. 8. WATSON AND MR. E. L. GILL ON THE

centre of the bone. This ornament much resembles that of Stegocephalian
skulls, and is unique amongst Dipnoi.

The palate is beautifully shown in the skeleton. The parasphenoid has a
wide but incompletely preserved stem, which expands into an elongated but
relatively very narrow lozenge, with the margin of which the pterygoids
articulate.

The pterygoids cover a very large area, meeting one another in a long
symphysis in front of the parasphenoid and passing out to the lateral margins,
which are quite straight and make an acute angle with one another. The
hinder end of the pterygoid is widened, is attached to the parasphenoid by a
very long suture, and produced into a rounded corner in the quadrate region.

The dentition of each pterygoid consists of a marginal row of large,
compressed, conical teeth and of a very large number of small, almost
hemispherical denticles covering a narrow area within the margin. These
denticles are rather regularly arranged in a series of straight lines, running
parallel to one another from caudal and lateral to cranial and mesial ;
that is, they do not agree with the normal direction of teeth-rows seen in the
pterygoids of Mipterus and Sagenodus.

The two prevomers of the skeleton are preserved ; each consists of a bone
with a cylindrical notch on the dorsal surface, which supports a row of four
denticles exactly similar to those of the “ splenial.”

There are in the skeleton three small bones, each bearing small, elongated
teeth ; two of them lie in close association with a prevomer: it is certain
that they are not lower jaw elements, and the conclusion seems irresistible
that they are premaxille and maxille corresponding to those found by
Watson and Day in Phaneropleuron.

The two rami of the lower jaw are separable at the symphysis, and each
-consists of at least three, almost certainly of four, bones.

The angular is a large bone exactly like that of Ctenodus in shape and
‘bearing a similar row of sensory pits. It articulates by a fine suture
with a splenial (dentary of Ceratodus), which presumably extended to the
symphysis.

There is a rather large dentary, whose distinctness from the splenial
cannot be proved, which bears an irregular strip of crowded small denticles.
The prearticular (splenial of Ceratodus) extends from the articular region,
toward and probably up to the symphysis ; it bears only a sinvle series of
teeth on its upper edge, which bite outside and are exactly similar to the
marginal teeth of the pterygoid.

The opercular apparatus seems to consist only of a large oval operculum,
agreeing closely in shape with that of Sagenodus.

There are a pair of badly-preserved ceratohyals, agreeing closely with
those of Ceratodus.

STRUCTURE OF CERTAIN PALHOZOIC DIPNOI. 205

DIPTERUS VALENCIENNESI, Sedg. & Murch.

Dipterus valenciennesi is a small fish, found very abundantly in the western
end of the mainland of Orkney and at Banniskirk, Achanarras, and other
localities in Caithness, and, except at Hdderton Burn, only very rarely in the
nodules of the Moray Firth.

It was distineuished from the more recent D). platycephalus by Pander on
account of its smaller size, lack of shiny surface on the cranial bones and
scales, and lack of well-ossified anterior ends of the skull and mandible.
Dr. Traquair, presumably because large specimens which in these characters
agree with D. platycephalus do occur at Achanarras, Gamrie, and Cromarty
and in association with a small fish which he regarded as D. valenciennesi at

Eire. 32:

Dipterus valenciennesi, Sedg. & Murch. Right lateral aspect of a crushed head. x 2.
From a specimen in D. M.S. Watson’s collection.
Cuav., clavicle; Curt. cleithrum; L.G. 3-6, lateral gulars, nos. 3-6; Mp., mandible ;
Mx., maxilla; Op., operculum; P.G., principal gular; P.TEM., post-temporal ;
8.Cx., supra-cleithrum ; 8.Op., sub-operculum.

the Gloup in Orkney, combined the two species. Watson and Day pointed
out a distinguislring feature in the fusion in D. platycephalus of the tabulars
and interparietal with the bones immediately in front of them, which remain
separate in D. valenciennesi.

Amongst the hundreds of specimens of D. valenciennesi which we have
seen, only three show the bones of the cranial roof clearly ; few show anything
of the gular apparatus, atid only one shows the lateral surface of the head
clearly.

206 PROF. D. M.S. WATSON AND MR. E. L. GILL ON THE

Our existing knowledge of the structure of the head still depends in the
main on Pander’s excellent description and figures. The only addition is
Dr. Traquair’s well-known restoration.

One specimen (fig. 32), now in D. M.S. Watson’s collection, shows the
side of the head nearly perfectly. Its structure will be best understood
from the figure; the interesting features are the relatively small- number
(seven) of circumorbitals, the presence of two small elements in the cheek,
representing some part of the greatly-developed bones in this region in
Osteclepids, and the presence of a very slender, toothless maxilla, lying below
the orbit.

Cray.

Cues.
Dipterus valenciennesi, Sedg. & Murch. Ventral surface of head. x 2.
From a specimen in D, M.S. Watson’s collection,
Ana., angular; Cuav., clavicle; Crier, cleithrum; L.G. 1-8, lateral gulars, nos. 1-3;
M.G., median gular; Op., operculum; Po.Sp., post-splenial=preangular; P.G.,
principal gular; S.Op., sub-operculum; Sp., splenial.

There is an indication of an inner ring of circumorbitals, already repre-
sented in Dr. Traquair’s restoration.

This specimen shows the operculum and a section across the suboperculum,
that bone having been driven outward and its upper half thereby removed
in the counter slab when crushed down on the rigid clavicular arch. The
sub-operculum is followed by a large principal gular, which supports a large.
lateral gular.

In a triangular space between these two gulars and the articular region of
the lower jaw lie three small bones, which can only be lateral gulars. These.
little bones are also shown in identically the same form in the specimen
No. 770, Hugh Miller Collection, Royal Scottish Museum.

The gular apparatus is, however, best seen in-the original of fig. 33,
a small specimen crushed directly vertically and viewed from the ventral

STRUCTURE OF CERTAIN PALZ@OZOIC DIPNOI. 207

surface. Here the space between the rami of the lower jaw is mainly occupied
by the two pairs of large gular plates already described; but in advance of
these are three other small bones, two of which are obviously a pair meeting
in the middle Jine, whilst the other more lateral element which on the right
side separates them from the anterior end of the large gular is concealed on
the left side by a forward displacement of that bone. At the spot where the
four large gulars meet is a small median lozenge-shaped element, already
figured, as were the large gulars, by Pander.

No specimens known to us show the structure of the shoulder-girdle
completely.

There are a scale-like post-temporal and supra-cleithrum connecting the
upper end of the cleithrum with the tabular. The cleithrum is an elongated,
narrow bone with a recess on its outer surface for the reception of the hinder
edge of the operculum. It is so rigidly attached to the clavicle, that that
bone usually retains its natural position and shape in the fossils, having
resisted the crushing which disarticulates most of the other bones.

There is some evidence (Peach Coll., No. 35, Royal Scottish Museum) that
this attachment is effected by a special downwardly-projecting process on the
inner surface of the cleithrum, which is received in a recess in the clavicle :
that, in fact, the structure here is exactly as in Sageriodus.

The clavicle is a massive bone turning inward and forward onto the ventral
surface, and with its fellow filling up the triangular space between the
principal gulars.

There is some evidence of a large scale in the position of an interclayicle,

DIPTERUS PLATYCEPHALUS, Ag.

We are unable to add much to the existing knowledge of the skull of
D. platycephalus. No specimen known to us shows the circumorbitals in
intelligible form, and none gives an altogether satisfactory view of the
opercular region. No. 1059, Hugh Miller Collection, Edinburgh, and
Pander’s figures, Taf. 3, fig. 17, and Taf. 4, fig. 26, show only three bones
on each side—an operculum, sub-operculum, anda gular; no other specimens
show additional elements, and it is thus possible that the apparatus was far
more reduced than in D. valenciennesi, although the material does not admit
of definite statements.

The structure of the lower jaw (cf. fig. 34, p. 208) is perfectly shown by
No. L.10858 of the Manchester Museum. The general features of the
morphology were accurately figured by Traquair, but that author was not
acquainted with certain very important characters, vividly shown in our

specimen.

The dentaries are small elements forming a rim to the anterior end of the
mandible ; the two bones are indistinguishably fused. In section their outer
surface forms nearly three quadrants of a circle passing smoouhly from the

LINN. JOURN.—ZOOLOGY, VOL. XXXV. _ 14

208 PROF. D. M. S. WATSON AND MR. E. L. GILL ON THE

lower upward into the oral surface ; this surface is covered with the shiny
punctate surface now recognized as characteristic of a cosmoid bone. This
region projects forward, and is sharply marked off from the rest of the lower

jaw by depressions on the nearly vertical lateral surfaces.

The dentary articulates with three other bones which lie on the outer
surface of the jaw ; these are, a very small splenial lying anteriorly on the
flat under surface of the chin, a somewhat larger post-splenial which forms
part of the floor of the lateral depression, in addition to a large part of
the ventral surface, and a very large angular which extends backward to
articulate with the articular.

Fie. 34,

Art Fr. Art.

Den.

Po.Sp, Ang. ; Ang. FoSp. Op.
Dipterus platycephalus (Ag.). Dorsal and ventral views of the lower jaw, with inner and
outer aspects of the left ramus. x14. Restored from No. L. 10858, Manchester Museum.

ANG., angular; Arv., articular; Drn., dentary; Po.Sp., post-splenial = preangular ;
Pr.ARrv., pre-articular; Sp., splenial.

The presence of splenial, post-splenial, and angular elements is clearly
confirmed by No. 1878.5.166, Royal Scottish Museum, where these bones
are seen from the dorsal aspect, the tooth-bearing bones being removed.

The articular is a very large bone, with a well-defined condylar surface
formed by a deep cylindrical excavation running across its hinder surface

STRUCTURE OF CERTAIN PALMOZOIC DIPNOT. 209

and passing very obliquely backward. The bone is continued forward for a
considerable but uncertain distance in contact with the inner surface of the
pre-articular.'

The pre-articular is the bone usually called splenial in Ceratodus. In
Dipterus platycephalus it is of “great length, extending from the articular
facet to a symphysis with its fellow, which extends forward nearly to the
anterior end of the jaw.

Tts hinder part is a vertically placed thin sheet of bone, tightly applied to
the inner surface of the articular, with a free upper margin which forms the
border of the supra-Meckelian vacuity, and with its lower border in contact
with a depressed strip of bone which appears to be a part of the angular, but
is perhaps, as Dr. Stensio has suggested to us, really the ossified Meckel’s
cartilage. It is possible that in the naturally articulated jaw the strip was
completely covered by the pre-articular, and that in consequence the ramus is
represented as of too great a depth in fig. 34.

From a point not far in front of the articular to the hinder end of the
dentary the upper edge of the pre-articular is turned outward and the bone
thickened ; anteriorly this edge meets the angular, forming the border of the
very large supra-Meckelian vacuity.

This out-turned edge of the pre-articular bears the tooth-plate, which is a
thick pad with denticulated ridges radiating from a point on the inner border
rather behind the middle. The pattern of these teeth-plates differs consider-
ably in individuals ; they commonly extend much further forward than in
fig. 34.

The pre-articular is completed by turning inward as a flat, nearly horizontal
plate to the symphysis, which lies on the dorsal surface of the splenials.

In the reconstructions represented in fig. 34 the ramus may be made too
deep and wide. The depth illustrated depends on the actual depth in the
specimen ; the width is fixed by that of the hinder end of the dentary and of
the tooth-bearing part of the pre-articular. At the most, the inaccuracy can
only be of the order of one or two millimetres.

The course of the lateral-line canal on the lower jaw is well shown in the
specimen. Along the lower border of the angular it is represented by an
irregular and often double series of very small pits.

On the post-splenial are two definite lines of large pits, one passing
forward parallel to and just within the lower margin of the lateral depression,
the other along the hinder border of the bone; in addition, there are other
seattered pits. The splenial has two pits near and parallel to its hinder
margin:

The new information about Dipterus recorded above adds very con-
siderably to the evidence in support of the view first definitely stated by
L. Dollo that Dipterus is by far the most pa e, as it is the oldest
known Dipnoan.

210 PROF. D. M. §. WATSON AND MR. E. L. GILL ON THE

Dipterus is now shown to resemble the Osteolepids in the following
ways :—
1. In the typical cosmoid structure of its scales.
2. In the general body form, possession of two separate dorsal and an
anal fin, and a heterocercal tail with an epichordal lobe.
3. In the number and relations of the dermal bones of the top of the
head.
4. In the presence of splenial and post-splenial bones in the lower jaw.
5. In the possession of a very elaborate opercular apparatus, including
opercular and sub-opercular, a series of lateral gulars, a pair of
principal gulars, and a median gular.
6. In the structure of its paired fins.
These resemblances are of .a fundamental character, and imply a community
of origin of the two groups.
oe principal features in which Dipterus differs from an Osteolepid are :—
. The loss of the hyomandibular as a bone playing a part in the support
of the quadrate.
2. The suppression of the suprapterygoid ossicles and the ‘oe of
a symphysis between the pterygoids anteriorly, and of a sutural
attachment of the pterygoids to the edges of the parasphenoid.
3. The loss of the palatine and ectopterygoids, and the development of a
“‘tooth-plate ~ on the pterygoid.
4. The loss of the coronoids and the development of the tooth-plate on
the pre-articular.
5. The great reduction of the marginal tooth-bearing bones, the pre-
maxillee, maxillee, and dentaries.
6. The forward inclination of the quadrate, with a correlated shortening
of the lower jaw and a great reduction of the cheek-plates.
7. The removal of the external nares from a point on the upper surface
to a position below the lip.
8. The absence of a movable joint between the parietals and frontals,
and of an unossified region of the basis cranii between the basi-
occipital and basisphenoid.

Numbers 1-6 of these differences are obviously dependent on different
food habits, the Dipnoi having adopted a diet which required trituration,
whilst the Osteolepids are predaceous, swallowing their prey entire.

Difference number 7 may perhaps have arisen not:as an adaptation, but as
a mere result of the mode of development of the olfactory organ in a fish
which has a much reduced maxilla.

The differences recorded under No. 8 ean only be explained by supposing
these peculiar characters to have been acquired by the Osteolepids after their
separation from the Dipnoi.

STRUCTURE OF CERTAIN PALMOZOIC DIPNOT. PALL

Thus we are driven to conclude that the Dipnoi, although they have
undoubtedly been derived from a stock whose general morphology was
Osteolepid, were separated from that stock before its members acquired their
most peculiar and diagnostic features.

Comparison of the head of Dipnoi with that of an Osteolepid, the homo-
logies of whose bones are obvious in the light of the structure of Palzeozoic
amphibia, throws great light on the determination of the elements of the
skull.

The very: interesting observations recorded by Prof. Goodrich in his
general work on Fishes (1909) were the first correctly to lay down the main
lines of the comparison between the skulls of Dipnoi and Osteolepids,
especially in their recognition of the distribution of the lateral-line canals on
the top of the head.

Gregory (1915) further extended this comparison, and finally a comparison
of the skull-roof of D. valenciennesi with that of the Osteolepids led
Watson and Day to an identification of nearly all the bones which compose
its apparently inexplicable mosaic.

This interpretation depends on the belief that in J. platycephalus the
tabulars and post-parietal have fused with the parietals and supratemporals.
The lack of a specimen of this species with the skull in articulation with the
trunk renders the evidence incomplete, but the fact that the posterior row of
three bones in all later Dipnoi houses the occipital cross-commissure of the
lateral line shows that it includes these bones; and the well-known pair
of grooves on tha median occipital, which are for a line of pit ergans and are
obviously homologous with a similar pair of grooves on the parietals of
Osteolepids and some other fish, show that this bone includes the parietals. _

In the lower jaw of Dipterus the dentary, angular and articular are readily
identifiable, and the relations of the splenial and post-splenial to the dentary
and angular are so exactly those which obtain in Osteolepids and indeed in
Labyrinthodont amphibia as to leave no doubt of their homologies The
only remaining bone, that which bears the tooth-plate, cannot be the splenial,
as has been previously believed, because an undoubted splenial occurs in the
same Jaw. It can only be pre-articular or coronoid: the application of its
hinder end to the inner surface of the articular, its large size and position on
the inner surface of the ramus, show that it is a pre-articular, and the
symphysis which it makes with its fellow can be matched in the case of
the pre-articular of Megalichthys.

In all Osteolepids and Labyrinthodonts the teeth-bearing bones of the upper
and lower jaws have certain quite definite relations to one another. The
dentary bites within the maxilla, the teeth on the coronoids interlock with
those on the palatine and ectopterygoid, and the upper edge of the pre-
articular, which is usually covered with a granulation of small denticles, faces
although it does not touch the similarly armed pterygoid. If, as seems to

212 PROF. D. M. 8. WATSON AND MR. E. L. GILL ON THE

be clearly the case, the tooth-plate on the pre-articular of Dupterus be a
development of this customary granulation, then the similar upper jaw
structure on which it grinds must have been derived from the similar
granulation on the pterygoid. It is improbable that this large tooth-bearing
bone on the palate can be a conjoint pterygoid and palatine as is usually
believed, because the palatine teeth of Osteolepids are always long tusks,
unlikely in any case to be worked up into so typical a crushing structure as
the Dipnoan dental plate, and were the palatine preserved, we should
expect to find that the lower plate was of coronoid derivation. Nothing in
the structure of the Dipterus lower jaw supports this view, which can, how-
ever, never be disproved because of the possibility (although we think
general great improbability) of a fusion of bones.

The opereular apparatus of D. valenciennesi is readily interpretable in
comparison with that of an Osteolepid. Its unusual feature, the direct,
contact between the principal gular and the sub-opercular, is probably due to
the great reduction of the hinder lateral gulars which is associated with the
shortening of the jaw. The meeting of the two rows of lateral gulars in
the front is paralleled in Megalichthys (No. 28308, Museum of Practical
Geology ).

The problem of the relationships of the known Dipnoan genera to one
another is still incapable of satisfactory solution, because of paucity of material.

The position of Dipterus valenciennesi, at the base of the series, seems to be
made certain not only by the fact that it is actually the oldest known species,
but by the close comparison which can be drawn between its structure and
that of an Osteolepid.

From this form the series D. platycephalus, Pentlandia, Scaumenacia,
Phaneropleuron, first suggested by Prof. Dollo and supported from new
evidence by Watson and Day, seem to haye arisen. These forms first
appear in time in the order named, and show a steady progressive change,
resulting in a reduction of the dermal bones in the front of the head, a
reduction of the ossifications in the chondrocranium, a loss of the inter-
frontal and internasal elements, and a gradual fusion of the median fins
with one another. All known features of their-structure are consistent with
direct descent of any form from that which precedes it.

The new facts about the structure of Sagenodus and Ctenodus brought
forward in this paper show that Watson and Day were not justified in
separating them widely from one another and in deriving Sagenodus from
Phaneropleuron.

The descriptions we have given show that the two animals are closely
related, that indeed the Lower Carboniferous “* C. interruptus” is in its teeth
in many respects intermediate between the Coal-Measure forms of the two
genera. Ctenodus has the less reduced skull-roof, which by its retention of

STRUCTURE OF CERTAIN PALHOZOIC DIPNOT. 213

interfrontal and internasal bones demands descent from a form not later than
D. platycephalus, and cannot have arisen from the much more reduced skulls
of Pentlandia, Scaumenacia, and Phaneropleuron.

The Sagenodus skull-roof can readily be derived from that of Ctenodus.
In its dentition Sagenodus is much more primitive than Ctenodus in retaining
the Dipterine arrangement of radiating ridges on the tooth-plates ; in fact,
Sagenodus and Ctenodus seem to afford an illustration of the fact, of not
uncommon occurrence and great theoretical interest, that if one of two allied
closely-related forms is more specialized in a certain region than the other,
it will be less advanced in the structure of some other region.

The comparison between the structures of Sagenodus and Ceratodus
included in the description of the former genus seems to us to establish the
descent of the latter from the former animal.

The Dipnoi from the Old Red Sandstone probably lived in unusual
conditions. J. Barrell has brought forward a mass of evidence to show that,
in common with the other fish of the Old Red Sandstone, they were
inhabitants of an arid region with seasonal rainfall, living in rivers which
were liable to dry up during part of the year and in shallow and
impersistent lakes. Although we believe that this view cannot be upheld
in its entirety—for it is difficult to conceive of the Caithness flags being
deposited “anywhere except in a very extensive and permanent sheet of
water—it is undoubtedly well founded in its general conclusions.

The Coal-Measure Dipnoi lived under entirely different conditions in pools,
often, as in the case of that in which the reof of the Low Main Seam at
Newsham was deposited, of very considerable size and permanence. These
pools seem to have lain in the midst of the coal-producing forests in a
elimate which was in no way arid. Thus this difference in habitat at once
affords an explanation of the absence of any direct descendant of the
Phaneropleuron line and the occurrence of a stock not known from the
Upper Devonian.

There remain for consideration the remarkable Uronemus and Conchopoma.

These animals have been associated with one another, though only very
doubtfully, by Traquair and Smith Woodward, because of the replacement
in them of typical dental plates by isolated small denticles.

Comparison of the figures and description given in this paper will show
that there are no valid reasons for believing in the close affinity—that, in
fact, they differ so greatly as to be in all probability merely functionally
parallel modifications of very different stocks. Uronemus has a skull-roof
retaining interfrontal and internasal bones, but much reduced by the loss of
the bones on the lateral edge of the temporal region behind the squamosal,
and of others forming the roof of the skull. Conchopoma, with a much more
reduced skull-roof, loses the internasal, has a much enlarged interfrontal,
exhibits a fusion of the frontal, intertemporal, and post-frontal, and retains

214 PROF. D. M. S. WATSON AND MR. E£. L. GILL ON THE
s

the bone behind the squamosal. So far as the skull-roof is concerned, Uro-
nemus might be derived from Ctenodus, Conchopoma from Sagenodus.

In the palate the two genera differ very widely. Uronemus has a small
parasphenoid, with a long posterior stem and a lozenge-shaped palatal part
completely devoid of teeth. The pterygoids form an exceptionally large
part of the palate, have a single series of large teeth along their outer margin,
and an area covered with small teeth which show no trace of arrangement in
radiating ridges. Conchopoma has a unique parasphenoid with a long, slender,
posterior stem, and an enormously enlarged palatal part extending far forward
and covered with an irregular development of small denticles. The ptery-
goids form narrow slips along the lateral borders of the parasphenoid, bear
only a few small teeth, and present no trace of the enlarged marginal teeth
of Uronemus.

Both forms are unexpectedly primitive in retaining dentaries in the lower
jaw, in addition to those splenials which have usually been called by that
name in Ceratodus,

It now falls to be considered whether all the ancestors of these fish had
only isolated denticles, or whether they have arisen separately by the breaking
up of dental plates.

Semon showed that in development the dentition of Ceratodus begins as a
series of isolated denticles supported by a net-work of bony spicules, and that
the tooth-plates of the adult are built up byithe confluence of such denticles.

This mode of development is consistent with the view that Uronemus may
have arisen from a form with tooth-plates, because its isolated denticles
may merely result from the carrying on to adult life of a structure which
occurred in larval stages. It is probable that a stage with distinct unfused
denticles formed a larger part of the life-history in early Dipnoi than it-does
in Ceratodus ; indeed, the small plate figured by Pander as D. tuberculutus
(Taf. 5, figs. 20-21) seems actually to consist of individual denticles placed
on a bony base.

The whole structure of the Dipnoan skull, the short mouth and forwardly-
directed quadrate, the rigid attachment of the pterygoids to the basis cranii,
and the reduction of the hyomandibular, all point to the Dipnoan stock being
specially modified for the use of a highly-developed crushing dentition, All
these changes can be paralleled in those other groups of fishes which have
developed analogous tooth-plates.

Thus we believe that a typical tooth-plate like that of Dipterus was probably
of very early introduction into Dipnoan structure, its production having in
fact gone on pari passu with the other correlated changes in the head.

We are thus led to believe that Uronemus and Conchopoma are derived
from fish which had tooth-plates, and as no such fish either has or needs a
parasphenoid dentition, that the well-developed dentieles on that bone in the
latter fish are new developments.

STRUOTURE OF CERTAIN PALAJOZOIC DIPNOI. ~ 215

SUMMARY.

This paper includes the first complete accounts of the cireumorbital region
and opercular apparatus of Dipterus valenciennesi, and of the lower jaw of
D, platycephalus. It gives very nearly complete descriptions of the skull,
Jower jaw, and clavicular apparatus of Sagenodus and Ctenodus which much
extend our knowledge of these fish. In it the structures of the anterior end
of the rare fish Uronemus and Conchopoma are described and figured for the
first time.

Dipterus is shown to be directly comparable with Osteolepids in the
structure of the opercular apparatus and the lower jaw, in addition to the
many previously known resemblances. It is thereby shown that, as its early
date would indicate, it is the most primitive known Dipnoan.

Ctenodus and Sayenodus prove to be closely allied, and a detailed com-
parison shows so great a similarity between the latter fish and Ceratodus as
to leave no doubt that it is essentially ancestral to it.

Uronemus and Conchopoma resemble one another only in the reduction in
them of the tooth-plates to isolated denticles. In the structure of the palate
and of the reof of the head they differ so much that they must represent
widely-separated stocks.

In the main, the trends of Dipnoan development suggested by Watson and
Day are confirmed. It is, however, pointed out that the structure of the
neural cranium of the Osteolepids, as described by Bryant in ELusthenopteron,
is such that the Dipnoi cannot be direct descendants of that group, but that
with it and the Amphibia they arose together from common ancestors at a
time before the Middle Devonian.

_ We have to thank Dr. A. Smith Woodward, of the British Museum,

Dr. Kitchin, of the Museum of Practical Geology, Dr. Tattersall, of the
Manchester Museum, and especially Drs. Eagle Clarke and J. Ritchie, of the
Royal Scottish Museum, for the use of the materials in their care.

List oF PAPERS CITED.

1835. AG@assiz, L.— Recherches sur les Poissons Fossiles, vol. ii.

1844, —— Poissons Fossiles des Vieux Grés Rouge.

1898. Doxto, L.—Sur la phylogenie des Dipneustes. Bull. Soc. Belze de Géol., vol. ix.

1889. rirscu, A.—Fauna der Gaskohle und der Kalksteine der Permformation Boéhmens,
Bd. i. Heft 2, pp. 65-92, pls. 71-80.

1909, Goopricu, E. S.—Vertebrata craniata, in . R. Lankester’s ‘Treatise on Zovlogy.’

x London, 1909.

1915, Greacory, W. K.—Present status of the Problem of the origin of the Tetrapoda.
New York Acad. Sci. Annals, vol. xxvi.

1871. GuntHER, A.— Description of Ceratodus. Phil. Trans, 1871, p. 586.

LINN. JOURN.—ZOOLOGY, VOL. XXXY. 15

216

1868.

THE SEPRUOCTURE OF CERTAIN PALHOZOIC DIPNOL.

Hancock, A., & T. Arrury.—Notes on the various species of Ctenodus obtained
from the shales of the Northumberland Coalfield. Ann. & Mag. Nat.
Hist., vol. i. po. 77-87.

1869. —— Notes on the remains of some Reptiles and Fishes from the shales of the
Northumberland Coalfield. Ann. & Mag. Nat. Hist., vol. i. pp. 266-278,
346-578, 3 pls.

1872. —— A few Remarks on Dipterus and Ctencdus, and on their Relationship to
Ceratodus Forstert, Krefft. Ann. & Mag. Nat. Hist., vol. vii. pp. 190-
198.

1872. —— Descriptive notes etc. on a few Fish Remains found in the Coal Measures at
Newsham. Ann. & Mag. Nat. Hist., vol. ix. pp. 269-272.

1868, Knrr, R.—Ueber Conchopoma, nov. gen. et sp. aus dem Rothliegenden yon Lebach.
Sitz. d. k. Akad. d. Wiss. z. Wien, Bd. lvii. pp. 1-28, pls. 1-4.

188]. Mratn, L. C.—On Ctenodus. Proc. York. Geol. & Polytechnic Soc., vol. vii. p. 291.

1849. Mitter, H.—Footprints of the Creator,

1858. PAnpErR, C. H.—Ueber die Ctenodipterinen des Devonischen Systems. St. Peters-
burg, 18658.

1891. TritEr, F.—Ueber den Schidel eines fossilen Dipnoers, Ceratodus Sturti. Abh. d.
k. k. Geol, Reichsanst. Bd. xy. pp. 1-39, 4 pls.

1878. Traquarr, R. H.—On the genera Dipierus, etc. Ann. & Mag. Nat. Hist. ser. 5,
vol. ii, pp. 1-17,

188]. —— Notice of new Fish Remains from the Blackband Ironstone of Borough Lee,
near Edinburgh. Geol. Mag., Dee. 2, vol. viii. pp. 84-57.

1882. —— Notice of new Fish Rensains, etc. No. III. Geol. Mag., Dec. 2, vol. ix.
pp: 540-546.

1888. —— Notes on the Nomenclature of the Fishes of the Old Red Sandstone of Great
Britain. Geol. Mag., Dec. 3, vol. v. pp. 507-517.

1889. —— Ona new Species of Dipterus. Geol. Mag. Dec. 3, vol. vi. pp. 97-99, pl. ii.

1890. —— Notice of new and little-known Fish Remains from the Blackhand Ironstone of
Borough Lee, near Edinburgh. Geol. Mag., Dec. 3, vol. vii.

1896. —— ‘The extinct Vertebrata of the Moray Firth Area, in J. A. Harvie-Brown and
T. E. Buckley’s ‘ Vertebrate Fauna of the Moray Basin,’ vol. ii.

1900. —— Presidential Address to Section D. Rept. Brit. Assoc., Bradford Meeting,
pp: 768-783.

1916. Watson, D. M. S., & H. Day.—Notes on some Paleozoic Fishes. Manchester

Memoirs, vol. lx. pp. 1-48, 3 pls.

. Wituiston, S. W.—A new species of Sagenodus from the Kansas Coal Measures.

Kansas University Quarterly, 1899.

. Woopwarp, A. S.—Catalogue of Fossil Fishes in the British Museum (Natural

History), vol. ii.

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THE JOURNAL

OF

THE LINNKEAN SOCIETY,

SA sult
SS

ZOOLOGY.

CONTENTS.

I. On the Method of Oviposition and the Egg of Lyctus brunneus,
Steph. By A. M. Arson, F.E.S. (Communicated by Dr.
A. D. Ivns, F.L.8.) (Plate 12 and 2 Text-figures.) ............ 217

II. Some Echinoderms from West Australia. By Huserr Lyman
Ciark, Museum of Comparative Zoology, Cambridge, Mass.,
U.S.A. (Communicated by Prof. W. J. Daxiy, F.L.S.)
(Geletesl Ss) Wremcemaer sei ctage cise acs x nspqeecienGc nek ccaueturndce cincvle sess vat 229

III. Courtship Activities in the Red-throated Diver (Cotymbus
stellatus, Pontopp.) ; together with a discussion of the Evolution
of Courtship in Birds. By Jurian 8. Huxuiry, M.A. (Com-
municated by Prof. EH. S. Goopricn, M.A., F.R.S., Sec.L.8.)
(Plates 14, 15, and 4 Text-figures.)

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THE METHOD OF OVIPOSITION OF LYCLUS BRUNNEUS. 217

On the Method of Oviposition and the Ege of Lyctus brunneus, Steph. By’
A. M. Arson, F.E.S. (Communicated by Dr. A. D. Inms, F.L.S.)

(PuatE 12 and 2 Text-ficures.)

[Read 15th February, 1923. ]

CONTENTS.

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INTRODUCTION.

In April 1920, the writer commenced an investigation on Lyctws—Powder-
post—beetles attacking seasoned hardwoods, with a view to devising control
measures against their ravages.

This paper is a part of one originally written dealing with all those various
aspects of LZ. brunneus which had not been recorded, but owing to the cost
of getting the whole published, it has been found necessary to cut it up into.
sections, and this constitutes the first part.

There are two species of beetles of the genus Lyctus, family Lyctidee, so
far recorded in the British Isles. They are Lyctus (Xylotrogus) brunneus,
Steph., and Lyctus linearis, Goeze (canaliculatus, Fab., oblongus, Oliv.,
striatus, Melsh., unipunctatus, Herbst). Both species breed in dry or
seasoned hardwoods, usually attacking the sapwood only, although L. brunneus
was found attacking the sap- and heartwood of mahogany (Ahaya sp.), and
Noerdlinger (1862) recorded LZ. linearis as attacking sap- and heartwood of
black locust (Gleditschia triacanthos).

In 1891 Fowler recorded L. brunneus as very rare, and L. linearis (canali-
culatus) as common; the contrary is now the case. Hopkins (1911)
considers brunnews—now widely distributed throughout the world—to be of
S. American origin, and linearis of Huropean origin.

REVIEW oF THE LITERATURE.
Amongst the mass of literature on Powder-post beetles, there has been very
little published upon ZL. brunneus, and what there is deals mainly with its
LINN. JOURN.—ZOOLOGY, VOL. XXXY. 16

218 MR. A. M. ALTSON ON THE METHOD OF

ravages, but on the other Lyctus beetles there are some descriptions of
oviposition, and in three cases an egg is described.

In 1853 Heeger figured and dlasominad an egg, a legless larva, and a pupa,
which he noodle to cn pubescens, Panz.

In 1855 Noerdlinger described the larva, pupa, and adult of L. linearis
(canaliculatus),and in an account of the habits stated that the eggs were
deposited in cracks or fissures.

In 1874 Kaltenbach refers to the habits of the larva of linearis (canali-
culatus) and to that of pubescens, quoting Heeger as his authority in the
case of the latter.

In 1876 Perris described and figured the larva of linearis (canaliculatus),
and disputed the accuracy of Heeger’s description of a legless larva of
pubescens.

In 1880 Kittel described—after Noerdlinger (1855)—the larva, pupa, and
adult of linearis (canaliculatus).

In 1883 Dugés described and figured the larva, pupa, and adult of
L. planicollis, Le Conte (L. carlbonarius, Waltl.), to prove that the legless
larva described and figured by Heeger could not belong to the genus Lyctus.

In 1890 Rye ee to the disaonoes of a legless larva of “‘a species of
Lyctus” (?).

In 1898 Xambeu described the life-history of linearis (canaliculatus)
including a description of the egg, and stated that the eggs were deposited in
cracks or fissures; this description was corroborated Ibe iperean (1900), and
since by several abions:

In 1916 Snyder described the egg and manner of oviposition of planicollis,
and figured the egg. This egg differs from that of pubescens as described
and figured by Heeger, and from that of linearis as described by Xambeu,
but it is practically identical in appearance with the egg of brunneus found
by the writer. This similarity between the eggs of planicollis and brunneus
is maintained up to the time of maturation of the larva (Pl. 12. fig. 4), but
the method of the larva’s hatching differs.

Oviposition in brunneus is similar to that of planicollis, and it is not as
described by French in 1918, who states :—‘‘ The female (brunneus) deposits
her eggs on the outside, underside, and ends of the timber.” The eggs are
deposited inside the timber.

MATERIAL USED.

Most of the infested material from which L. brunneus was bred was obtained
in London from hardwoods stored in a timber yard, where the presence of the
beetle had been known for a few years.

Pieces of infested wood were collected in April 1920. The beetle was
then still in the larval stage. In May of the same year, a small quantity of
the infested material was placed in a refrigerator with a constant temperature

OVIPOSITION AND THE EGG OF LYCTUS BRUNNEUS. 219

of 8° C. to retard development; this material was withdrawn in July
1920 and enabled the writer to verify the observations on oviposition and the
egg at leisure.

The infested woods collected were West African mahogany (Khaya sp.),
““ Wainscot” oak (Quercus Robur), and “Italian”? walnut (Juglans regia).
Beetles were also bred from locust (Robinia pseudacacia).

Untortunately no living specimens of L. linearis were obtained, although
‘efforts were made in various directions to procure some. No beetles emerged
from two pieces of oak—hitherto infested with 1. linearis—which were sent
ever from Paris, nor were any living larvee or pupee found. Dr. C.J. Gahan
informed the writer, that no record of the capture of this beetle in the British
Isles had come to his notice for some years.

Marine.

The beetles are sexually mature when they emerge. Mating takes place
immediately after emergence either at dusk or during the night. It was
observed in daytime; and on several occasions two beetles were found
occupying the same pupal chamber, but in no instance was mating observed
to last very long. Individual males fertilize several females, therein differing
from ZL. linearis as described by Xambeu (1898), who stated coition lasted
the entire night and the male then dies. In brumneus, females considerably
outnumber the males.

The length of the life of the females when free to mate and oviposit
averaged about six weeks ; the males lived two to three weeks. The food of
the adults consists of particles of wood-tissue.

OVIPOSITION.

The ovipositor is an exceedingly long and flexible organ, and when fully
extruded (Pl. 12. fig. 1) it is nearly the length of the beetle.

Oviposition begins two to three days after mating and takes place, at dusk
or during the night, in the tracheze, or vessels, or pores of the wood.
‘Consequently those woods in which the vessels are most numerous are most
liable to heavy attacks.

The female either projects the ovipositor directly into the vessel, or it is
eurved down and bent forward in the vessel underneath the body. Before
-actually inserting the ovipositor into a vessel, a preliminary examination of
the surface is made with the ventral pygidial palps, and after selecting a
vessel the ovipositor is slowly inserted, and apparently a further examination
is made within the vessel by the vaginal palps (p, text-fig. 1) on the apex
of the ovipositor ; when, if the conditions are suitable, two or more eggs are
deposited. The number, however, depends upon the suitability and capacity
of the vessel. For, when an obstruction is encountered——in the form of
‘broken down transverse walls, etc.—only one or two may be deposited, or the

ovipositer will-be withdrawn and another vessel tried.
Ge

220 MR. A. M. ALTSON ON THE METHOD OF

The eggs are always laid longitudinally in the vessel and in juxtaposition,
the anterior pole of one being end onto the posterior pole of the next.

A female will retain the ovipositor in the vessel for several minutes, and
repeat the process in other vessels until her supply of ripe eggs is exhausted.

The egg issues from the vaginal orifice (vo., text-fig. 1) between the basal.
pieces (bp.), and is guided by the vaginal palps (p.).

Trext-ria. 1.

Apex of the ovipositor, ventral. an., anus ; 6c., bursa copulatrix ; bp., basal
piece; fr., forked rod; p., vaginal palp; 7d. chitinised rod;
vo., vaginal orifice. Camera lucida. (X128.)

From observations on a beetle when ovipositing, it was noticed that she
moved forward once before withdrawing the ovipositor ; apparently the
movement was just far enough to allow for a space for the second egg to be

deposited in

OVIPOSITION AND THE EGG OF LYCTUS BRUNNEUS. 221

Oviposition also takes place—in the case of previously infested wood—in
those vessels which have been bitten across in the pupal chamber and its
extension to the exit-hole.

It was observed that, when the beetles are feeding on the surface of the
wood, they always bite the tissues transversely; this habit—if the wood is
longitudinally split—not infrequently cuts open a vessel and creates a point
of access to it. In several instances, examination has shown that this opening
in the vessel has been used for the deposition of eggs.

The eggs are deposited in the vessels at varying distances from the point
of access of the ovipositor, but the anterior pole of the last egg laid is seldom
less than 1 mm. from it. In one specific instance in which three eg
found, the distance from the point of acvess to the posterior pole of the first
egg deposited was 3°75 mm. ; on two occasions five eggs in juxtaposition were
found, in these cases the ovipositor had been inserted more than 5 mm.

The eggs were difficult to find and very easily broken when shaving off the
wood-tissues in search of them, and when exposed, the task of removing them
or “digging ” them out is an exceedingly difficult one. The writer estimates
that on an average 75 per cent. of the eges were lost in trying to locate them,
and probably 60 per cent. of the located eggs were damaged or destroyed in

gs were

endeavouring to extract them from the vessels.

From the examination of several ovaries, it was found that in the case
of fertilized females, only a small number of eggs mature at a time,
eight to twelve beiny the usual number found collected in the calices.
In the case of unfertilized females, the ripe eggs continue to pass into the
calices until the latter become swollen and the death of the female ensues.

Toe Hae,

Pl. 12. fig. 2 is a photograph of deposited eggs in situ; part of the vessel
and surrounding tissues have been removed.

The egg is translucent white and cylindrical; tapering towards and
rounded at the posterior pole. The anterior pole is rounded, but continuing
from it, as if broadly attached to it, is a long, slender tube-like process
which terminates in a round protuberance.

This process varies considerably in length even in those eggs laid by one
female. The process was never observed to be attached to the walls of the
vessel, but when two or more eggs had been deposited in the same vessel it
was usually found adhering to the egg next to it.

The chorion is creased longitudinally, giving the egg the appearance of
being marked with longitudinal strive, which concentrating at the anterior
pole are more pronounced in this area. These striations and the process or
strand were found to owe their origin to the action of chitinised sete lining
a pair of valves situated at the junction of the oviducts, and are the result
of the pressure exerted by the valve upon the egg as it is forced through on
its passage.

222 MR, A. M. ALTSON ON THE METHOD OF

A ripe egg removed from the calices of a female does not show the:
process or striations, and would thus agree with Xambeu’s (1898) descrip-
tion of the egg of L. linearis.

The recently deposited egg (PI. 12. fig. 3) has a distinct granular appear-
ance, due to the exceptionally large yolk granules.

The size of the deposited eggs was found to range from 1:25 mm. to:
0-8 mm. in length without the process, which varied in length from 0:2 mm.
to 0°35 mm., and the width of the egg ranged from 0°15 mm. to 0°175 mm.
During maturation the ege slightly increases in size.

In eight specific instances, comprising 13 eggs under observation, the
young larva reached maturity 15 days after the egg was laid. That is,
movements of the larva were visible within the chorion at that period.
The actual number of hours any of these eggs took to reach this stage is not
known, as the time at which oviposition took place was not observed. But
the small pieces of wood in which the beetles were allowed to oviposit were
put in their cages—glass-topped tins—in the evening and collected the
following morning, 12 to 16 hours later.

At the time of maturation or when the first movements of the larva are
visible, it occupies approximately half the length of the egg (Pl. 12. fig. 4).
and is situated in the posterior portion ; the anterior portion, from the base
of the process to the head of the larva, is occupied by a mass which consists
of large yolk granules and fat bodies. The writer has observed that this.
extraordinary mass” constitutes in L. brunneus the initial food of the young
larva whilst still enclosed within the chorion.

In the case of L. planicollis it is stated that: “In hatching, the larva
backs out of the egg” (Snyder, 1916). It is the reverse with L. brunneus.
So soon as the larva is matured, it commences to eat the residual yolk-mass
situated in the anterior portion of the egg by means of its mandibles, and
travels forward to do so. If an egg is completely exposed in the vessel,
the larva is unable to consume the entire residual yolk-mass, owing to its
movements and the lack of “overhead”? support—the wall of the vessel—it
usually breaks the chorion and works its way out of the vessel. But if small
strips of tissue, such as parts of a medullary ray, are left above part of the
egg, the larva is. able to maintain its position and consume its food, and at
the same time the observer is enabled to follow its normal movements.

An examination of longitudinal and transverse sections of a small number
of eggs in different stages of development, suggest that the phenomenon of
the residual yolk-mass is due to the blastoderm enveloping only a part of the
yolk. It was thought that it might be a case of polyembryony, with one or
more embryos abortive, but there was no evidence found to support this.

* Owing to the lack of an existing term for this “ yolk-mass”—no analogous yolk-mass.

being known—the term residual yolk-mass has been suggested and is used in this paper to.
denote it.

OVIPOSITION AND THE EGG OF LYCTUS BRUNNEUS. 223

Text-fig. 2 gives a series of diagrammatic sketches showing the matura-
tion of the egg in four stages.

The movements of the larva are easily discernible within the intact
covering of the chorion, and it is not until the larva has been feeding for
some time that the chorion becomes broken by its movements at the posterior
end and later injured in the process of the consumption of its initial food.

Pl. 12. fig. 4 depicts an egg in which the larva has commenced feeding on
the residual yolk-mass in which its head is partially buried. (The bend in
the egg was due to the movement of the larva when dropped into Carnoy II.)
The photo shows that the chorion at the posterior end is broken, and that it
is crumpled in the part occupied by the larva, whereas it is quite taut

around the anterior part.
TExtT-FIG. 2.

oO
S 0)
oeeeaees ae C, ft 04

np sberrpiecs, A

°
ry
°

0 PO O00," OP v
05 0080 L202

Development of the egg (diagrammatic). 1. Egg twelve hours old. 2. Several days later.
3. About ten days old. 4, Mature egg. :

a., anterior pole; ch., chorion; ec., embryonic area; em., embryo; fym., formation of
residual yolk-mass; p., posterior pole; pc., process or strand; sym., residual yolk-
mass; y., yolk.

Locating THE Hees.

In the first instance the beetles were caged in a cavity (14 inches in
diameter) cut with a brace and bit in pieces of mahogany (4 inches by
3 inches by 1 inch thick). The cavity, which was full of cracks, fissures,
and crevices, was about 3 inch deep, and it and the surface of the piece of
wood were covered with a piece of glass held in position with elastic bands.

Shaving off the wood with a sealpel under binoculars in search of the

224 MR. A. M. ALTSON ON THE METHOD OF
'

eggs, after the manner of oviposition was established, proved far from
satisfactory and extremely laborious owing to the mass of wood which had
to be cut away from the sides to shave down those parts to get near the
vessels which had been opened transversely and accessible to the beetles.
And so another and entirely successful method was adopted.

Glass-topped tins were used as cages. Into ihese small pieces of
mahogany were put for the beetles to oviposit in. The sizes. of these
pieces, which were split on all faces longitudinally with the vessels, and
cut transversely at the ends, ranged from 1 to 2 inches in length by one-
eighth to about three-eighths of an inch in width and thickness.

The search for the eggs in these pieces was carried out as above and was
mainly confined to the extremities, unless a vessel had been fractured in
splitting or bitten open by a beetle.

No attempt was made to find eggs in planks or in the “ field.”

Note on Lyrervs riwearrs, GOEZE

As no living specimens of L. linearis were obtained, the writer’s intention,
to study the early stages of this insect with a view to critically examining
the descriptions of the manner of oviposition and the egg as published
by Noerdlinger (1855) and Xambeu (1898), did not materialise.

Several writers since 1898 to 1920 have published corroborative accounts
of Xambevw’s description of the egg and manner of oviposition, But it has
recently been found that in 1917 Hopkins and Snyder—after the latter’s
discovery of the egg and manner of oviposition of L. planicollis—published
a paper in which they described the life-histories of L. linearis, L. parallelo-
pipedus, Melsh., L. cavicollis, Lee., and L. planicollis, as being identical
except as to the time of the emergence of the adults. The inference to be
drawn is that the egg and manner of oviposition is similar in these four
species and consequently similar to the egg and manner of oviposition of
L. brunneus as described in this paper.

Therefore Xambeu’s description of a strandless egg deposited in cracks,
fissures, or crevices can no longer stand.

It will probably be found that this method and manner of oviposition—
depositing the eggs in the tracheze, vessels, or pores—is a generic charac-
teristic of the ‘aod infesting Lyctus beetles.

CONCLUSIONS.

The phenomenon of the egg of L. brunneus is, so far as it has been
possible to ascertain, unlike any case of embryological development recorded,
not only in the records of Entomology, but in those of Zoology.as a whole ;
a case in which within the egg is produced the young larva’s initial food.

The closest analogy found is that of a case of polyembryony discussed by
Gatenby (1919) in a review on the early development of the egg and the

OVIPOSITION AND THE EGG OF LYCTUS: BRUNNEUS. 225

formation and maturation of ‘the larvee of a polyembryenic Lncyrtid
(Parasitic Hymenoptera), in which he refers to the existence of abortive
embryos. He states: “It is remarkable to find that in the polyembryonic
Hymenoptera a large region of the egg is entirely discarded. In fact, just
that region of the egg which would have formed the head, brain, etce., of
the embryo is rejected.”

It is the same area in the egg of L. brunneus which becomes the larva’s
initial food.

The similarity of the egg of L. brunneus to that of L. planicollis—as
described and figured by Snyder (1916)—in their appearance up to the time
of maturation, suggests that the embryological phenomenon of the former
species must also exist in the latter.

SUMMARY.

1. The method and manner of oviposition in LZ. brunneus is established and
is found to be the same as in L. planicollis, which is that of depositing its eggs
in the tracheze, vessels, or pores, and under the surface of the wood.

2. The eges incubate in 15 days. The young larva, which occupies barely
half the length of the egg, does not hatch out at once, but proceeds to eat the
residual yolk-mass contained in the anterior part of the egg.

ACKNOWLEDGMENTS.

The invéstigation, of which this paper records part of the results, was
suggested by Prof. H. Maxwell Lefroy, Imperial College of Science, to
whom the writer has to express his thanks, and to the Committee of the
Scientific and Industrial Research Department, for a grant to carry on
the work.

The writer is also indebted to Dr. C. J. Gahan, Keeper of the Depart-
ment of Hntomology, Natural History Museum, for identifying specimens
of L. brunneus; to Prof. Percy Groom, Imperial College of Science, for
identifying the various species of timber used in this work; to Dr. L. T.
Hogben, lately Lecturer in Zoology, Imperial College of Science, for his advice
and assistance in regard to the ege of L. brunneus ; to Dr. Hugh Scott,
Cambridge Museum, for specimens of L. linearis ; to M. P. Lesne, Muséum
d’Histoire Naturelle, Paris, for sending two pieces of oak infested by
L. linearis ; to Dr. J. W. Munro, Forest Hntomologist, Board of Agriculture,
and to Mr. W. Dallimore, Royal Botanic Gardens, Kew, for material.

Tn addition, thanks are due to Dr. Av D. Imms, Rothamsted Experimental
Station, for his advice and assistance in connection with the’ publication of
this paper ; and to Prof. S. MacDougall for his efforts to get the original
paper published as a whole.

226 MR. A. M. ALTSON ON THE METHOD OF

REFERENCES.

Berwese, A. (1909).—‘Gli Insetti.’ Milan.

Bureav, L. (1900).—“ Le Lycte canaliculé (Lyctus canaliculatus Fab.) et les ravages quik.
fait dans les parquets et autres bois ouvrés (moeurs, expertise,
jugements) ”: pp. 169-201. Bull. Soc. Sci. Nat. lOuest France,
tome x.

Dueks, E. (1883).—“ Metamorphoses du Lyctus planicollis Le Conte”: pp. 54-58, pl. i.
Annales de la Soc. Ent. de Belg., tome xxvii.

Fowxer, W. W. (1891).—‘ The Coleoptera of the British Islands’: vols. iv. and vi. (1913).

Frencu, C., Junr. (1918).—“ Furniture and Timber Boring Insects”: pp. 214-221. Journ.
Dept. Agric. Victoria, Australia.

Froeeatr, W. W. (1907).—‘ Australian Insects’: pp. 170-171.

Froeeatt, W. W. (1920).—“ The Powder-Post Beetle and its Parasite”: pp. 273-276,
2 figs. Agric. Gaz. N.S.W., Sydney, vol. xxxi. No. 4, April.

Ganan, C. J. (1920).—‘‘ Furniture Beetles, their Life-history and how to Check or Prevent
the Damage caused by the Worm.” Brit. Mus. (Nat. Hist.) Econ-
Ser. No. 11. London.

GaTEnBy, J. Bronts. (1919).—“ Polyembryony in Parasitic Hymenoptera—A Review
pp. 175-196. Q. J. M.S. New Series, vol. lxiii, London.

Hereer, E. (1853).—“ Beitriige zur Naturgeschichte der Insekten”: pp. 938-989, pl. v.
Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften,
Math.-Naturw., bd. xi. Vienna.

Hewneeoy, L. E. (1904).—‘ Les Insectes.’ Paris.

Hopkins, A. D. (1911).—*“ A Revision of the Powder-Post Beetles of the family Lyctide
of United States and Europe.’ By E. J. Kraus. Appendix:
“Notes on Habits and Distribution with list of described Species.”
By A. D. Hopkins. Tech. Ser. Bull. No.20, Part II. U.S. Dept.
Agric. Bur, Ent.

Hopkins, A. D., & SyypEr, T. E. (1917).—“ Powder-Post damage by Lyctus Beetles to
seasoned Hardwood.” Farm. Bull. 778, U.S. Dept. Agric. Bur. Ent.

Katrensacu, J. H. (1874).—‘ Die Pflanzenfeinde aus der Klasse der Insekten’: pp. 185,
546, 644. Stuttgart.

Kerr, J. Granam, (1919).—‘ Text-book of Embryology.’ Vol. ii. VIER London.

Keene E., & K. Herper. (1899).—‘ Text-book of Embryology.’ Vols. i-ii, Trans-
lated by M. Bernard and M. F. Woodward. London.

Kirret, G. (1880).—‘Systematische Uebersicht der Kiifer, welche in Baiern und der
nichsten Umgebung vorkommen”: pp. 104-105. Correspondenz-
blatt. Zoologisch-Mineralogischer Verein in Regensburg, bd. 34.

MacBring, E. W. (1914).—‘ Text-book of Embryology.’ Vol. i. Invertebrata. London.

Norrpuinecer, H. von. (1855).—* Die kleinen Feinde der Landwirthschaft, oder Abhand-
lung der in Feld, Garten und Haus schadlichen, oder lastigen
Kerfe, sonstigen Gleiderthierchen, Wiirmer und Schnecken, mit
besonderer Beriichsichtigung ihrer natiirlichen Feinde und der
gegen sie anwendbaren Schutzmittel,’ etc.: pp. 189-191. Stutt-
gart und Augsburg.

Norrpiinerr, H. von. (1862).—“ Der Splintkifer, Lyctus canaliculatus L.” Kritische
Blatter fiir Forst- und Jagdwissenschaft, von Dr. W. Pfeil: pp. 284—
238, bd. 48. Leipzig.

Pacxarp, A. 8. (1909).—‘ A Text-book of Entomology.’ New York.

Perris, Ep, (1876).—“Laryes des Coléoptéres”: pp. 60-63, figs. 247-250. Ann, Soe. Linn.
de Lyon, tome xxiii.

oH A

Aiknes Journ. Linn. Soc, ZOOL VoL XXXV. PL.

A™M.A Phot .ond

Westwoca
Bequest

LYCTUS BRUNNEUS Steph.

OVIPOSITION AND THE EGG OF LYCTUS BRUNNEUS. 227

RuprertsBerGER, M. (1880).— Biologie der Kifer Europas’: p. 178. Donau.

RupERTSBERGER, M. (1894).—‘ Die Biologische Literatur tiber die Kifer Huropas von
1880 an’: p. 181. Linz a. d, Donau.

Ryn, I. C. (1894).—‘ British Beetles,’ second ed., revised and in part re-written by
W.. W. Fowler.

Smita, J. B. (1907).—“ Powder-Post Beetles”: p. 39. New Jersey Agric. Exp. Sta. Bull,
208.

Snyper, T. EK. (1916).—“ Ege and Manner of Oviposition of Lyctus planicollis.” Reprinted

: from Journ. Agric. Res. vol. vi. No. 7, Dept. Agric. Washington,

D.C.

Sreppine, H. P. (1914)—‘ Indian Forest Insects of Keonomic Importance.’ Coleoptera,
p- 177.

SrepuEns, J. F. (1880).—‘Illustrations of British Entomology: Mandibulata,’ vol. iii.
pp- 116-117, pl. 18. fig. 4.

XamBerv. (1898).—“ Mceurs et Métamorphoses du Lyctus canaliculatus Fabricius”: pp. 69-
72. Bull. Soe. Sci. Nat. ’Ouest France, tome viii.

EXPLANATION OF PLATE 12.

Fig. 1. Photomicrograph of female Z. drunneus, with ovipositor fully extended ; magnified)
7 diameters.
2. Two eges im situ in a piece of mahogany.
3. Photomicrograph of an egg, 12-16 hours old.
4. Photomicrograph ot a mature ege.

(Fig. 1 by C. Gunns; figs. 2-4 by A. M. A.)

thee

ey

(tiers Os ara
ae Sty Liar rey

Nagst a

vat

ECHINODERMS FROM WEST. AUSTRALIA. 22%

Some Echinoderms from West Australia. By Husrrr Lyman Crark,
Museum of Comparative Zoology, Cambridge, Mass., U.S.A. (Com-
municated by Prof. W. J. Daxin, F.L.S.)

[Percy Sladen Trust Expedition to the Abrolhos Islands under the
leadership of Prof. W, J. Daxry,]

(PLATE 13.)

[Read Ist February, 1923. ]!

Tue collection of echinoderms placed in my hands by Professor Dakin *
prove to be of great interest even though the number of species represented
is small. The specimens were taken off the coast of West Australia, chiefly
among Houtman’s Abrolhos Islands, but a considerable number were dredged
near Fremantle, while one interesting specimen comes from Broome. Many
of the species are represented by but one or two specimens, and on this
account it is not possible to determine the relative abundance of the
different forms.

The 143 specimens represent 46 species, of which two (Ophiactis savignyi,
Echinocardium cordatum) are practically cosmopolitan. Of the others, 27 are
tropical species, most of which are common in the Hast Indies; their
occurrence at the Abrolhos is notable because those islands are so far south
of the usual range of the species; few, if any, extend their range nearly so
far southward on the eastern side of the continent. :

There are, in the present collection, 8 species which occur on the southern
or south-eastern coasts of Australia between Sydney and Perth, though most
of them are known from only a few widely separated stations. These, which
may be called the typical Australian species, are :—

Astropecten preissi Miller & Troschel.
LIuidia maculata australasie Doderlein.
Asterina gunnii Gray.

Coscinasterias calamaria (Gray),
Ophiothria spongicola Stimpson.
Amblypneustes pallidus (Lamarck).
FHeliocidaris erythrogramma (Valenciennes).
Breynia australasie (Leach).

* I beg to express here my sincere thanks to Professor Dakin for the opportunity of
studying this valuable collection, I would also thank my friend Mr. Austin H. Clark for
very important assistance in connection with bibliographical data, which were not available.

to me,

230 MR. H. L. CLARK ON SOME

There are 9 species which seem to be endemic; at least they are not yet
‘known from anywhere but the coast of West Australia :—
Pentagonaster stibarus H. L. Clark.
Anthenea australie Doderlein.
Anthenea globigera Déderlein.
Nectria macrobrachia, sp. nov.
Peiricia obesa, sp. nov.
Parasterina crassa (Gray).
Uniophora dyscrita, sp. nov.
Ophiothrix michaelseni Koehler.
Centrostephanus tenuispinus H. L. Clark.

The following species were taken only off Fremantle :—

India maculata australasie Doderlein.
Astropecten preissti Miiller & Treschel.
Asterina gunnii Gray.

Parasterina crassa (Gray).

Ophiothrix michaelsent Koehler.
EHehinocardinm cordatum (Pennant).

Three species were found both at Fremantle and at the Abrolhos :—

Anthenea australie Doderlein.
Ophiothriv spongicola Stimpson.
Oplwothria stelligera Lyman.

From Broome alone, comes a specimen of

Euryale aspera Lamarck.

Although Gray described several sea-stars from “Swan River” and
“¢ Western Australia” as far back as 1840 (Ann. Mag. Nat. Hist. vol. vi.
Dec. 1840, p. 281) and 1847 (Proc. Zool. Soc. London, pp. 75-83), almost
nothing else was published dealing with the echinoderms of that region until
after the close of the nineteenth century. In 1907, Koehler reported on the
collection of ophiurans made by Michaelsen and Hartmeyer between Cape
Naturaliste and Sharks Bay in 1905. (See their ‘Die Fauna Siidwest-
Australiens, Bd. i. Lief. 4.) There were 28 species in this collection, of
which only three were new, and most of the others were well-known Indo-
Pacific species. In 1911, A. H. Clark published his invaluable report on
the ‘Recent Crinoids of Australia” (Mem. Austral. Mus. vol. iv. pp. 703-804)
and also a report on the collections made by Michaelsen and Hartmeyer
(Fauna Siidwest-Australiens, Bd. iii. Lief. 13), the two papers giving a
complete list of Western Australian Crinoids. In 1914, Déderlein reported
-on the echini brought back by Michaelsen and Hartmeyer (Fauna Siidwest-

ECHINODERMS FROM WEST AUSTRALIA. 231

Austral. Bd. iv. Lief. 12), which included 16 species of which three were new.
The same year, Mr. B. Alexander published (1914, Rec. West Aust. Mus.
vol. i. pt. 1, pp. 108-112) a list of the echinoderms of Western Australia
found in the Western Australian Museum at Perth. This list was based on
papers by A. H. Clark and myself published in the same part of the
““Records,”? dealing with the crinoids and the other echinoderms res- .
pectively. In his list Alexander records 86 species, of which half a dozen
are holothurians. As there are no holothurians in the collection sent me
by Professor Dakin I shall ignore that class, and the present remarks deal
only witb the actinogonidiate echinoderms*. Alexander includes only ten of
the 28 ophiurans of Koehler’s 1907 list and only five or six of Déderlein’s
list of echini, there being some question about certain identifications in this
group.

In 1918, Mortensen (Kungl. Svenska Vet.-Akad. Handl., Bd. 58. no. 9)
published a report on echini collected by Mjéberg at Cape Jaubert and
Broome, a region from which almost nothing was previously known. This
-eollection contained 14 species, of which only four are found in Déderlein’s
list from the south-western coast. In 1919 appeared Gislén’s (Kungl. Svenska
Vet.-Akad. Handl., Bd. 59. no. 4) admirable report on the crinoids collected
by Mjoberg, in which is given a careful and very valuable account of seven
species, one of which had not previously been reported from Western
Australia.

Up to the present time therefore, 125 species of echinoderms, not including
holothurians, have been reliably reported from Western Australia, and there
care perhaps 10 or 11 species which could be added to this list on the strength
of old records which, may not properly be ignored. This gives us a total of
at least 1485 species occurring in the region, already reported, but as the
Dakin collection contains, no fewer than a dozen species not hitherto recorded,
it is evident that the total number of echinoderms occurring on the western
side of the Australian continent certainly exceeds 150, and it is probably in
excess of 200.

It is interesting to examine separately the list of species occurring at the
Abrolhos, as those islands are said to contain the southernmost coral reefs in
the world. Including the species here recorded, the list of forms known
from the Abrolhos is as follows :— ;

CRINOIDS :—

Comatella nigra (P. H. Carp.). Comanthus parvicirra (J. Miill.).

C. stelligera (P. H. Carp.). C. polycnemis A. H. Clark.
Comatula solaris (Lam.). Amphimetra jacquinoti (J. Miull.).
C. pectinata (L.). Lanwprometra gyges (Bell).
Comanthus annulata (Bell). Ohyometra serripinna (P. TH. Carp.).

* (The Holothurians from West Australia are in the hands of Dr. Jos. Pearson, Colombo
Museum.— W. J. D.]

232 : MR. H. L. CLARK ON SOME

ASTEROIDS :—

Nectria macrobrachia, sp. noy. Fromia andamanensis Koeh.

Pentagonaster stibarus H. L. Clark. F.. elegans H. L. Clark.

Stellaster incet Gray. Bunaster lithodes Fisher.

Anthenea australia Dod, Petricia obesa, sp. nov.

Anthenea globigera Dod. Asterina burtonit Gray,
OPHIUROIDS :—

Ophiactis savignyi (M. & T.). Ophiothrix stelligera Lym.

Ophiothrix longipeda (Lam.). Ophionereis porrecta Lim.

O. michaelseni Koeh, Ophiocoma brevipes Peters.

O. spongicola Stimpson, Ophioplocus imbricatus (M. & T.).
EicHInoips :—

Prionocidaris baculosa var. annulifera Salmaeis virgulata Ag. & Des.

(Lam.). ; Pseudoboletia indiana (Mich.).

P. bispinosa var. chinensis (Déd.). Tripneustes gratilla (L.).

Centrechinus savignyt (Mich.). Heliocidaris erythrogramma (Val.).

C. setosus (Leske). Echinometra mathaei Bl.

Centrostephanus tenuispinus A. L. Clark. Breynia australasie (Leach).
Amblypneustes pallidus (Lam.).

OF these 40 species, now known from the Abrolhos Islands, three-fourths-
may be reckoned as tropical species, for L is tropicopolitan, 25 occur in the
Hast Indies, and 4 others occur along the coast of northern or at least north-
eastern Australia. Of the remaining 10,7 are endemic and 3 are South
Australian forms. Of the endemic species, 3 (Nectria macrobrachia, Petricia
obesa, and Centrostephanus tenuispinus) are most nearly allied to species from
the southern coasts of Australia. It is evident, therefore, that the Abrolhos
have received their echinoderm fauna chiefly from the north, and that only
a very small number of characteristically Australian forms have as yet found
a footing among those islands.

CRINOIDEA.

CoMATELLA NIGRA.

Actinometra nigra P. H. Carpenter, 1888, ‘ Challenger’ Comat. p. 304.
Comatella nigra A. H. Clark, 1908; Smithson. Misc. Coll. li. p. 207,

A single well-marked specimen dredged off Long Island, Abrolhos. It is.
nearly black, has 40 arms about 110 mm. long, and the very powerful cirri
are XXII, 25-28. The disk is about 32 mm. across, orally. This species
has not been recorded hitherto from western Australia, though it is known
from the Aru Islands and I found it at the Murray Islands in 1913.

CoMATELLA STELLIGERA.
Actinometra steligera P. H. Carpenter, 1888, ‘ Challenger’ Comat. p. 308.
Comatella steligera A. H. Clark, 1908 ; Smithson. Misc. Coll. hi. p. 207.
A single specimen of this northern species was taken on the shores of
Wooded Isle. It is bright yellow-brown, in its preserved condition, is.

ECHINODERMS FROM WEST AUSTRALIA. 233

25 mm. across the disk, and has 25 arms: 90 mm. long more or less.
Cirri XXX, 19-22. There is only one record of C. stelligera from a point
anywhere nearly so far south as Wooded Isle, and that is the isolated and
dubious record from Port Jackson. The species ranges from Ceylon to
Samoa and is very common at the Murray Islands, east of Torres Strait, but
is not known from the coast of Queensland.

CoMATULA SOLARIS.

Lamarck, 1816, Anim. s. Vert. ii. p. 533.

A specimen with the 10 arms, 125 mm. + long and about 7 mm. wide,
near base, was taken at Hast Wallaby Island. It is almost black, but has a
longitudinal light stripe on the dorsal side of each of the arms, which are
remarkably stout. A second specimen not quite so large, taken in the
dredge off Long Island, has no trace of the light stripe on the dorsal side of
the arms. It has the cirri XIII, 18.

ComMATULA PECTINATA.
Asterias pectinata Linné, 1758, Syst. Nat. ed. x. p. 663,
Comatula pectinata A. H. Clark, 1908, Proc. U.S. Nat. Mus. xxxiii. p. 685.

This comatulid seems to be rather common at the Abrolhos, as there are
13 specimens in the present collection: 9 dredged off Long Island and 4
dredged near First Island. All are brown, pale brown, or yellow-brown in
colour. They are small, only two or three having arms 100 mm. long. The
cirri range from I-XIV, with segments 10-14, but in no case are they
arranged in pairs at the corners of the centrodorsal as they are in C. purpurea.
Otherwise these specimens would, because of their small size, be more
naturally assigned to that species, which Mr. A. H. Clark has recorded from
“between Fremantle and Geraldton.” Several of the specimens examined
by Mr. Clark were not typical C. purpurea, and intergrade evidently with
C. pectinata. Gislén (1919) has found so much intergradation between
C. pectinata and C. purpurea that he retains the latter name as varietal only,
and I am inclined to agree with him that it is certainly not specific. Just
what the relation between C. purpurea and C. pectinata really is requires still
further study.

JOMANTHUS ANNULATA.
Actinometra annulata Bell, 1882, Proc. Zool. Soc. London, p. 535.
Comanthus (Vania) annulata A. H. Clark, 1911, Mem. Austral. Mus. iy. p. 757.

The seven specimens of this handsome species were all taken on the shores
of Wooded Isle. The number of arms ranges from 36 to 47 and their length
from 100 to 125 mm.; in the specimen with 36 arms every Br series is
4(3 +4), but in the one with 47 there are four II Br series, 2 and one
IV Br series,2. Cirri very weak, V-VIII, 12. The uniformity of these
specimens in coloration is their most notable feature, and in this they agree

LINN. JOURN.—ZOOLOGY, VOL. XXXY. 17

234. MR. H. L. CLARK ON SOME

with seven of the specimens A. H. Clark (1914, Rec. W. Austral. Mus. i.
p- 120) reeords from between Fremantle and Geraldton, except that they
apparently have a darker ground-colour. But they are all spotted with
uniformly small, circular greenish-yellow dots, generally quite distinct, but
obscured in some of the darkest specimens. In view of the extraordinary
diversity of colour of this comatulid, at the Murray Islands, it is remarkable
that no diversity at all is shown at the Abrolhos. Possibly this western
form may be worthy of a varietal name, but further field observations are
necessary before a decision can be reached.

CoMANTHUS PARVICIRRA.
Alecto parvicirra J. Miiller, 1841, Arch. f. Naturg. vii. p. 145.
Comanthus parvicirra A. WH, Clark, 1908, Smithson. Misc. Coll. li. p. 203.

There are seven comatulids from Wooded Isle which seem to represent this
species, though the colour is the yellow-brown characteristic of C. luteofusca.
They range in size from those with arms 35-40 mm. long up to those whose
arms exceed 100 mm. ‘The disk is 7-12 mm. across. The arms range from
12 to 23 innumber. The cirri are few and weak, and have the characteristic
form and proportions.

AMPHIMETRA JACQUINOTI.

Comatula jacquinoti J. Miiller, 1846, Monatsh. d. k, preuss. Akad. Wiss. p. 178.
Amphimetra jacquinoti A. H. Clark, 1914, Rec. W. Austral. Mus. i. p. 124.

There are five specimens of this fine comatulid, but all are more or less
badly broken ; two were “dredged outside Wallaby Group,” while the other
three were ‘‘ dredged off Long Island.” They are each about 12 mm. across
the disk, and the arms were apparently about 100 mm. long. The cirri are
XXI-XXVI, 26-32, 33-39, 36-41, with the segments all much wider than
long and the dorsal teeth beginning at the 12th-15th segment. The calyx
and arms in each specimen are a dirty cream-colour, while the cirri are of
that shade only at the base, becoming purple distally.

LAMPROMETRA GYGES.
Antedon gyges Bell, 1884, ‘ Alert’ Rep. p. 160.
Lamprometra gyges A. H. Clark, 1913, Proc. Biol. Soc. Wash. xxvi. p. 144.

There are two light-brown specimens of this species collected along the shore
at Wooded Isle. The arms are about 100 mm. long, and their segments
proximally are so closely opposed to each other that the basal part of the
arm is noticeably smooth and regular. The cirri are XXX-X XXII, 25-30 ;
one specimen shows twenty-three additional cirrus-sockets. The cirri are
less brown, more grey than the calyx. In one specimen there can be
distinguished along the dorsal side of each fully-developed arm an incon-
spicuous longitudinal whitish line.

ECHINODERMS FROM WEST AUSTRALIA. 235

OLIGOMETRA SERRIPINNA.
Antedon serripinna P. H. Carpenter, 1881, Notes from the Leyden Museum, vol. iii.

pp. 175, 182.
Oligometra serripinna A. H. Clark, 1908, Proc. Biol. Soc. Washington, vol. xxi. p. 126.

At Long Isiand a single small specimen of this comatulid was taken, but
it is not typical. The ridge across the segments of the cirri is very incon-
Spicuous, and none of the pinnule segments have conspicuous projections.
There are ten arms about 35 mm. long, and the cirri are XII,17. P, has 10
segments, P, has the same number but is conspicuously bigger, while P; also
has 10 segments and is about equal to Py. In colour the general impression
is olive-cream and purple, the former being the ground-colour. There is a
distinct wide purple line up each side of Br; and B, basally, but this soon
breaks up into spots, and disappears in the joints or in lines across the
segments at their margins. There is no regularity of arrangement.

The occurrence of sermpinna at the Abrolhos is unexpected, as it has not
been hitherto recorded from Australia, but Mr. A. H. Clark considers this
specimen undoubtedly Carpenter’s species and not O. carpenteri Bell, which
has been recorded from northern Australia several times. (A. J. D.)

ASTEROIDEA.

ASTROPECTEN PREISSII.
Miller & Troschel, 1843, Arch. f. Naturg., jhrg. ix. 1, p. 119.

There are three Astropectens, dredged off Fremantle, which seem to be
undoubtedly this species. They have lost all indication of their original
colour and are now dingy light-brown. ‘The smallest has R = 30, r= 6, and
br = 6°5 mm.; thus R=5r. Superomarginal plates all unarmed. Infero-
marginals with a single wide, flat, pointed marginal spine, below which are
two very much smaller and more slender spines, of which the adoral is
smaller ; about four somewhat smaller spines form a longitudinal, well-
spaced series on the ventral surface of the plate. A second specimen has
R= 70,r= 10, and by = 13 mm.; hence R=7r. On onearm three of the
distal superomarginals bear small but distinct spinelets. The armature of
the inferomarginals is essentially as in the young individual, but the marginal
spines and those below are markedly heavier. The third specimen has
R= 120, r=15, and br = 17 mm.; hence R= Sr. There are no supero-
marginal spinelets, and the inferomarginal armature is also quite typical.

LUIDIA MACULATA AUSTRALIA.
Luidia austrahe Doderlein, 1920, ‘Siboga’ Ast. pt. ii, Luidia, p. 266.
An adult specimen, taken near Fremantle, has the usual seven rays, and
the measurements are R= 225, r= 29, and br =31 mm. The distal

paxillee are characteristically large, and there are some pedicellarize on the
Nias

236 MR. H. L. CLARK ON SOME

lateral paxillee near the arm-bases. The arms are rather wide near the tip,
not tapering so much as in some Chinese specimens. Although I have not
seen any really intermediate specimens, I think the Australian form is more
probably a local subspecies than a distinct species. Déderlein himself
suggests that it should perhaps be considered “nur als eine Lokalform.”
Material from the northern coast of Australia and the southern Hast Indies
is essential for a proper solution of the problem.

NEcrriA MACROBRACHIA *, sp. nov. (Pl. 18. figs. 5, 6.)

Diagnosis : Disk small ; arms relatively long and cylindrical. Granulation
of abactinal plates, near tips of rays, coarse, crowded, and prismatic;
actinal granulation noticeably prismatic. Abactinal plates of disk and arm-
bases less paxilliform and more crowded than in the other members of the
genus.

Description of holotype: Rays 5. R=60 mm.; r=17 mm.; R=3-5r.
Br=16 mm. Br at middle of ray=10 mm. ; at tip, 6mm. Disk relatively
small, only a little elevated ; arms relatively narrow, for the genus, some-
what flattened basally, but nearly cylindrical or terate for the distal half.
Abactinal plates on disk and on basal two-thirds or more of rays, large, low,
flat, more or less irregularly hexagonal with rounded corners, well spaced
but not widely separated, of dissimilar size; they are connected with each
other by heavy radiating ossicles, in the spaces between which arise papules
in groups of 4-14. Hach plate is covered by a coat of low, more or less
convex, polygonal granules ; there are about 40-50 on a plate 2—-2°5 mm. in
diameter, besides a marginal series of about 25 distinctly larger granules.
The marginal series of adjacent plates are in close contact even on. disk,
except here and there at the angles. Distal part of rays covered by similar
but coarser granules, quite closely crowded and without indication of
marginal series. Madreporite small, about 2 mm. across, situated half-way
between centre and margin of disk.

Superomarginal plates about 25, very similar to the abactinal plates in
covering and appearance ; interradial ones much higher than long ; distally
and especially close to tip of ray, the superomarginals are so closely crowded
against each other, the abactinal plates, and the inferomarginals that they
can be distinguished only imperfeetly. Inferomarginals about 27, but distal
ones very difficult to make out. The whole distal end of the ray is so closely
covered with coarse unequal granules that plate limits cannot be distin-
guished. Intermarginal, and even inframarginal, papillee are evident near
base of ray.

* nacpds—long+fpaxiov=arm, in reference to the characteristically long and relatively
slender arms.

ECHINODERMS FROM WEST AUSTRALIA. DONG

Actinolateral areas small; along the mid-interradial lines there are about
six series of actinolateral plates, but the outermost consists of only one or two
plates, the next two are little longer, the next is perhaps 10 mm. long, the
second series does not quite reach the middle of the ray and the first extends
far out, almost to the tip of the ray ; all these plates are covered by coarse
prismatic granules, much coarser than those of the abactinal surface ; the
largest and most prismatic granules are nearest the oral plates.

Adambulacral plates more than 50, the distalmost hard to distinguish ;
they form a slight undulating margin to the furrow ; each plate carries
3 furrow spines (distally only 2), which are subequal, not notably prismatic,
blunt, and slightly thicker at tip than at base; distally, as a rule, the
adoral is the smaller of the two; near the base of the arm the spinelets are
about 2 mm. Jong, and *50--70 mm. thick at tip. On the surface of each
adambulacral plate are 3-5 very short, thick, and prismatic spines; the two
largest of these adjoin the furrow spines, while the remaining 1-3 are near
and resemble the prismatic granules on the first series of actinolaterals.
Oral plates not conspicuous or peculiar ; even the oral spines are no larger
than those on the adjoining adambulacrals. No pedicellarize were seen
any where.

Colour light yellowish-grey.

‘There are two specimens of this interesting new JVectria, collected along .
the shore in the “ Pelsart group”; one label says “ Pelsart Island.” The
interbranchial septa are calcified, and the general appearance is so much like
Nectria that there can be little question of the generic position. And yet the
abactinal plates are much less paxilliform, and are much more crowded on
the disk and arm-bases than in the other members of the genus. The disk
is also very distinctly smaller, and the arms are narrower at base, wider at
tip, and more nearly cylindrical than in either N. ocellata or .N. ocellijera.
The granulation of the actinal plates, especially at the base of the arms, is
noticeably prismatic in W. macrobrachia, and that of the abactinal plates at
the tips of the rays is coarse, crowded, and prismatic ; these differences in
granulation, which seem trivial when put into words, are very conspicuous
when specimens are compared.

The paratype of NV. macrobrachia is a badly injured individual, much
smaller than the one described. There are but three rays ; two, side by side,
seem to have been bitten off very close to the disk, apparently at different
times, as they show different degrees of healing. The rays present are
strongly curved and contracted, but apparently R=42 mm. and r=11, so
that Ris almost 4r. The granulation and colour are exactly the same as in
the holotype. The armature of the adambulacral plates is also similar, but
the number of furrow spines is not reduced to two until ‘almost the very tip
‘of the arm is reached.

238 MR. H. L. CLARK ON SOME

PENTAGONASTER STIBARUS.

Hi. L. Clark, 1914, Rec. Austral, Mus. i. p. 136,

Two little pentagonal sea-stars from Pelsart Island appear to be the young
of this species. The smaller is 17 mm. across, R=9 mm., and has 73 flat
abactinal plates besides the madreporite, 4 superomarginals on each side,
and a terminal plate on each ray, 99 plates altogether abactinally. The
distal marginal plate on each side of each ray is the larger, but is not
conspicuously enlarged. Orally there are 4 inferomarginals on each side,
and the distal ones on each ray are noticeably larger than the proximal.
There are 12 (on 2) or 13 (on 3) plates on each of the interradial areas, and
there are 17 adambulacral plates on each furrow margin. The other
specimen is a trifle larger (18°5 mm. across), but has only 55 flat abactinal
plates besides the 20 marginals, 5 terminals, and madreporite, 81 plates
altogether on the dorsal side. On the ventral surface there are 14 (in
one area 15) plates in each interradial area. A few pedicellariz occur
on the abactinal surface of each specimen, but there are none on the oral
surface.

That these little sea-stars are really the young of stibarus seems clear as a
result of comparing them with the young of P. diibent. The latter, have the
abactinal plates swollen, the rays longer and narrower, and, at least in some
cases, actinal pedicellarice are present.

At Wooded Island a larger specimen of P. stibarus was taken, 27 mm.
across, R=15 mm. _ There are only 4 marginal plates still in each series, on
each side of the distinctly pentagonal sea-star ; the distal plate is distinctly
the larger in every case, Ina specimen of P. diibent of the same size there
are 40 superomarginals and 50 inferomarginals, 8 and 10 respectively on each
side of the animal, which is not, however, at all pentagonal, since R is
15 mm. and r only 8 mm.

STELLASTER INCEL.

Gray, 1847, Proc. Zool. Soc. London, pt. xv. p. 76.

Two specimens of this well-known sea-star were dredged off Long Island.
The larger specimen has R=90 mm. and the smaller 80 mm. There are
2-4 low, blunt spine-like tubercles on the carinal line, on the disk at the
base of each arm. The number and arrangement of the inferomarginal
spines show great diversity ; whereas the number of inferomarginal plates
is about 17 in each series, the number of spines ranges from 3 to 11.
The present specimens have lost all colour.

ANTHENEA AUSTRALIA.
Déderlein, 1915, Jahrb. Nassau. Ver. f. Naturk. Wiesbaden, Ixviii. p. 52.
Two Antheneas may well be referred to this species. The genus is a
perplexing one, and Doderlein’s revision of it is a very admirable piece of

ECHINODERMS FROM WEST AUSTRALIA. 239

work. His recognition of this species seems to be amply justified. The
smaller of the two specimens at hand has R=33 mm. and r=15, while
the larger has the measurements 65 and 30 mm. respectively. Hyen the
latter is only three-fourths as large as Déderlein’s type. According to the
label the smaller specimen was “ dredged between Rat and Pelsart Group.”
It differs from the larger in the paucity of granules and pedicellariz on
the abactinal surface, which is nearly.smooth. Hven the superomarginal
plates have very small groups of granules, especially near the interradius.
At the base of each arm, on the carinal line, 7 mm. from centre of disk,
is a low but conspicuous tubercle. There are two smaller tubercles near
the anus. Similar tubercles can be seen in the larger specimen, but as
they have not increased in size with the growth of the sea-star, they are
no longer conspicuous among the numerous granules and pedicellariz with
which the abactinal surface is covered. This larger specimen was dredged
off Fremantle. Its colour (dry) is brown-olive, with granules, tubercles, and
pedicellarize nearly white or at least very light brownish and the madre-
porite chocolate-brown ; on the oral surface the granules and pedicellarize
are so numerous and crowded that the general effect is much lighter than
dorsally. The smaller specimen is grey-brown aboye and light yellowish-
brown below. The larger specimen is very similar to specimens of
A, pentagonula of the same size, but it is evidently stili immature.

ANTHENEA GLOBIGERA.

Déderlein, 1915, Jahrb. Nassau. Ver. f. Naturk. Wieshaden, Ixviii. p. 50.

A very fine Anthenea, taken by “‘shore-collecting, Wallaby Group,” is
undoubtedly to be referred to this well-marked species. It is larger than
Déderlein’s type, in which R=59 and r=30 mm., for R=70 and
r= 34 mm. The radial series of tubercles is nearly wanting, though the
5 primary ones are evident. Large dorsal pedicellarise are numerous and
conspicuous. The tubercles on the marginal plates are smaller than in
Déderlein’s type. The dorsal surface is dark purple-brown, with tubercles,
pedicellarize, and madreporite conspicuously light-coloured in contrast ; the
oral surface is wood-brown, the tubercles, spinelets, and pedicellarize much
lighter.

FROMIA ANDAMANENSIS.

Koehler, 1909, ‘ Investigator’ Ast. p. 105.

Six specimens of a Fromia have been a source of great perplexity in the
study of the Abrolhos sea-stars, but I have failed to find any satisfactory
reason for not referring them to andamanensis. The unique holotype of
that species is from the Andaman Islands, without more definite locality,
and while it is recorded as from ‘238-290 fathoms,” I think there is
undoubtedly a mistake about the depth. Certainly the specimens from the

240 MR. H. L. CLARK ON SOME

Abrelhos are typical littoral sea-stars. None is exactly like Koehler’s
figure of F. andamanensis, but the one most like it is only a trifle larger
and has the rays just a little bit more slender. The abactinal plating is
very much as Koehler shows it, but the rays are not so flattened. This
specimen was taken at Pelsart Island, and no two of the rays are the same
length. The longest has R = 32 mm., the shortest only 17. The colour is
very light, almost a dirty whitish (dry). A specimen of about the same
size from “ Wallaby Group. Shore” is brown in colour, and the abactinal
plates are no more numerous than in the Pelsart specimen. At the base of
the ray one can count six or perhaps seven longitudinal series of these
plates. A larger specimen from the Wallaby station has R = 35 mm., and
there are nine or ten series of abactinal plates, which are noticeably smaller
and of more uniform size than in the other specimen. In this larger
specimen the rays are nearly equal, only one being noticeably shorter than
the others, and they are distinctly terete and not flattened, the height at
the base being 9 mm, and the width 11. The colour is brown as in the
smaller specimen. Shore-collecting at Long Island yielded an individual
very much like those from the Wallaby Group with R=35 mm., but having
the rays a trifle more flattened (11°5 x 85 mm.). Dredging off Long
Island yielded a slightly larger specimen, in which the longest R = 40 mm.,
but br = 10 mm. and height of arm at base about 8mm. This specimen
is also much lighter coloured, nearly white. Finally, from the reef-flat at
Pelsart Isle is a much larger Fromia, with R= 54 mm., br=12, and
height of arm at base only 7 mm., of a pale brown colour and having 10 or
11 longitudinal series of abactinal plates. The disconcerting feature of this
specimen is that many of the adambulacral plates on the basal half of
the rays have three furrow spines. ‘Thus the flatness of the rays and the
adambulacral armature approach closely to ’. milleporella. On the other
hand, the large number of series of abactinal plates and the more slender
rays give this large /romia a different facies from ordinary F’. milleporella,
and taken in connection with the presence of only two furrow spines on
most of the plates, warrant us in considering F’. andamenensis as a distinct
species. Information regarding the colour in life would perhaps be decisive,
but the labels with the present specimens are blank on that point.

FROMIA ELEGANS.

H. L. Clark, 1921, Mchinoderm Fauna of Torres Strait, p. 43.

There are two Fromias in the collection which are labelled ‘‘ Colour—
Brown dark. Locality—I1st Island. Dredge.” The smaller has all the rays
broken ; it resembles the small specimen of F’. andamanensis from Pelsart
Isle, but the abactinal plates are noticeably larger. The other specimen is
perfect and has R= 40 mm., r and br = 10 mm. The disk and rays are
quite flat, the height of the arms at base being only about 6-7 mm.

- ECHINODERMS FROM WEST AUSTRALIA. 24.1

T cannot find a single reliable character by which to separate this specimen
from F. elegans, yet the abactinal plates are not so large or well defined
as in that species. The coloration is now pale brownish, but in life it
must have been very similar to that of F. elegans, if we may trust the
label. On the whole it seems better to refer these two specimens from
“Ist Island” to #. elegans than to inelfide them under the name
LF, andamanensis.

BUNASTER LITHODES.

W.K. Fisher, 1917, Proc. Biol. Soc. Wash. xxx. p. 91.

There are four little ophidiasterids in the collections, two from Pelsart
Island and two from Wooded Island, which undoubtedly belong to the
genus Bunaster. They are certainly not B. ritteri, for pedicellariz are
present and there are no “ball and socket” plates, and they are not
B. uniserialis, as comparison with the holotype of that species shows.
After very prolonged study it seems best to consider them as young
B. lithodes, in the holotype of which species R = 22 mm. In none of the
Abrolhos specimens does R exceed 10mm. The most noticeable resemblance
between these specimens and Bb. lithodes is seen in the finely shagreen-like
covering (aside from the abactinal plates) of the upper half of the animal ;
in B. uniserialis the space between the abactinal plates is occupied by a
coarse granulation, as it is in B. ritterd also. In two points the Abrolhos
specimens are unlike B. lithodes; the subambulacral spines are very little
longer than broad, and it is hard to see more than a single row of actino-
lateral plates even at the base of the arm. These two points may be
dependent, however, upon maturity, and the characters are certainly liable
to growth-changes. In view, then, of the obvious immaturity of these little
Bunasters, I think they may well be referred to B. lithodes.

The two specimens from Pelsart Island have R= 8 or 9mm. They are
shell-pink, faintly variegated with whitish and decidedly lighter below than
above. One of the Wooded Island specimens has R = 9 mm., and the rays
are distinctly more slender than in the others. Its colour is also a very
much darker pink, The fourth specimen has R= 10 mm.; the rays are
stout and the pink colour has been entirely bleached.

PETRICIA OBESA*, sp. noy. (PI, 18. figs. 1, 2.)

Diagnosis : Rays wide, rounded at tip. Abactinal skeleton concealed by
the very thick skin with which it is covered. No tubercles, spinelets, or even
granules on distal marginal plates.

Description of holotype: Rays 5. R= 65 mm.; r= 30mm.; R= 2-17r.
Br = 32mm. Br at middle of ray = 24 mm.; at 5 mm. from tip, 15 mm.
Disk large, only slightly elevated; arms wide, slightly arched, very blunt

* Obesus = fat, in reference to the wide, blunt rays and the thick skin.

242 MR. H. L. CLARK ON SOME

and rounded at tip. Abactinal skeleton almost completely hidden, even
in the thoroughly dry specimens, by the very thick skin which covers the
whole animal. YPapule numerous, in large irregularly-defined and more or
less coalescent groups, absent only from the terminal fifth of the rays.
Anus well marked by the fine calcareous papillee which surround it, nearly
central in position. Madreporite 10 mm. from anus, small (2 mm.
across) but distinct. Interradial pedicellarize huge and very conspicuous,
the valves 4-5 mm. long. No other pedicellarize are present.

Marginal plates almost completely concealed by the thick skin ; the infero-
marginals appear to be larger than those of the upper series, and are larger
distally than interradially ; there are about a dozen in each series. They
bear no spinelets, nor is the surface at all rough or shagreen-like. Inter-
radial areas covered by thick, smooth skin, through which one can distin-
guish some actinolateral plates, of which the series adjoining the furrow
reaches to about the middle of the arm. Adambulacral plates about 50 on
each side of the furrow ; each plate carries two (rarely three and even more
rarely only one) spines, 2—-2°5 mm. long, shaped like a bowling-pin, blunt,
and even a little capitate at the tip. These spines are connected clear
to their tips by a fleshy fold of skin. Outside this told is the series of
subambulaeral spines, which are stouter than the furrow spines and are
buried nearly to their tips in skin, though in the distal part of the ray they
are more free. On some plates at the base of the arms there are two
subambulacral spines, but as a rule there is a single one on each plate.
Oral plates are not distinguishable, but they carry five, rarely six, spines
on each side of the jaw, the most proximal largest, about 3mm. long. On the
surface of the jaw are four short, thick spinelets, two on each side, one
proximal, one distal, buried in skin, and very similar to the subambulacral
spines. Colour (dry) dull chocolate, of somewhat varied hues, the darker
with a purplish cast ; adambulacral spines and jaws of interradial pedicellariz,
nearly white.

There are two specimens of this remarkable sea-star from Pelsart Island,
but they differ so much from each other in colour that it is hard to believe
they are of the same species. Unfortunately the labels say nothing as to
the colour in life, so we do not know how much of the present difference
may be artificial. The paratype is dull greenish-yellow ; everywhere the
outer surface layer of skin has cracked, permitting the deeper layer, which
is nearly white, to show through. This is particularly marked on the
actinal interradial areas. The interradial pedicellariz are smaller and
much less conspicuous in the paratype, but there are no other differences
worth noting. ‘he size is essentially the same.

The occurrence of Petricia on the western side of the continent is very
interesting, but the Abrolhos species is very different from P. vernicina, the
species which occurs at Port Jackson and along the southern coast. The

ECHINODERMS FROM WEST AUSTRALIA. 243

chief points of distinction are the wider and terminally rounded rays,
the thick skin which conceals the skeletal plates and is not at all smooth and
shiny when dry (the feature from which P. verncina gets its appropriate
name), and the absence of tubercles, spinelets, or even granules on the distal
marginal plates, in P. obesa. The West Australian species seems to be
somewhat larger than the one from the east coast, the largest specimens of
which have R=50-60 mm.

ASTERINA BURTONII.

Gray, 1840, Ann. Mag. Nat. Hist. vi. p. 289.

There are half a dozen very typical specimens of this little sea-star, one
dredged off Long Island and the other from Wooded Island. The largest
has R=26 mm. None of the specimens has retained any of its natural
colour. The discovery of this species at the Abrolhos extends its known
range very far to the south on the Australian coast.

ASTERINA GUNNII.

Gray, 1840, Ann. Mag. Nat, Hist. vi. p. 289.

There is a badly damaged small specimen (R=27 mm.) of this southern
species in the collection, but it is not from the Abrolhos. It was dredged off
Garden Island near Fremantle, which is probably about the northern limit
of the species on the western coast.

PARASTERINA CRASSA Fisher. (PI. 18. figs. 3, 4.)
Patiria (?) crassa Gray, 1847, Proc. Zool. Soc. London, p. 88.
Parasterina crassa Fisher, 1908, Smithson. Mise. Coll. lii. p. 90.

A sea-star dredged off Fremantle is apparently to be referred to this
species, as it answers well to Perrier’s (1875, Arch. Zool. Exp. y. p. 142)
rather detailed description except for its much larger size. Gray makes no
reference to the size of his specimen, but Perrier says “ d=10 mm.” From
the context one would infer that d meant the distance from tip to tip of
alternate rays—that is, the diameter of the entire animal; but in that case the
specimens in the British Museum are tiny indeed, and it seems incredible
that the description given could apply to so small an individual. If d refers to
the diameter of the disk, then Perrier’s specimen was only about half as large
as the one in hand. Possibly the 10 is a misprint for 100, but in that case
the present specimen is much smaller than those in the British Museum.
Perrier refers several times to resemblances to Pentanogaster. These led me
for a time to feel sure that my specimen could not be; P. crassa, as I see
nothing in which it is the least like Pentagonaster, save possibly the adam-
bulacral armature, where a slight resemblance might be imagined.

_..The present specimen has R=36 mm. and r=9, and R is therefore equal
to 4r; Perrier says R=32r, a difference easily understood if his specimen
had r only 5 mm. Perrier says that the large ossicles in the dorsal skeleton

244 MR. H. L. CLARK ON SOME

are more numerous than the small ones, and then says the papulee are
surrounded by the small ones, which certainly implies a considerable number
of the latter. In the present specimen, near the base of the arm there are
about five small plates to each one of the large ones. At the tips of the
rays, however, the large plates become more numerous. All plates, both on
the dorsal and oral surfaces of the sea-star, are closely covered_with cylin-
drical spinelets like those occurring in Asterina. Dorsally the spinelets end
in 2-4, usually 3, glassy points of which one is larger than the other ; ven-
trally there are more glassy points on each spinelet, and they are subequal.
Hven on the ventral plates, I cannot see that these crowded spinelets resemble
a“ oranulation”” as Perrier says. Gray says the colour of his dry specimen
was pale yellow, and the one before me might be called dirty yellow.
Perrier says that some of those in the British Museum were ‘“ encore variés
de jaune et de blanc,” but immediately states that the spines on the large
ossicles were “bleus”; probably either “blanc” or “bleus” is a typo-
graphical error.

COSsCINASTERIAS CALAMARIA.
Asterias calamaria Gray, 1840, Ann, Mag. Nat. Hist. vi. p. 179.
Coscinasterias calamarta Perrier, 1894, ‘Travailleur’ et ‘Talisman’ Stell. p. 106.

A sea-star dredged near Fremantle is the only representative in the col-
lection of this characteristic species of southern Australia. It is a small
specimen in poor condition, only three arms remaining attached to the disk,
which is but 10 mm. across. The rays are about 55 mm. long, and there
evidently were eleven of them originally.

UNIOPHORA DYSCRITA *, sp. nov.

Diagnosis: Disk moderate, little elevated ; arms short, wide, and little
arched. Inferomarginal, actinal, and adambulacral spines flattened, cbisel-
like at tip. No intermarginal plates. No shagreen-like areas on supero-
marginal plates. Abactinal spinelets of two kinds, blunt cylindrical and
capitate conical.

Description of holotype: Rays 5. R=55mm.; r=13mm.; R=4r+.
Br=15 mm. Br at middle of ray, 13 mm.; at 5 mm. from tip, 7 mm.
Disk moderate, little elevated; arms wide and little arched. The specimen
is dry and flattened, and it is hard to tell how much elevated and arched the
upper surface may have been in life. Abactinal skeleton of rather heavy
plates forming an irregular and very open network, the meshes of which are
four distinct series only at the base of the arm; at the middle of the arm
the lateral series are still evident, but the median series are quite replaced
by irregularly-arranged meshes 3-5 mm. across, more or less occupied, at
least at centre, by papulee. Basally there are 4—12 papulve in the mesh-areas.

* §voKpiros=hard to determine, in reference to the difficulty of distinguishing the species
of Uniophora.

ECHINODERMS FROM WEST AUSTRALIA. 245.

Marginal plates very distinct, the upper series forming a very evident boun-
dary to the abactinal surface of the arm. Intermarginal papule numerous.
At the base of each arm a carinal series of plates is evident and one lateral
series on each side, but on most of the abactinal surface of each arm, as well
as on the disk, there is no definite arrangement of the plates evident. Most
of the abactinal and superomarginal plates carry spinelets and minute pedi-
cellariee. The latter are scattered and not very numerous. The spinelets are
of two sorts: nearly cylindrical very blunt ones about 50-75 mm. long and
the diameter one-third to one-half as much; and stout, markedly conically
capitate spinelets 1-1-5 mm. long and about one millimetre in diameter ; on
the marginal plates these heavy spinelets are conical rather than capitate.
There is never more than one of the stout spinelets on a plate, but one finds on
each side of it, usually, 1-3 of the more slender little spinelets ; occasionally
the stout spinelet is solitary. On the more conspicuous abactinal plates the
spinelets form transverse series, sometimes without any large one at the middle.
Madreporite 2 or 3mm. from the disk margin, small, scarcely 2 mm. in
diameter, elevated and surrounded by a circlet of 15-20 small spinelets.

Inferomarginal plates about 28, each with a stout spine, farely accom-
panied by a single much smaller one; distally the spines are low, blunt,
conically, but proximally they are flattened and the tips are chisel-like, or
deeply channelled on the upper side, or divided into 2 or 3 very short
branches. These proximal spines are about 2 mm. long with the tip a
millimetre or more wide. Actinolateral spines in two series continuing
nearly to tip of ray ; at base of arm there seem to be two series of actino-
lateral plates, but it is hard to determine how far out on the arm they are
continued ; the spines are similar to those on the inferomarginal plates, but
are smaller; there is seldom more than one to a plate. Adambulacral spines
about 2 mm. long, flat, blunt, and rounded, or a little widened at tip, in two
regular crowded series; the spines of the outer series are only a little, if
any, larger than those next the furrow. Oral plates, each with three spines
of which the distal is smallest and most like the adambulacral spines, while
the most proximal is 3 mm. or more long and yery flat and chisel-like.
Tube-feet crowded, in four series. Pedicellarize are very few on the actinal
surface; here and there are scattered minute forcipiform pedicellarize, and
near the base of the ray careful search reveals a few small forficiform pedi-
cellarize, with short, erect jaws and no stalk ; they are about as wide as long.

Colour, of dry specimen, dull yellowish-grey.

One specimen dredged at Garden Island, near Fremantle.

In view of the confusion already existing in the genus Uniophora, it may
seem of doubtful utility to describe a new species based on a single speci-
men, but after examining a considerable number of Uniophoras from South
Australia, representing at least two species and perhaps more, I am satisfied
that this West Australian form is at least distinct from any of those hitherto

246 MR. H. L. CLARK ON SOME

described. It is particularly characterised by very flat chisel-like infero-
marginal, actinal, and adumbulacral spines, but another feature in which it
differs from South Australian specimens is the absence of shagreen-like
“pebbled” areas on the lower part of the superomarginal plates. These are
quite conspicuous in most of the specimens from South Australia, but can
barely be distinguished on one or two plates of the specimen from Garden
Island. It should also be mentioned that the present specimen does not
seem to have any intermarginal plates, such as occur in some Uniophore.
None of the South Australian specimens have the grey colour of U. dyscrita,
but, owing to the poor condition of the type, this may be of no significance
at all.

OPHIUROIDEA.

HuRYALE ASPERA.

Lamarck, 1816, Anim. s. Vert. ii. p, 538.

A single specimen of this “ basket-fish,” 20 mm. across the disk, is labelled
as having been brought from Broome. It is very light-coloured, possibly
somewhat bleached, but is otherwise in good condition. The species was
previously known from north-western Australia only from two young speci-
mens taken by the ‘ Gazelle.’

OPHIACTIS SAVIGNYI.
Ophiolepis savignyt Miller & Troschel, 1842, Syst. Ast. p. 95.
Ophiactis savignyt Liungman, 1867, Ofv. Kong. Vet.-Akad. Forh. xxiii. p. 323.
A single specimen of this tropicopolitan species is in the collection from
Wooded Island. It has six arms and the disk is 3 mm. across.

OPHIOTHRIX MICHAELSENI.

Koehler, 1907, Fauna Siidwest-Australiens: Ophiuroidea, i. p. 250.

While I do not feel at all sure of the validity of this species, since the
‘section of the genus to which it belongs is in very great confusion, there is
a specimen of OjstheRadn at hand, taken at Garden Island near Fremantle,
which is almost certainly identical with Koehler’s specimens and may well
bear their name until the group is revised.. It is 13 mm. across the disk, and
although the arms are all broken, enough is left (65-115 mm.) to show that
they are very long. The disk is grey, the arm-spines pale brown, and the
upper side of the arms is indistinctly banded with dark and light slate-colour ;
there isa narrow, more or less interrupted, light line along the middle of the
upper-arm surface.

OPHIOTHRIX SPONGICOLA.
Stimpson, 1855, Proc. Acad. Nat. Sei. Philad. vii. p. 385.
This is one of the few characteristic species of the couihone coasts of
Australia which oceur at the Abrolhos. In the present collection there are

ECHINODERMS FROM WEST AUSTRALIA. 247

three specimens from ‘“ Garden Island, near Fremantle ; dredged,” one from
“off Fremantle,’ one from “ Wooded Isle,’ and four from “ Long Island.”
None of the specimens is large, the disk diameters ranging from 4 to
8 mm.

OPHIOTHRIX STELLIGERA,

Lyman, 1874, Bull. M. C. Z. iii. p. 287.

A single small specimen, from Garden Island, near Fremantle, has the disk
only 8 mm. across and the arms about 20 mm. long. Seven larger specimens,
from “ off Long Island,” have the disk 35-7 mm. across. Of these Abrolhos
‘specimens, four show spinelets among the thorny stumps on the disk. All
the specimens are much alike in their indistinctive, light greyish- or reddish-
brown coloration, and show the white stripe on the upper surface of the arms.
In the largest specimen a curious anomaly occurs in that on the proximal
part of the arm the white stripe is black. Distally there is a white stripe
delimited by a black line on each side, but as this passes towards the base of
the arm, the black encroaches more and more on the white, and finally
obliterates it altogether, the two black lines coalescing. Were the arms
broken off near the base, it would be hard to believe that the brittle-star was
really an example of stelligera.

OPHIONEREIS PORRECTA. ;

Lyman, 1860, Proc. Boston Soc. Nat. Hist. vii. p. 260.

The occurrence of this species in the Abrolhos is most interesting, as it
-does not seem to be common anywhere in the Hast Indies and is not yet
known from the mainland coast of Australia, although it is not rare at the
Murray Islands, in the Torres Strait region. There are five specimens at
hand from Wooded Island and two from Pelsart. One of the latter is only
4mm. across the disk and has the arms about 20 mm. long. The disk is
nearly pure white, save for the dark brown distal tips of the radial shields.
‘The other half dozen specimens are 8-12 mm. across the disk.

OPHIOCOMA BREVIPES var. VARIEGATA.

Ophiocoma variegata EK. A. Smith, 1876, Ann, Mag. Nat. Hist. (4) xvii. p. 39.
Ophiocoma brevipes var. variegata H. L. Clark, 1921, Echinoderms of Torres Strait,
p. 180.

It is rather remarkable that this widespread and variable Ophiocoma
should be the only representative of the family taken at the Abrolhos. It
was found both at Wooded Island (8 specimens) and at Long Island, Pelsart
Group (4 specimens). The specimens range in disk-diameter from 12 to
29mm. All are evidently to be referred to the variety variegata, but two
have the black-spotted disk of the form that has been called O. déderleinz and
-two have the reticulated disk of the so-called O. dentata. None is like
O. insularia or the typical O. brevipes.

248 MR. H. L. CLARK ON SOME
OPHIOPLOCUS IMBRICATUS. i
Ophiolepis imbricata Miller & Troschel, 1842, Syst. Ast. p. 93.
Ophioplocus imbricatus Lyman, 1861, Proc. Boston Soc. Nat. Hist. viii. p. 73, footnote.
There are two adults, light-coloured specimens of this species in the
collection, taken:on the reef at Long Island, Pelsart Group.

ECHINOIDEA. *

PRIONOCIDARIS BACULOSA var. ANNULIFERA,
Cidarites annulifera Lamarck, 1816, Anim. s. Vert. iii. p. 57.
Prionocidarts baculosa var. annulifera Mortensen, 1918, Kungl. Svenska Vetensk.
Handl. lviii. no. 9, p. 8.

A very fine specimen of this handsome sea-urchin was dredged among the
islands of the Wallaby Group. It is 49 mm. in horizontal diameter, and the
largest primary spines, which are ali more or less conspicuously banded with
reddish-purple and greenish-yellow, are about 57 mm. long and 5 mm. in
diameter near the base. The number and arrangement of the conspicuous.
red-purple spots on the collar of the primaries show great diversity.

PRIONOCIDARIS BISPINOSA var. CHINENSIS.
Doderlein, 1903, Jena Denkschr. viii. p. 697.

A small cidarid dredged off Long Island resembles so closely Déderlein’s
description and figure that I refer it to this variety with little hesitation, even
though the type-locality is so far distant from the Abrolhos. The specimen
is 35 mm. in diameter. Most of the primary spines are broken or missing,
but the unspotted red-brown collar is characteristic, and the distinctive
thorns are well-developed on the upper spines. The longest ones, 33 mm.
long, 2 mm. in diameter at the collar, cylindrical, and without thorns, are at.
the mid-zone. The secondary spines are brownish-red, those on the ambulacra
being slightly darker than those on the interambulacra and with a purplish

tinge.

CrNTRECHINUS SAVIGNYI.

Diadema savignyi Michelin, 1845, Rev. Mag. Zool. p. 15.
Centrechinus savignyt H. L. Clark, 1921, Echinoderm Fauna of Torres Strait, p. 145.

A specimen 85 mm. in diameter was taken in the shore-collecting at
es
Wooded Island.

CENTRECHINUS SETOSUS.
Echinometra setosa Leske, 1778, Add. ad. Klein, p. 36.
Centrechinus setosus Jackson, 1912, Phylogeny of Echini, p. 28.
A specimen 80 mm. in diameter was taken during the shore-collecting in
the Pelsart Group. The abactinal white spots are large and conspicuous, and
easily distinguish the species from the preceding.

s

ECHINODERMS FROM WEST AUSTRALIA. 249

CENTROSTEPHANUS TENUISPINUS.
H. L. Clark, 1914, Ree. W. Austral. Mus. i. p. 162.
There are three half-grown but typical examples of this western species,
one of which was taken in the lagoon at Pelsart Isle, while the other two
were dredged off Long Island.

AMBLYPNEUSTES PALLIDUS.
Echinus pallidus Lamarck, 1816, Anim. s. Vert, iii, p. 48.
Amblypneustes pallidus Valenciennes, 1846, Voy. ‘ Venus,’ Zoophytes, pl. ii. fig. 1.

A small Amblypneustes from Wooded Island, about 20 mm. in diameter
and nearly 20 mm. high, is best referred to this species, although the diamond-
shaped markings on the interambulacra are very faint. The test is faintly
pink, the secondary and miliary spines dirty whitish, and the primaries are
pale red. There are no spines on the anal system, and the tuberculation of
the test is that which is characteristic of A. pallidus.

SALMACIS VIRGULATA ALEXANDRI.
Salmacis alexandri Bell, 1884, ‘ Alert’ Rep. p. 118.
Salmacis virguiata alevandri Déderlein, 1914, Ech. Stidwest. Aust. p. 454.

This urchin has already been recorded from Geraldton and from Sharks
Bay, so the occurrence of two large specimens (74 and 79 mm. respectively
in horizontal diameter) from the Abrolhos is not surprising. They were
dredged off Long Island. One has a low ambitus and flat oval surface, though
the test is fairly high, while the other has a lower test and the ambitus much
nearer to the mid-zone. In the latter specimen the primary spines are green,
though the test is light purple, only the primaries near the peristome being
somewhat purple near the base. In the larger specimen the primaries, us
well as the test, are purple, but those below the ambitus are much longer and
more brightly coloured than those above, and all are tipped with white.

PskUDOBOLETIA INDIANA.

Toxopneustes indianus Michelin, 1862, Wch. et Stel.: Annéxe A, in Maillard’s Notes
sur Bourbon, p. 5.
Pseudoboletia indiana A. Agassiz, 1872, Rey. Ich. pt. 1, p. 153.

The occurrence of a Pseudoboletia on the western side of the Australian
continent is a discovery of more than ordinary interest, as the genus has not
hitherto been reported from Australia or the Torres Strait region. But
the present specimen, 50 mm. in diameter, with a dirty white coloration
lacking all indication of pink, is unmistakable. It was dredged off Long
Island.

TRIPNEUSTES GRATILLA.
Echinus gratilla Linné, 1758, Syst. Nat. ed. x. p. 664,
Tripneustes gratilla Lovén, 1887, Ech. Linn. p. 77.
Two specimens, 68 and 86 mm. in diameter, with dark test and white
LINN. JOURN.—ZOOLOGY, VOL. XXXV. 18

250 MR. H. L. CLARK ON SOME

spines, were taken at Pelsart Island, thus extending the known range of the
species on the western side of Australia considerably to the south. On the
mainland coast it is known only as far south as Sharks Bay.

HELIOCIDARIS ERYTHROGRAMMA.

Echinus erythrogrammus Valenciennes, 1846, Voy. ‘ Venus, Zoophytes, pl. vii. fig. 1.
Heliocidaris erythrogramma Agassiz & Desor, 1846, Ann. Sci. Nat. Zool. (8) vi. p. 871
There are half a dozen specimens of this well-known species of the southern
Australian coast, but all are young and the specific characters are not well
marked. Three from the lagoon, Pelsart Group, 25-30 mm. in diameter and
13-14 mm. high, have purplish tests and bronze-green spines; the largest
primaries are 15 mm. long, and not quite a millimetre in diameter at the base.
There is little doubt that these are normal H. erythrogramma. There is a
very similar specimen from Hast Wallaby Island. Two specimens from
Wooded Island have a different appearance, and yet differ more from each
other than either one does from the Pelsart specimens. One is about 28 mm.
in diameter, scarcely 13 mm. high, and has the tubercles much more con-
spicuous than in the others. The primary spines are purple, but the secon-
daries show an evident bronze-green coloration. Most of the primaries are
broken, but it is obvious that all were more or less stunted ; particularly all
the spines above the ambitus are relatively short and thick and blunt.
Apparently this specimen lived beneath a rock or among rocks where surf
or tidal currents were strong. The other Wooded Island specimen has the
test very light-coloured, with a green tinge abactinally, and the spines are
green tinged with purple at the tip, at least orally. The largest primaries
are 10-12 mm. long, with the diameter at base distinctly more than a
millimetre. This individual looks like a different species from the others, and
may be a young H. tuberculata, but it is highly improbable that that species
occurs in the Abrolhos. Probably this little green specimen was collected
on an eel-grass bottom in still water, such an environment as favours the

green colour and the better-developed spines.

ECHINOMETRA MATHAEI.

Echinus mathaei de Blainville, 1825, Dict. Sci. Nat. xxxvii. p. 94.
Echinometra mathaei de Blainyille, 1880, Dict. Sci. Nat. lx. p. 206.

There is a specimen 47 mm. long, 41 mm. wide, and 26 mm. high from
North Island, and the label says ‘“‘ Very common everywhere.” There are
also two small specimens (15-17 mm. long) from “ Long Island, shore.”
The abactinai tube-feet of the large specimen contain large numbers of the
triradiate spicules which Déderlein considers typical of his genus Mortensenia,
but I am not prepared to admit that their presence is even a good specific
character, and 1 think these specimens are more properly recorded as
EE. mathaei than as E. oblonga. But they are evidently identical with those
which Déderlein (1914, Fauna Siidwest-Austral. Bd. iv. Lfg. 12, p. 487)
identifies as Mortensenia oblonga, which were collected in Sharks Bay.

Clark

ECHINODERMS FROM

co),

Valera

=

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(a
re)

WEST AUSTRALIA

ECHINODERMS FROM WEST AUSTRALIA. 251

BRBHYNIA AUSTRALASLE. t
Spatangus australasie Leach, 1815, Zool. Misc. ii. p. 68.
Breynia australasie Gray, 1855, Cat. Ree. Ech. pt. 1, p. 46.
A single fine specimen from North {Island shows that the characteristic
species of Australia is not wanting at the Abrolhos.

EXCHINOCARDIUM CORDATUM.

Echinus cordatus Pennant, 1777, Brit. Zool. iv. p. 69.
Echinocardium cordatus Gray, 1848, Brit. Rad. p. 6.

Two specimens of this cosmopolitan species are in the collection, They
were dredged off Garden Island, near Fremantle. The larger is 32 mm. long,
22 mm. wide, and 13 mm. high.

EXPLANATION OF PLATE 13.
Fi

eo. 1. Petricia obesa, sp. n., aboral view, $rds nat, size.

2. 3 9p oral view, * .

8. Parasternia crassa (Gray) Fisher, oral view, nat. size,
4 P p aboral view, ,, ,,
5. Nectria macrobrachia, sp. n., aboral view, {ths nat, size,

” ” oral view, ” ”?

18*

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 253

Courtship Activities in the Red-throated Diver (Colymbus stellatus Pontopp.) :
together with a discussion of the Evolution of Courtship in- Birds”.
By Jutran 8. Huxuny, M.A. (Communicated by Prof. H. 8. Goop-
rico, M.A., F.R.S., Sec.L.8.)

(PuateEs 14, 15, and 4 Text-figures.)

~ [Read 2nd November, 1922.)

Tae following paper is a continuation of previous work on the sexual
relationships in species of birds in which both sexes possess bright colours
and also exhibit similar sexual ceremonies. The reader is referred to papers
on the Great Crested Grebe and on the Dabchick (Huxley, *14 and ’19).
It is unnecessary to recapitulate at any length, but certain general con-
clusions may be stated.

1. In most, possibly all monogamous birds, a pre-mating and post-mating
period may be distinguished. “Courtship” activities usually occur during
both of these periods; but the majority of the “courtship” actions which
are to be found described in the literature occur in the post-mating period,
and therefore cannot be operative in any true form of sexual selection as
imagined by Darwin. So far as pre-mating ceremonies occur and are
effective in the choice of mates, they can of course find place in a scheme of
true sexual selection.

2. In birds in which the sexes are quite or almost similar, and both
adorned with bright colours or special plumes, or other structures, which are
displayed or otherwise used in sexual ceremonies, both sexes play a quite or
nearly similar réle during “courtship”; very often ceremonies occur in
which both sexes simultaneously play a similar rdle ; for such ceremonies the
term “mutual” is employed.

3. These mutual ceremonies may be “self-exhausting ”—that is, may end
in the birds resuming the ordinary routine of life ; or they may be a means
of raising the emotional tone as a direct or indirect excitant to coition. In
the Grebe, where self-exhausting display ceremonies were the rule, special
pre-coition ceremonies were found, in which attitudes resembling those
employed at coition were adopted.

4. An association of various of the birds’ other activities with the sexual
ceremonies was often observed. The “handling” of nest-material is fre-
quently so associated, as has been noted by many previous observers ; but
actions like those of preening or of shaking the head, although originally
quite without sexual significance, may also be used as part of the raw
material of sexual ceremonies.

* No, 21 of the Results of the Oxford University Expedition to Spitsbergen in 1921.

254. MR. JULIAN S. HUXLEY ON

5. In many birds similar in the two sexes both male and female incubate.
In many but not all such species the relieving of one bird by the other on
the nest is the occasion for a special sexual ceremony, which in its turn may
or may not be used at other times. This ceremony of nest-relief appears to
be absent in the Grebe.

With this brief introduction, we may pass to the observations on the Red-
throated Diver. These were made while the writer was on the Oxford
University Expedition to Spitsbergen, 1921. Most were made on Prince
Charles Foreland, an island 60 miles long, off the west coast, on which a
small party of us camped for 11 days, from June 30 to July 10. Numerous
small freshwater pools are found here, as in many other places near the coast
in Spitsbergen ; and ona great many of these, at any rate on the north-
western and western coasts, Divers are to be found nesting. We were
fortunate in being encamped on the shores of Richard Lagoon, a sheet of
brackish water about a mile wide and 5 miles long, communicating with
the sea by one narrow opening. This lagoon was the scene of much of the
Divers’ courtship, both pre- and post-mating, though especially the former.
Much of the birds’ feeding was done on the open sea: but for rest and court-
ship the lagoon was chosen. Even after incubation had begun, the birds
appeared to come down to the lagoon when not sitting ; it was the scene of
veritable social gatherings of Divers, four or five being commonly on its waters
together, and eight having been seen there on one occasion. Sometimes they
would fish, progressing by means of long dives, as does the Crested Grebe ;
at other times they would stay quiet, resting or preening themselves. But
during the period of our stay we never observed two or more Divers on the
lagoon without some courtship action following within a comparatively short
time. The male and female are similar in appearance; the female is smaller
than the male, although the wing and beak measurements of species overlap
somewhat. However, the female appears to have a much less massive neck
than the male, and a separation of the sexes can usually be made with

reasonable probability.

Description of Chief Courtship Actions.

Tt will be best to begin with a description of the chief types of actions
seen, following this with their probable interpretation.

1. “ Plesiosaur race.” This ceremony was so christened because the
attitude adopted by the birds in its performance made them resemble miniature
Plesiosaurs half out of water (fig. 1). Usually two or three birds took part
in it, sometimes four. The birds depressed the hinder half of the body below
the water ; the body was held at an angle so that the breast and shoulders
were stuck out ; the neck was stuck upwards and forwards in a stiff position,
the head and beak inclined somewhat forward, again somewhat stiffly. In
this attitude the birds swam through the water, accompanying one another.

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 255

Their passage through the water seemed to be accompanied by a good deal of
resistance, since a wave showed at their breasts. They would go for some
distance in this way; then would often turn and continue the process in the
opposite direction. The bird which had led in one direction, it seemed, might
be replaced by another as leader after turning. The whole ceremony did not
give the idea of a true pursuit, such as is seen among many birds, but more
of a race, or of something still more ceremonial and formal.

The most usual arrangement is for one bird to lead, the other one or two
being half a length or less behind. Various modifications of the pose are
seen. Sometimes the body is at a very low angle, the neck little arched, the
head and bill almost horizontal ; at others the tail is more submerged, the
breast more elevated, the neck much arched, and the head and bill pointed
‘downwards; it isin this pose that the likeness to a Plesiosaur is most marked.
Occasionally almost the whole of the body is submerged. As to the general
appearance, I quote from my notes :—“ There is a strange stream-line effect
about the creatures. They must be paddling with great energy to keep up

Fie. 1.

é Red-throated Diver: Plesiosaurus-race ceremony.
the speed.” ‘There is often a snaky look about them when in the semi-
vertical attitude.” Altogether the effect as of tension, of emotional ritual, so
familiar to all those who have watched birds during courtship, is marked.

A still further modification of the pose is sometimes seen in what may be
called the “wing Plesiosaur attitude.’ In this the wings are stretched out
nearly at right angles to the body, perhaps 2/3 spread, the point inclined a
little backwards, and somewhat drooped, so that the upper surface is plainly
visible. This might be adopted by all the birds in a “trace,” or by the
“‘pursuer”’ alone in a “race” of two birds only. The same attitudes and
relative positions were maintained by one such pair of birds for over
50 yards.

Occasionally birds might be seen in a still more nearly vertical attitude.
When this was so they did not progress fast.

A peculiar note invariably accompanied this ceremony. I find it described
in my notes as “a growl with a bubble in it” or a “rolling growl.” This is
repeated a number of times, and appears to be used just before or at the
beginning of the ceremony proper. This note did not seem to be used at any
other time, except in the “snake-ceremony ” (see p. 257).

256 MR. JULIAN S. HUXLEY ON

From the beginning we noticed that birds engaged in this and in other
sexual ceremonies, or more often when under the influence of sexual excite-
ment, but not actually engaged in a ceremony, often engaged in curious and
apparently meaningless actions. ‘The most frequent was the dipping of the
beak in the water (fig. 2), often repeated several times within a few minutes ;
a variation of this consisted in “looking into the water ”’—in other words, the
submerging of the beak and fore-part of the head for some little time ; finally,
shaking of the head was observed, though much more rarely than in the Grebe.

Fie 2.

Red-throated Diver: snake-ceremony.

It became quite certain, as we continued our observations, that these actions, °
or at least the two former, were definitely associated with sexual excitement.
Like the head-shaking and ‘“‘habit-preening” of the Grebe (Huxley, *15),
they appear to represent actions normally performed with some other
function, or, if functionless, with non-sexual associations, which have become,
through some peculiarity of the psychological mechanism, secondarily
associated with sexual excitement, and used as physical expression of sexual
emotion. This appears also to be the case with a special form of dive which

I call the “splash-dive.” In this, instead of submerging quietly, almost

Fie. 3.

Red-throated Diver: emergence ceremony.

without a ripple, as is done when the birds are diving for food or to escape
danger, the bird gives a sharp kick with the legs as it goes under, sending a
shower of spray into the air; furthermore, the dive is fora very short distance,
the bird emerging generally about 5 or 6 yards away. This, too, is associated
with apparently all forms of sexual excitement, and otherwise is used only
under the influence of the emotion of anger (see Van Oordt and Huxley,
722). Here we have apparently a normal form of activity, which is used in a
modified form when associated with sexual emotion.

—

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 257

2. A very remarkable ceremony, unfortunately only observed clearly on
one or two occasions, was the dive followed by vertical emergence((figs. 3 & 4).
In the best-observed case two birds out of a party of three were close
together. Suddenly one—almost certainly a female—dived (with a splash).
The other—almost certainly a male—immediately spread its wings and half
flew, half sputtered along the surface of the water for 5 or 6 yards. The bird
which had dived then came up out of the water close to the other, in an
almost vertical position (fig. 3). The neck was held so as to continue the line
of the body; the head bent down so that the bill made a small angle with the
breast. The emergence was fairly slow. It stayed in this position for perhaps
5 seconds, then settled down on the water. Here we have a ceremony which
is extremely similar to one which I have described in the Crested Grebe,
although not quite so elaborate. Its chief feature of interest is that, although
both birds play active parts in it (the flying-off and waiting of the non-diving
bird is an active part, which culminates in the Grebe with a special display),
yet the parts are different. If we may judge from the Grebe, either part

Fre 4.

Red-throated Diver: another sketch of an emergence ceremony.

may at different times be played by a bird of either sex. This is rendered
more probable by the fact that the more active réle on this occasion was
apparently taken by a female.

3. A ceremony in many ways resembling the Plesiosaurus race is also
common, but apparently only among birds which have mated up. It is
always accompanied by a ery which is the same as, or extremely similar to,
the “roll-growl” already described. The two birds swim, one leading, for
some distance with their necks arched so that the tip of the bill is submerged.
The bill is also held open, in order to give the cry. The bodies, however, are
not submerged posteriorly, but kept in normal swimming position, and there
is not a marked wave thrown off the breast, nor does motion appear to be at
all violent. Asin the Plesiosaur race, the birds may turn, and the former
leader become the second. This ceremony might be performed on the small
nesting-pools or on the lagoon. Owing to the snaky look of the neck when
in this attitude, I shall call this the snake-ceremony (fig. 2).

There remain peculiar actions associated with coition ; these, however, may
best be described later, in connection with the account of a particular pair.

258 MR. JULIAN S. HUXLEY ON

Activities of a Particular Mated Pair.

T will now proceed to describe in some detail the actions of one pair which
I had under close observation for a few days, since I believe that in this way
a fuller insight is gained by the reader (as it certainly is by the observer at
the time) into the psychology of the birds.

On a small tarn about a mile from camp a single Diver was observed on:
June 30. The tarn was then only partially thawed, and nest-building was
out of the question. It would appear, however, that the birds had already
“staked out a claim” to this special nesting territory, as ig known to be done
by many if not all monogamous birds before actual nest-building begins (see.
HK. Howard, 20). A single bird was again observed here on July 4; it was
restless and remained on the tarn for a long time ; nest-building had almost
certainly begun, as the event showed. On July 5 both birds of the pair
were seen on the tarn, and one visited the spot where the nest afterwards.
proved to be. It only stayed half a minute or less, and then returned to the
water. The birds were continually giving a mewing call, very similar to:
that given by solitary birds on the water, or by sitting birds when man or a
Skua threatens danger. The callas given on this occasion, however, I record
as “louder, longer, and more emotional than when given by a solitary bird.”
Usually the two birds would howl almost, but not quite, simultaneously.
There was certainly some sexual significance attaching to this ‘“duet-
howling.”

At one moment the typical “roll-growl” was given. This I take as
proof that a snake-ceremony occurred, although the birds were unfortunately
ifivisible below the bigh bank. On my approach to pitch my observation
tent, the birds rose together and made off. I take it as certain that no egg
was as yet laid, since, once this has happened, the nest is sat on almost con-
tinuously to prevent Skuas stealing the egg.

Next day (July 6) I spent about 9 hours at the tarn. It was noticed that
whenever the howl or mewing note was given, the neck was stretched out
nearly or quite parallel with the water. This was confirmed on many sub-
sequent occasions. The action seems to be inseparable from the particular
note. It is probable that it renders the bird less conspicuous (see Van Oordt
and Huxley, loc. cit.). The note was given repeatedly by the solitary bird
(female) that was there at intervals from noon to 3 P.M. Occasionally the
bill would be dipped as in sexual excitement. At one time she indulged in
what were obviously practice dives, just in and out, emerging two to four
lengths away. This was repeated five times in quick succession.

Shortly after, the male arrived. The female speedily joined him, and the
pair swam towards me, the female leading, with bill slightly open. When
close to the bank, the female suddenly performed a remarkable action,
obyiously of a sexually stimulating nature. She stretched her neck forwards
at an angle of perliaps 30° with the horizontal, the head and beak also.

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 259

pointing up, but at a less angle. The neck itself was straight and rigid. In
this pose she swam close up to the male, crossing his bows, so to speak.
Whatever its significance, the performance was exciting to watch. There
was a tenseness about the bird’s attitude, a rigidity, which has been com-
mented on by other writers, notably by H. Selous, in connection with various
sexual ceremonies in other birds (Selous, 701-02, 05a, 056). I can best
describe the impression it made on me by saying that it was like that
apparently produced by certain sexual dances of savage tribes—the whole
thing fraught with the significance of sexual emotion, and mysterious in the
sense of being thus emotionally charged far beyond the level of ordinary life,
but completely natural and without restraint. It is fairly clear that, even in
animals, the emotional tension during sexual excitement is far higher than at
almost any other time, and that the impression given to the observer is,
therefore, not wholly a subjective one.

On this occasion the male was absolutely unresponsive to the female’s
“stimulating” action. The pair went off together ; after some time the
female started to go by a long route across dry land to the nest, but soon
gave it up. The pair then swam, the female again leading, to a little bar
covered with moss on the far shore. This bar J afterwards examined, and
found that on it there was a rudimentary nest, built almost entirely of the
moss which was here abundant, both on the bar and under the surface of the
shallow water near by. Another such “nest” was found on the shore of
a second tarn on which a second pair was breeding. Very similar structures
are built by the Crested Grebe ; and in both eases their function appears to
be the :ame—they are the places on (or near) which the act of coition oceurs-
Just before arriving here both birds simultaneously “looked into the water ”
(vide supra) for about 20 seconds ; there was alsoa good deal of beak-dipping
by both male and female. On arriving at the bar the female turned and
gave an energetic splash-dive. The male responded by a less energetic
splash-dive. The male then crawled out on the mossy bar, onto what I later
found to be the pairing-nest; there he stood upright, stamped alternately with
his two feet several times, and then sank down as if brooding. He plucked
small fragments of moss in his beak, and apparently placed them round
himself as if adding to the nest. The female meanwhile was swimming close
to the bar, in water scarcely deeper than her draught, giving repeated splash-
dives. Her tail (which of course was very short, as in all Divers and Grebes)
was repeatedly waggled up and down. The association of this motion with
copulation in birds is obvious and well-known. However, she did not land,
but swam across to the right and back, giving several more splash-dives ;
the male got up and scrambled back into the water. The pair then
went to the nearest point to the true nest, both diving twice en route (not
splash-dives) ; the female led, at a fast pace. After one short abortive
excursion overland towards the nest, followed by swimming off the shore, she

260 MR. JULIAN S. HUXLEY ON

went right up to the nest and settled down on it. The male dozed on the
open water with his head under his wing. It was remarkable that since the
arrival of the male not a single sound had been uttered by either bird.

About half an hour later the male flew off. The female continued to
brood. At one time a Richardson’s Skua came down close to the nest. This
was the signal for the Diver to give a series of mewing calls, with neck out-
stretched in the direction of the enemy. (Occasionally the neck may even be
directed slightly downwards.) About 7 p.m. I went off to carry out some other
work. On returning, she was on the tarn. I visited the nest, and found it
a low heap of mud with a shallow depression, and a moderate amount of
moss round the rim. There were also several pieces of slate on it, but it was
not clear whether their presence was not purely accidental.

On my returning to the hiding-tent, the bird soon came back to the nest.
At 7.45 p.m. the male came back to the tarn. The female continued to sit
without making any sign for a few minutes, but then suddenly left the nest
and joined the male. Both appeared excited, and “looked into the water ”
several times in quick succession. Immediately afterwards they both assumed
a very poor “snake attitude,” with the neck only slightly arched. This
lasted but a short time; the female then gave five splash-dives in rapid
succession. The pair then indulged in a snake-ceremony, swimming towards
me almost parallel, neither markedly leading. It was, however, a poor affair
compared with those at full intensity of emotion. They then swam close up to
the moss-bank on which was the pairing-nest, the female leading ; she swam
close in and was obviously excited, but the male did not follow. After both
had had a spell of rest on the open water, the male looked up and soon
started a “roll-erow!l ” (the first sound giyen by either bird since his arrival).
The female soon joined in, and a typical snake-ceremony followed. They
then returned, the female leading, to the moss-bank. The male scrambled
out again on to the bank, but at the other end from the nest. He trampled
as before, but less often, and settled down, but did not pluck any moss. The
female, however, after first pecking at the moss on the bank, turned, put
her head under waier, and proceeded to pull large pieces of moss up from
the bottom. These she then jerked backwards over her shoulder, dropping
them carelessly into the water and showing no further interest in them.
This she did eight or nine times in quick succession. After pecking up
moss, both birds usually gave a sharp, emphatic shake of the head.

The male then got off, made for the shore near the true nest, and sat
there some time on the mud. Then both came out onto open water, and
a good deal of mewing in duet went on.

The female then led the way to the moss-bank, where (I quote from my
notes) “she beached herself, stern just in water, breast on land, head
down. The male followed, mounted onto her, standing neariy upright,
and copulated or attempted to copulate (it did not look as if he was

COURTSHIP ACTIVITIES IN- THE RED-THROATED DIVER. 261

successful, but this is hard to judge). He then walked up her body and off
her right shoulder (in this whole action behaving very much like the Crested
Grebe).”

The female remained perfectly motionless for one or two minutes, in the
same extended position; the male, after sitting on the bank for a slightly
less time, took to the water. The female than sat up, plucked up pieces of
moss as before, but fewer and with less energy, and then joined her mate on
the water. After swimming, with the male leading, they turned; the female
gave a short flight of a few yards, the male swimming rapidly after her ; the
male then gave a similar short flight. Both shook their heads emphatically
several times (compare the Crested Grebe). After a short time the female
got up, followed at once by the male, and both flew off westwards and were
lost to sight. This abandonment of the nest with an ege in it was remark-
able ; possibly they were more upset by the presence of the tent than they
appeared to be. In any case, apart from the presence of a pair of birds
which may or may not have been the same, for a short time next day, the
tarn was abandoned, and the egg fell a prey to Skuas within 24 hours.

Another nest was discovered on a neighbouring tarn on July 7. This
already contained two eggs, although this tarn had been still later in thawing
than the other, and had ice extending for some distance in from all its
margins on July 2. This tarn was within a short distance of camp, and
the “roll-growl,” signifying the mutual “snake-ceremony,” was frequently
heard from it (e. g. 5.30 a.m. July 9). This shows that mutual “courtship ”
continues, as in the Grebe, throughout the incubation period. On investi-
gating the banks, a ‘‘ cock’s nest,” similar to the one already described but
rather more rudimentary, was discovered, also on a rather mossy part of the
shore (as a matter of fact, the only mossy part in this tarn); doubtless it, too,
subserved the function of coition.

The association of the mossy bar with the rudimentary nest is interesting.
The true nest often contains some moss; and the behaviour of the female,
above recorded, in plucking up moss from the bank shows that in the Diver,
as in many other birds, the handling of nest-material has a sexually
emotional association. It is presumably this emotional association which
prompts the choice of a mossy bank as the site of the pairing-nest. There is,
however, far less nest-material employed than by the Grebes; and
accordingly nest-material plays a smaller part in the Divers’ sexual
ceremonies.

On my arriving at the hiding-tent on the evening of July 9, the female,
who was sitting, was rather suspicious, left the nest, and swam about
restlessly on the pool for some time. Although alone, she frequently dipped
her head half into the water, as previously deseribed, for 5 to 15 seconds. Thus
this habit, which is undoubtedly the main “associational”? habit of this
species, may be associated with other emotions than that of sexual excitement—

262 MR. JULIAN S. HUXLEY ON

in this case, with that of anxiety. It was here evident that it was per-
formed especially when the bird was particularly nervous; for instance,
whenever she approached the nest+region. This she did repeatedly, sheering
off six or eight times before finally plucking up courage to ascend the nest.
‘The “ looking in the water ” alternated with a curious action of the wings,
first one and then the other being raised very slightly. This was not
seen at any other time. The male arrived on the tarn about 6 p.m. The
female flew along the water to his neighbourhood, uttering a repeated cooing
note. Both made towards the nest, the female leading, swimming very low
in the water and cooing; she made one short dive. Both then cooed in
unison. No ceremony followed, however, and the female soon went on to
the nest. The male shortly afterwards flew off, but returned within half an
hour. No excitement occurred within the next hour. After this he several
times came up close to the nest, and gave the cooing call, which was some-
times given by the female in response. Soon after this he flew off. Here we
have evidence of sexual ceremonies in the morning, but a very “ dull evening.”
Probably the mutual ceremonies are practised less and less as incubation
proceeds.

Next day (the 10th) I was in the tent from 2.45 to 5 p.m. The male did
not appear at all. The tent was now pitched at 6 yards from the nest. The
bird was markedly less nervous than the day before, but every time that she
approached the nest from the open water, she dipped her bill several times
before venturing on the difficult overland traverse. This corroborates the
interpretation of the “ looking into the water”? as an expression of emotion.
On the 11th, the last day we spent on the Foreland, I spent 12.30 to 2 a.m.
at this tarn. The male again did not appear. The female appeared to be
sitting whenever I was watching ; but there is no doubt that the male is to
be found sitting in this species (Witherby, 00, &e.). During this period
the three birds previously referred to (p. 257) flew down and, after circling
round, alighted on the tarn. All were very alert in expression, and were
repeatedly dipping their beaks into the water—?. e. under the influence
of emotional excitement. From>their sizes, I should say there were two
females and one male. After a few minutes they rose, circled round again,
and again settled. One separated from the other two. The (apparent) male of
these two arched its neck very slightly, as if going into the snake position ;
on this, the other bird dived and emerged in the vertical “ Penguin ” attitude,
as already described. The immediate fluttering over the water by the
remaining bird on the female’s diving indicates that this is a regularly-
‘occurring ceremony and that the emergence was expected ; the same is true
in the Grebe.

The ‘‘ Penguin” bird dived again, with a splash; but before it had
emerged, the other two had both, for some unknown reason, flown off. The
diving bird emerged in an abortive “ Penguin” attitude, looked round to see

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 263

where the other was, and then it, too, rose and followed the others. They all
three made off in wide sweeps, and were finally lost to view some miles away.

I take it that these three birds were not yet paired up for the season.
Possibly here, as in the Crested Grebe (Pike, 13), some birds do not nest, but
continue “ courtship ” actions throughout the season. The long flight to the
tarn and away again so soon after is of interest; it may have been a pursuit ;
but, if so, it is interesting that the bird which took the chief initiative in
courtship on the tarn was probably a female.

Other Observations on the Red-throated Diver's Courtship.

Before proceeding further, it will be as well to mention the actions
observed by othersin the same species. HH. Selous (12) made observations on
a family in Scotland, during July, at a time when the young were hatched,
and-were being looked after by both parents. Although he was watching
steadily for over. three weeks, he only saw “courtship” behaviour on three
occasions. This in itself is of interest, as showing that solong as the breeding-
season lasts—z.e. at any rate so long as the instinct for caring for the young
persists, and possibly until the autumn moult—the possibility of sexual
ceremonies appearing in special circumstances will also persist. The obser-
vations of Pike (713) are of interest in this connection. He states that a large
percentage of tlle Crested Grebes on a certain breeding area, although mated
up, failed, for some unknown reason, to build or lay in the season when he
was there: these birds, and apparently none of the other pairs, continued
their well-known mutual courtship late into the season (7. e. after the normal
time for incubation was over). Facts bearing on this problem are well
known in other species ; for instance, in many birds, the final liberation of
the parents from feeding the young, when these are old enough to look after
themselves, is often marked by a recrudescence of various sexual activities.
These are similar to the truly epigamic activities of the early part of the
season, though they now, of course, have no function. Similarly, in birds
which rear two or more broods, there is usually a recrudescence of epigamic
ceremonies in the interval between broods. Looking at the matter from an
opposite angle, we find that the song of many birds, e.g. the Nightingale,
ceases immediately the young are hatched—i. e. as soon as the male embarks
upon the new duty of helping feed the young. Mr. Eliot Howard informs
me that in birds such as the Buntings, in which the males occupy territory
long before the females arrive, the volume of song in the first period is
greater than in the period immediately after the arrival of the females. This
has been corroborated for other species, e.g. the Blackbird (Dewar, ’20).

In a yery similar way, the female, on reaching a certain level of endocrine
stimulation, will permit and indeed solicit coition ; her emotional activity
will also find relief or expression in the building of the nest. Coition brings
about egg-laying, and nest-building brings about the finishing of the nest;

264. MR. JULIAN S. HUXLEY ON

a new period is thus introduced in which new objects are introduced to the
bird. Her excitation, so far as it is determined internally, remains at the
same level, but her activities are determined also by the objects presented,
and she now begins to incubate. In the absence of nest and eggs she would
revert to the behaviour of the preceding stage—coition and nest-building
once more, as is shown by the behaviour of birds whose nest and eggs are
destroyed after incubation has begun; but in the absence of the internal
stimulation by the secretion of the gonad, she would neither brood, nor be
eager for coition, nor build a nest.

The annual cycle of a bird is therefore divisible into a series of periods,
determined (a) by the state of internal stimulation, (b) by external objects—
the presence or absence of mate, eggs, young, &e. These latter will
determine the division of the breeding-period into a number of sub-periods.
In the third place, (c), an influence on bebaviour is exerted by general
environment—e. g. by the state of the weather, the abundance of food, &c.

So long, therefore, as the internal stimulation persists at the same intensity,
the sub-periods will be mainly determined by the external objects presented
to the bird. But, while any one sub-period will be characterized by one
predominant type of behaviour, the other types of behaviour will be latent
in the bird all the time, and may emerge under special circumstances,
though rarely, so long as the breeding-season (7. e. the secretion of the gonad)
continues.

To return to Selous’s observations on the Diver. The male had just fed one
of the chicks ; he then approached the part of the pool where the female and
other chick were swimming ; the female also swam towards him. On the
two birds coming to within a certain distance, both made two short
flights towards each other over the surface of the water; each time the
flight ended by the birds “as it were walking on the water, as a Penguin
walks on the land, bolt upright, with the whole of the white expanse of the
under surface....showing.” It is like “a Penguin running a few steps,
and then, for an appreciable period, standing upon the water, the naked legs
and feet just hidden by it.” ‘On another occasion, the male of another pair
was seen to alight on the water on its return from the sea, and to end its
alighting in the following manner: it “ footed it, for a little, towards another
one, presumably its mate, .... maintaining with the help of its wings the
upright Penguin-like attitude.”

On the third occasion, the female had started to give a special note with
head stretched forward ; she then began advancing towards the male in a
series of little plunges. This stimulated the male to do the same. Hach time
the cry was given, the neck was arched and held rigid. They eventually met,
and then quieted down.

It is interesting to note that none of the actions recorded by Selous are
identical with those seen by me. This is probably due, at least in part, to.

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 265

the different period at the time of observation. On the other hand, they are
built up out of the same elements.

Bent (19) says of all the species of Diver, as well as of numerous other
Pygopodes, that they migrate in pairs, or at least arrive at the breeding
localities in pairs, and select the nest-site as soon as possible. He believes
they pair for life. In all species it appears probable that both sexes incubate
(although from my personal observations it appears that one bird, pre-
sumably the female, takes much longer spells than the other ; and see Turner,
113):

Courtney, quoted by Chapman (°22), gives a beautiful drawing of a common
courtship ceremony of the North American Loon (Colymbus ¢mmer), in which
one bird rises into the erect position, and runs along the surface of the water
in this attitude for some way near another.

Bent (loc. cit.) refers to the “ maniacal ” screeching laughter of this species
in the breeding-season.

Seidlitz (13), writing of the Black-throated Diver, says that each pair has
a well-marked and large territory (Rewer). He states that there are as
many unpaired as paired birds ; these are birds of one, and possibly of two
years of age. He does not mention whether these ever indulge in “ court-
ship’’ ceremonies. This would be a point of considerable interest.

Bahr (07) mentions the dipping of the beak into the water when the bird
is nervous and afraid to return to the nest. He also, on one occasion only,
saw a very remarkable performance by a single Red-throated Diver. The
bird ascended to a great height, and then descended headlong, twisting and
turning as it did so, the air rushing through the feathers making a noise
“like a train.” Such descents are, of course, well known in many birds,
e.g. Rook. They are common in Louisiana Heron, Little Blue Heron, and
Snowy Egret on returning to their rookery, and I have on one occasion seen
a somewhat similar performance by our common British Heron. It is
probable that these actions should be classed as play, and not as courtship.

Hatch, quoted by Bent ((oc. cit.), states that C. immer has a remarkable
family habit. When the young are (apparently) well-grown, they sleep with
their parents on the pool. In the early morning, before sunrise, the male
parent gives a special call; all draw near, and, after a little manceuvring, start
to run over the water side by side in a line “‘at incredible speed,” with the
wings about a quarter-extended, the whole motive power being derived from
the feet. They run thus for about 400 yards, then wheel, and return to the
starting point. This is repeated several times and suddenly stops. It would
be interesting to have further observations on this extraordinary ceremony.

INTERPRETATION.
It will be seen from the above that (as in many, and indeed in most, of
even our commoner birds) much remains to be known about the life-history
LINN, JOURN.—ZOOLOGY, VOL. XXXV, 19

266 MR. JULIAN S$. HUXLEY ON

and especially the relations of the sexes. Those who have opportunity,
patience, and a good glass, and are willing to take full notes, will find that
steady observation of almost any species of bird at the beginning of the
breeding-season, particularly if a single pair can be followed throughout,
will bring results which may be of very considerable interest, both from the
standpoint of pure biology and also from that of comparative psychology, as
well as being in itself a very fascinating occupation.

The following paragraphs must therefore be taken only as a provisional
interpretation of the observations which have so far been recorded.

The Red-throated Diver is monogamous ; possibly it pairs for life. If it
pairs for life, it follows that the pairs will have some form of association even
through the winter. In any case, however, a change occurs in spring.
Hither the loose association of male and female is replaced by a much more
intimate one, or else the birds have been really unmated during the winter,
and now do separate out into pairs. This must, of course, be the case with
birds which have not previously mated. In any event, the change is definite,
and warrants the term “ pairing-up” which is usually given to it. This
change is presumably associated with a change in the reproductive organs,
which start to secrete their specific hormones.

The pairing-up appears, at least with some individuals, to occur in the
winter haunts. When so, it is followed, probably shortly, by migration, in
pairs, to the breeding-grounds, where each pair as soon as possible stakes out
a territory for itself. The present species appears to breed on small pieces
of water, and probably the territory is, in most instances, coterminous with
the nesting-pool.

The fact that the pair migrate together and choose the nest-site and
territory together is in strong contrast to the occurrences in such Passeres
that have been fully investigated, such as the Sylviide and the Emberizinz,
in which the males alone stake out territory (Howard, ’20), and, if migration
occurs, migrate before the females. However, it seems to occur in many
Pygopodes (Bent, op. cit.); and in Herons (unpublished observations of my
own), although migration is in flocks, the choosing of the nest-site is under-
taken by the pair together.

Although from the evidence of others it appears certain that pairing-up
may occur before migration, yet my own observations appear to point to this
not being universal. The fact that the “ Plesiosaur” ceremony occurred
frequently during the first few days of our stay, but later was absent or very
rare, as well as the fact that the simplest interpretation of it was that it was
concerned with the “choosing” of mates, indicates that probably a good
many birds arrive unmated on the breeding-grounds. Possibly (I throw this
out merely as a suzgestion) the birds which have nested before, pair up in
winter quarters, while those which are about to nest for the first time do not
do so until they arriye on the breeding-grounds,

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 267

The “ Plesiosaur” or “race” ceremonies, then, appear to be competitions
fora mate. It is of interest to find that when three birds are participating
in these ceremonies, their respective sizes indicate that they may be either
two males and a female or two females anda male. This is in accordance
with the rest of the tacts observed about courtship in this species—_the two
sexes appear to play almost identical (or interchangeable) roles.

The only occasions when I observed the dive followed by the “ Penguin-
like” emergence, also seemed to be in this pre-mating period. The erect
“Penguin” attitude, however, as Selous’ observations show, is not only con-
fined to this period, although he does not record diving followed by this attitude.

There is, however, another possible interpretation of the participation of
three and four birds in these ceremonies, which may be best understood after
a brief digression. In the Buntings, as Mr. E. Howard kindly informs me,
during the period while male and female are both present on the territory,
but coition and nest-building have not yet begun, every day begins in the
same identical way. The male wakes first: later, the female emerges too
from her roosting-place : after a short time the male flies at her, eager for
coition. She however flies off, and there follows what Howard ealls the

“sexual fight,” when the female, dodging and twisting, is pursued closely in
her rapid flight by the male. Hventually the birds seem to get exhausted,
and alight. Such pursuit flights are of course well known among many
species. What is of special interest, however, is the fact that once the
flight has started, it is a source of great excitement to other males in the
vicinity, and, although these males may have been long in occupation of
territory, and mated, yet one or more will usually join the rightful male in
the pursuit. In passing, it should be observed that this instinct of other males —
to join in the excitement of these pursuits appears to be at the bottom of
those undignified skirmishes of the House-Sparrow, in which a single
unwilling female is mobbed by a number of males ; and see also Bent, op. cit.,
pp. 186-191, where the Californian Guillemot is described as becoming very
much excited at the sight of an act of coition by another pair.

It is conceivable that the Plesiosaur race of the Diver may be a mutual
‘‘nost-mating ” ceremony, essentially identical with the “ snake-ceremony ”
described as occurring later in the birds’ history, but, since it takes place
before the nest is built and other activities share the birds’ mind, differing in
its greater intensity. If so, then the participation of three or four birds in
the ceremony may be due to others joining in under the influence of
excitement—others which are either wholly unmated, or else are mated but
temporarily without their mates. Since there were almost always several
birds on the lagoon, while the nesting-pools hardly ever had more than a
single pair on them, and since further the greater level of emotional excite-
ment in the Plesiosaur ceremony is unmistakable, it would be quite
intelligible that the threes and fours were only seen on the lagoon, and only

performing in the Plesiosaur, not in the ‘‘ snake ”’ ceremony.
Lg)

265 MR. JULIAN S.- HUXLEY ON

For the present, however, I prefer to think of these as ceremonies
actually connected with mating-up. If so, they are interesting, as they
would be, so far as I know, the first pre-mating ceremonies described which
were “mutual,” in that both sexes played similar réles. Further observations
are urgently needed, not only on this particular point, but on the whole
problem of pre-mating ceremonies, especially in species with mutual post-
mating courtships.

In any case, the post-mating ceremonies are essentially mutual. The
commonest is the “ snake” ceremony, the incentive to which may, to judge
from my observations, be given by either sex. This ceremony, like the
pre-mating ones, may often be “self-exhausting ” in the sense that, as in the
Orested Grebe, it need by no means always or usually lead to coition, but
may be followed by a relapse into ordinary non-sexual routine. Here,
however, I should like to put on record that further observation on this
species and on Grebes has conyinced me that such ceremonies may often
have a stimulative effect, although that effect may be gradual and in a sense
cumulative. It is very common in the Grebe to see a pair repeat a series of
mutual ceremonies at frequent intervals, and finally go off in the direction of
the nest and pairing-platform ; this has also been observed in the same species
by Selous, and clearly occurs to some extent in the Red-throated Diver.
That in certain circuntstances the ceremony may be completely self-
exhausting is perfectly intelligible: like the song of most Passeres, the
performance of the ceremonies is definitely pleasurable in certain states of
emotional tension. Physiologically, it is then performed “ for its own sake” ;
it may or may not, according to various circumstances, have the further
biological function of raising the emotion to a pitch at which coition is
desired.

On the other hand, there is no evidence that in the Divers there exist special
attitudes and ceremonies especially used as a symbol of readiness to pair, as
in the Grebe. The approach to the pairing-platform is in itself a symbol of
some degree of readiness to pair. The emotion associated with this may
apparently again be dissipated by excited actions, such as the pulling of weed
from the bottom, and by the ascent of the pairing-platform without actual
coition.

The situation of the pairing-platform in a mossy situation is of interest.
Moss is often but apparently by no means always used as nest-material, and
in any case always in very small quantities ; the birds can never handle it in
such large amounts during nest-building as they do in the vicinity of the
pairing-platform. Two hypotheses might be considered in regard to this.
In the first place, it might be a reminiscence, the birds having in the past
employed more moss in nest-construction than at present. Or it might
represent a specialization ; once the divergence between nest and pairing-
platform had arisen, and the emotions connected with coition had become
attached exclusively to the latter, new ceremonies, based on the handling

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 269

of nest-material in the first instance, might have been developed in
association with the pairing-platform to a much greater pitch than they
had ever been formerly. If this is so, and offhand I am inclined to
accept it as the more probable, there has been an evolution of the pairing-
platform and the ceremonies connected with it, which is in many ways
similar to that of the Bower-birds’ bower.

Personally I have not seen nearly so many post-mating ceremonies in
this species as are to be observed in the Crested Grebe. This is undoubtedly
due in part to the paucity of my observations. The work of Selous shows
that “Penguin” ceremonies do occur in this period, and, it is interesting
to note, continue, though rarely, after the young are hatched. Thus the
physiological state (presumably dependent on gonadial secretion) during
which epigamic ceremonies can be performed extends, apparently in all
birds, up to the very end of the breeding-season ; it does not then issue so
frequently in ‘ courtship,” because incubation and the care of the young
afford other outlets for physical energy and emotional tension.

On the other hand, only three, or possibly four types of epigamie ceremony,
whether pre- or post-mating, have been observed at all in this species,
whereas in the Grebe six have been seen; and the greater abundance in
the latter is probably not only apparent but real, and associated with the
greater development of epigamic ornament in the Grebe.

Incubation appears to be undertaken mainly by one bird, the smaller of the
pair; but there are numerous records of males also incubating, and it is
probable that in the Divers, as in other forms with mutual courtship, there is
a sharing of the duties of incubation as well as those of caring for the young.
It is noteworthy that no ceremonies associated with nest-relief have been:
observed in the Divers or Grebes, whereas they are common in Herons (my
own observations), Pelicans (Chapman, ’08), &c. It would be important to
find out the range of size in male and female Divers. I am inclined to
believe that a single bimodal curve would be obtained. For a discussion of
sex-difference in birds with close similarity of male and female, see Huxley,
722 b. :

GENERAL Discussion.

It remains to discuss some of the general significance of such forms of
mutual courtship as are seen in the Divers, Grebes, Herons, and Hgrets—to
mention only birds whose sexual habits I have myself investigated.

As I have recently pointed out (Huxley, ’21), it is impossible in the
present state of our knowledge to maintain that Darwin’s original theory of
sexual selection is adequate to explain the origin of most of the sexual
ceremonies and adornments to be found in monogamous birds. ‘These
adornments are chiefly used in ceremonies which take place a/ter mating-up
has taken place for the season. There cannot therefore be a direct selection
as between one male and another in respect to them. It may be mentioned

270 MR. JULIAN S. HUXLEY ON

in passing that there is evidence of this direct competition (using the word
in a metaphorical sense) between males in at’any rate some polygamous
species, e.g. the Blackcock (Selous, 709). What, then, is the function of the
sexual adornments and ceremonies—for the most casual observation is enough
to convince anybody that they possess some function ?

It is to be observed that sexual ceremonies and colours or structures which
are displayed solely or chiefly during such ceremonies are only to be found
in animals of a certain grade of nervous organization. Observation on
newts, which are remarkable in that the males deposit their sperm in the
form of spermatophores first and afterwards go through their display
ceremonies, indicates that the function of the display is a stimulative one,
designed to stimulate the female to pick up the spermatophore (quoted in
Pyeraft, °13). The experimental] work of Sturtevant (15) on Drosophila
entirely corroborates this view. The male in this species performs special
actions with his wings when “ courting” the female. Sturtevant found that
males with the wings cut off, when imprisoned singly with females, succeeded
in copulating, but after a longer time than normal males in a similar
situation. On the other band, when each female was placed with two males,
one normal, the other with amputated wings, it was found that the wingless
males were successful in copulating almost as often as were the normals.
This can only be interpreted to mean that the wing-waving ceremony
stimulates the female in such a way that she is then ready to receive
any male in copulation. Morgan (19) appears to adopt this attitude in
viewing the-whole sexual selection problem, but his statements are not
very definite, and he does not bring first-hand observation to bear on the
question. ;

Observation of most of the post-mating courtship ceremonies of birds
bears out this interpretation. If we for a moment confine ourselves to
a consideration of birds with distinct sexual dimorphism, either of
colour and structure, or at least of behaviour, we can say (1) that the
female is always or usually at a lower level of sexual emotion than the
male; (2) that the nervous organization of birds is so advanced that
behaviour is no longer merely reflex, but that it depends in the main on
highly-developed instincts, each associated with a particular emotional
tone. The centres for the instinet require to be stimulated before the
impulse to the particular instinctive action is felt; there is a certain
minimum intensity of stimulation necessary, which, as we have just said,
is, in the case of the sexual instinct, generally higher in the female than
in the male; (3) that, accordingly, a necessary preliminary to coition is
the raising of the emotional state of the female to the level at which her
sexual instinct is stimulated to action.

If we like to sum it up from a slightly different angle, we can say that in
birds (as well as in other groups), mental organization has reached a pitch at
which the higher centres concerned with emotion play a predominant part in

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. PAUL

determining behaviour, and that a stimulation of the emotional centres is a
necessary preliminary to coition. In a precisely similar way, at an carlier
stage of evolution, a stage of general bodily organization was reached at
which internal fertlization was desirable or necessary. On this stage being
reached, it became imperative that coition should occur, and that, in practi-
cally all cases (the majority of birds constitute an exception), specialized
copulatory organs differing in the two sexes should be developed. Both (a)
copulatory organs and (6) most of the courtship ceremonies and epigamic
structures we have been considering can therefore be classed under the same
head—viz. as accessory sexual organs. I have heard this point epigram-
matically summed up in discussion by the remark that this type of male
courtship is a “ psychological penis.” In any event, it is clear that just as
no one has ever considered it necessary to postulate a special type of selection
to account for the origin of copulatory organs, so the origin of courtship, so
far as it is of this stimulative type, does not demand the special evolutionary
mechanism of Darwinian sexual selection to explain it.

The question may now be raised as to the relationship between this
“stimulative ” courtship in species with sexual dimorphism and courtship in
species where the appearance and behaviour of the sexes are similar. To
answer this we must go back for a moment and ask a more fundamental
question :—What is the reason, in the dimorphic species, for the difference in
sexual excitability between male and female? There may be some primary
difference between the sexes which tends in this direction, but there can be
little doubt that the marked accentuation of this, which is seen in most
dimorphic species among the higher animals, has for one main function, as
has been several times suggested (e. g. especially by Grocs,’98, and H. Howard,
07-14), the regulation of coition. If both sexes were highly excitable,
coition would occur too often ; on the other hand, it would be undesirable to
reduce the strength of the impulse too far, until it was near the lower level
at which it ran the risk of being extinguished under the pressure of accidental
unfavourable circumstances. (There is much evidence to show that, e. g., cold
and damp have a markedly depressing influence on the sexual activity of birds.)
An obvious method of ensuring the golden mean between the danger of
excess and that of possible suppression of the instinct is to have one sex with
a very strong instinct, or at least one which is readily excitable, while that
of the other requires considerable stimulation to arouse it. (We have already
seen that the instinct must be aroused through stimulation of the higher
centres.)

There is, however, another method of ensuring a similar result, and that is
to have the sexes at the same general level of excitability, the male less
excitable than in dimorphic forms, the female more so. In this case both
sexes will constantly be reaching the level of stimulation at which desire for
coition occurs. On the other hand, here again it is unlikely that both
will at the same moment spontaneously reach this level—unless both were

272 MR. JULIAN S. HUXLEY ON

keyed so high that an undue amount of coition resulted. Further, when the
level of readiness to pair is reached, it does not follow that the bird will be
under that amount of emotional tension which results in some physical
expression being necessary : in fact, this appears to be the accompaniment
of a definitely higher level of excitement. “‘ Courtship” ceremonies which
result in these circustances may therefore have one of twofunctions. In the
first place, they may be performed by one alone ; they may then convey to
the other that the first is ready to pair—-in other werds be informative in
function—or in addition be stimulative, and help to key the other up to the
requisite pitch. The stretching-out in the passive pairing attitude which
occurs on open water in Crested Grebes (Huxley, 14) and on the nest in
Dabchicks (Selous, °15) appears to be primarily such an informative |
symbol, although it is at least probable that it is also stimulative. On the
other hand, there are many other courtship actions, such as the majority at
least of the mutual post-mating ceremonies described both for the Diver and
the Grebe, which appear to be primarily, or in any case, primitively stimu-
lative in function. They are not symbolic, in the sense that they are not
representations of coition attitudes, and both birds take part in them, usually
though not always performing similar réles. These have as their original
function the raising of the level of sexual emotion—only here it is a mutual
raising of an originally similar level. It is true that, as in the Grebe, these
ceremonies are frequently what I have called self-exhausting, and do not
lead to coition, but this is probably secondary; it is also probable that
they may have other functions besides that of stimulating to coition.
Both these points, however, we must discuss later, as also the reason for
the existence of the two different methods of dimorphic and of mutual
courtship.

It is no doubt often difficult to distinguish between the two; but in the
Crested Grebe at least there appears to be a real distinction. Objectively
(“‘ behaviouristically ”’), e. g. the pairing-attitude is adopted only near the nest
or pairing-platform, and is almost always followed closely by coition ; it is
not accompanied by any of the usual signs of violent nervous activity, nor by
anything that can be called display. Per contra, the mutual ceremonies may
take place anywhere, may or may not be followed by coition, and, if se,
generally after some interval; they are associated with various signs of
general nervous activity, and do serve to display special structures and
colours.

Psychologically, it would appear that the coition-attitude is narrowly and
specifically associated with coition, and is ‘‘informative,’ denoting readiness
to perform the act itself; while the mutual display ceremonies are not
associated with coition in particular, but with general emotional excitement.
In so far, therefore, as they have a stimulative function, it is concerned with
raising the general emotional level to the pitch required to make the birds
repair to the neighbourhood of the nest—i. e. to bringing about the general

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 273

psycho-neural state within which the particular state of readiness to perform
the act of coition itself can only occur.

In so far as they are truly self-exhausting, and not stimulative, they must
either be considered as biologically functionless, as so many by-products of a
mental organization of the type required to execute the stimulative forms of
display (cf. the way in which song has in many cases become a biologically
useless outlet for very various emotional states, as well as subserving its
primary functions in connection with territory); or, as I have previously
suggested (Huxley, *14), their function must be concerned with keeping the
pair, in species in which both sexes share the duties of incubation and care
of young, more closely attached to each other throughout the season.

Here we must be content to note that both types are ‘“‘ accessory ” sexual
characters, in the sense that their main funetion, like that of copulatory
organs, is to facilitate the union of the gametes, and that they represent
different methods of arriving at the same result—viz., the raising of the
emotional level of the pair to a point at which coition will take place.

It may be suggested that to speak of excessive coition, as I have done, is
to use a phrase which it is difficult to justify biologically. It should be
remembered, however, that experiment has shown that, in mammals at least,
a male will, if females in the right condition are provided, continue to pair
until his seminal fluid contains very few spermatozoa (Hays, *18; Lloyd-
Jones & Hay, 718). Normally, of course, this is prevented in mammals by
the fact-that the females only attain the condition in which they are ready
for coition for very short periods at a time. A further disadvantage would
accrue from tie absorption of attention in the sexual act which is so marked
a feature in most animals. The more frequent this might be, the more the
species would be exposed to the attacks of enemies. The whole subject has
been fully dealt with by Groos in his interesting book ‘Die Spiele der
Tiere’ (98) ; and see also Mottram (14).

The regulation of coition, however, is undoubtedly not the only factor of
importance in the evolution of courtship. Previous workers on the subject
have been too exclusively preoccupied with the epigamic significance of
courtship. This was natural; but they should have remembered that the
organism is a whole. We find as a matter of fact that the form and extent
of courtship, nay, in some periods its very existence, is due to causes which
are not epigamic in origin, but connected with other fundamental biological
needs in relation with the annual cycle of the animal.

In many birds, as we shall find (see Howard, *20), it is of prime import-
ance for the welfare of the pair and its offspring that a well-defined territory
shall be occupied in which the nest is afterwards built, and from whose
produce the young are usually fed.. It isa biological advantage to occupy
territory early ; birds which did not do so until late would find all available
space taken up, and would have to fight for occupation. When the sexes are
dimorphic, one will usually be the more active in this task—almost invariably

274 MR. JULIAN S. HUXLEY ON

the male. But once he has occupied the territory, it will be again a biological
disadvantage for the female to put off too long her quest for territory-plus-
mate, or else she too will find herself forestalled.

On the other hand, the exigencies of the food-supply demand that ovi-
position should not occur until a certain date, varying naturally for different
species, but almost always considerably later than the date at which territory
is occupied. Thus we have two opposed advantages—one in the early
occupation of territory, one in the late starting of coition. AS a result, there
will be a period after both birds are on the territory—z.e. after pairing-up
has occurred—when it will be biologically undesirable that oviposition should
occur. This we may call, if we choose to adopt a not wholly suitable human
metaphor, the “engagement period.” Hxtremely little is known concerning
this period in species with mutual courtship: in many of them it is absent,
the birds only returning to the breeding-grounds, or only pairing up,
immediately before coition can and does occur. (This is apparently the case
in Snowy Hgret, Louisiana Heron, and many cliff-breeding birds.) In any
ease, what is known is so slight as to make discussion unprofitable.

In many dimorphic species, on the other hand, a good deal is known.
Such an “engagement period” occurs apparently in all small Passeres which
have nidicolous and insectivorous young, and which therefore, as Howard
has demonstrated, imperatively require a territory which is not merely a
nesting-area, but a hunting-ground from which the needs of the new-hatched
young may be supplied. As we shall see later, the dimorphism itself has
probably been evolved in relation to other biological needs of the species-—
viz. the necessity for concealing coloration and behaviour in the female.
Here, however, we shall take the dimorphism as given.

The simplest way of satisfying these biological requirements is to make the
snale enter in early spring on to a sexually-excited state in which he is
impelled to seek and occupy territory, to pair or. attempt to pair with any
female who stays in the territory, to assist in feeding young when hatched,
ete. So far as his endocrine excitation goes, he remains in the same phase
throughout the season.

With the female, however, the case is different. She must pass through
at least two phases of excitation—the first sufficient to impel her to abandon
the “neutral” non-sexual existence of the winter and to seek and if neces-
sary fight for (P1. 15. fig. 7) the company of a particular male, but not sufficient
for coition to take place; the second, more intense, impelling to coition and
nest-building ; when the eggs are laid, to incubation ; and, later still, to the
care of the young.

There will thus exist a period in which the male will be anxious for coition,
but the female will not permit it. Further, the biological causes for the
existence of such a period are not epigamic, not concerned with the relation
of the sexes, but are to be sought iv relation to the need for occupying

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 275

territory and the need for nesting at the proper time for supplying the food-
requirements of the young.

Such species, where territory is of major not of merely minor importance
( feeding territory as opposed to merely nesting territory), we may for con-
venience call “territory birds.” In them, during the female’s second phase,
there is comparatively little “‘courtship” on the part of the male. When
the female feels impelled to coition, she.adopts a particular attitude, and
the male immediately performs the act. The bulk of the ‘ courtship”
(excluding song) is due to the fact of the existence of the “engagement”
period when the male and female are in different states of endocrine exci-
tation, and this is a biological “accident” arising as a by-product of other
needs and functions.

The song, on the other hand, is, in territory birds (where it reaches by far
its highest development), connected primarily with territory—with attracting
females to, and warning other males off, occupied areas. The displays and
pursuit flights are merely an expression of the desire of the male for coition,
a desire which cannot be satisfied in the engagement period. The display is
in origin an adoption of the coition attitude itself, or something closely
resembling it, brought about naturally as a result of brain-structure when a
certain level of sexual excitement is reached ; the pursuit flight is in origin
an attempt to force the female to submit to coition.

Once, however, these actions had arisen in this way, they formed a possible
basis for further evolution. The fact that in large numbers of species of this
type the male is brightly coloured, and brightly coloured in such a way as to
display the bright colours in the above-mentioned type of ceremony, is
evidence, though of a merely circumstantial kind, that something of the sort
has occurred, and that the ceremonies, originally mere attempts at coition,
have later acquired a second and presumably stimulative function. The fact
that the males of some territory birds are not brighter than the females
(e.g. the Warblers) may well be accounted for on the supposition that in
these species protection is desirable for both sexes.

There is, further, interesting evidence to show that these ceremonies
probably do possess some stimulative function.

I am informed by Mr. Howard—and the statement has been in general
corroborated to: me by Mr. Jourdain and other field ornithologists—that in
seasons when the weather has been very cold and unpropitious during the time
of egg-laying and immediately before, the average number of eggs in a clutch
issmaller than usual, and that the proportion of infertile eggs is higher than
usual. This latter effect may in part be due to actual damage to fertilized
eggs soon after jaying, caused by the inclement conditions ; it may also be
due to failure of fertilization. The smaller number of eggs in a clutch must
be due to failures in ovulation. It is further well known that ovulation in
birds is, in part at least, under nervous control; the caressing of a female
virgin dove on the back of the neck with the hand causes ovulation.

MR. JULIAN S. HUXLEY ON

be
“1
lor)

Harper’s observations (discussed by Marshall, 722, pp. 134-135) make it clear
that the female pigeon will not ovulate unless stimulated emotionally. This
stimulus is normally provided by the male ; but two females isolated together
will often lay. Single females when isolated never lay (except occasionally
when overted, and then only a few eggs). The recent work of Chance (22)
indicates that in the common Cuckoo ovulation occurs under neryous control,
the sight of a pair of foster-parents building their nest acting as a stimulus.
Tt is obvious, further, that the willingness of female birds for coition is not
almost exclusively a matter of internal secretion as in mammals, but is largely
under emotional control. It is also well known to field observers that the
state of the weather has a marked effect upon the emotions and _ their
expression in birds ; song and display is regularly far poorer on a cold, dull
day than on a bright, warm one. Other things being equal, therefore,
anything which tends to counteract the depressing effect of bad-weather
conditions upon the emotions and the actions under their coutrol will be of
biological advantage in that the number of ovulations will be increased, and
also the number of coitions, and so the chances of fertilization for the eggs
after ovulation. There would thus in ordinary seasons be no marked
biological advantage gained from stimulative ceremonies; this would accrue
in exceptionally bad seasons only.

It appears, ag I have already indicated, that in birds with mutual courtship
the ceremonies do have a stimulative effect, although this is usually of a
general and indirect kind, keeping the level of sexual emotion up to a con-
stant pitch, and occasionally heightening it so that visits are made to the
pairing-platform, &c., where coition will usually occur. It is highly probable
that a similar general stimulation is exerted by the more primitive displays
of the “territory” birds, although here the stimulation is not mutual but
unilateral.

It is further clear that any display which did have such an effect would
reap the biological advantages, mentioned in the preceding paragraph, of
counteracting the bad effects of cold seasons. Asa matter of fact, even in
these bad seasons some nests will be found with full clutches of eggs and
normal proportion of fertile eggs. There is, therefore, a form of natural
selection actually in progress here, which (as often) is only operative in
particularly adverse conditions.

It is of very great importance that this point should be cleared up, and
that field ornithologists should undertake careful statistical work on_ the
relation between weather, number of eggs in a clutch, and percentage of
infertility. It is only so that firm bases can be found for biological theories.

It must now be asked, as regards stimulative displays, what form of
stimulus has been adopted and why. So far as males of dimorphic species
are concerned, what is the position? It is that they are subjected to a
mental state which is comparatively rare in organisms below man, of being
stimulated by a powerful emotion but being unable, so long as the female

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 277

remains passive, to express it physically by the natural channel proper to
the instinct which it accompanies. What will the consequence be? It will
be that the psycho-physical energy will discharge into other motor channels,
since it is debarred from discharging into its proper and normal channels.

Perhaps the most frequent way in which it is discharged in birds is by
rapid, restless movement, accompanied by drooping and partial spreading of
the wings, spreading of the tail, and erection of feathers, especially on the head
and breast. This type of action seems to have had a dual origin. In the first
place, it is often very similar to the actions normally adopted at coition, e. g., as
Mr. Howard informs me, in the Buntings. In the Warblers the similarity is
still marked, but to a lesser degree. As I have suggested, it is probably the
direct result of the unsatisfied desire for coition. In the second place,
however, it appears to represent the result of a general state of excitement,
quite apart from any representation of coition attitudes. It is often difficult
in any particular courtship ceremony to disentangle the share of the two
contributions. In the Grebe, for instance, the ordinary ceremonies are in no
way representations of coition actions, while in the Buntings they are almost
wholly so. In any event, the dual origin of the ceremonies as a whole is
clear—general expression of hyper-excitation combined with more or less of
the typical coition attitudes.

Once this type of action exists, however—and it will inevitably come into
being in some form or other in the conditions which we have outlined,—it
will naturally form the chief basis for the development of specialized court-
ship actions and courtship colours and structures belonging to the stimulative
type. When it is the wings and tail which are spread and the crown and
breast-feathers which are erected, it will be inevitable that, if further stimu-
lation is advantageous, it will in the main be achieved by making these parts
more conspicuous (so as to accentuate the difference between the normal
appearance of the bird and its appearance under the-influence of sexual
emotion); and this will be most readily done by the development of bright
colours on these parts, or by the formation of special structures in these
regions—structures which are usually displayed in full only under the
influence of sexual emotion. Since the form of the wings cannot be much
altered without interfering with the efficiency of flight, it follows that special
courtship structures will not often be developed in connection with them ;
and as a matter of fact we find that the chief special courtship structures
are crests on the crown (as in Peacock, Herons, Goldcrest, Peewit, Cranes,
&e.) or breast-plumes or other structures on the breast (asin Herons and
Egrets, Bustard, Umbrella-bird, &c.), or elongations or other specializations
of the tail (e. g. Pheasants, Humming-birds, Blackeock, Lyre-birds, &c.)
or of the tail-region (Peacock, &c.).

The erection af feathers on other parts than crown or breast is also common,
and its specialization gives us structures like the ruff of the Crested Grand
and that of the Ruff, the elongated scapulars of Herons, and those, both

278 MR. JULIAN 8. HUXLEY ON

elongated and dissected, of Egrets, together with some of the peculiarly
situated plumes of Humming-birds and Birds-of-Paradise.

It appears, then, that “courtship” consisted at first chiefly of non-
significant movements employed by the male as an outlet for the pent-up
sexual emotion, these being in part derived from those employed in coition,
and that special colours and structures displayed only mainly during
these movements are later specializations. If so, then we should expect
to find some species of birds in which the movements exist, but not the
special colours or structures—‘“‘ courtship ” actions without epigamie colours
or structures. This is as a matter of fact the case. The Sylviide, for
instance, give us many examples of species which are not dimorphic in
colour, or only so to an obyiously non-significant extent, but are dimorphic
in behaviour, usually the males alone performing special post-mating
ceremonies, apparently stimulative in function, in which wings, tail, and
feathers are moved in the general way that has been described
(BH. Howard, 20). Rarely, however, both birds perform ceremonies simul-
taneously, as is shown in fig.8 (PI. 15). But the two perform ordinary uni-
lateral ceremonies, and not in any common rhythm. There is no sign of a
unified mutual ceremony. It is an interesting intermediate type of behaviour.

Usually, however, the close observation of the courtship of a species
reveals other behaviour which cannot be brought under the rubric of
originally non-significant physical release of emotional tension. I refer to
actions which are to be thought of as having originally been developed in
connection with certain other functions, and later having been connected
with sex in courtship displays. Such actions have already been referred
to. Sometimes they are to a certain extent associated with sexual emotion,
as are all those connected with nesting-material ; sometimes they appear
to have no such connection, such as the preening, head-shaking, dipping
of the bill, &c., which we have noticed in Divers and Grebes. In Mute
Swans, I recently observed some interesting courtship-actions, and there
one of the prominent pieces of behaviour was the dipping of the whole
head and fore-part of the body below the water ; the action was strongly
reminiscent of an incipient attempt at reaching down to feed in the
characteristic swan manner. The birds also repeatedly preened themselves ina
manner very similar to that seen in the Grebes—i. e. a manner indicative of .
the action being used in what I may eall a ritual way, and without any
of its usual functional significance. I am confident that when once attention
has been drawn to this “ ritual” use of non-sexual actions during courtship
activities, it will be found to be of very wide occurrence. Prof. H. Balfour
informs me that it occurs commonly in the Gannet. The association of
nesting-material with courtship is certainly very widespread, and here we
may probably see the working of direct association. The sexual emotion
directly activates the centres connected with nést-building and the handling

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 279

of nest-material, because the two activities have been so often actually
connected. But we cannot adopt this comparatively simple explanation for
the incorporation of wholly non-sexual activities, such as preening, in court-
ship. So far as my personal experience goes (and so little attention has been
devoted to the matter that the literature is of scarcely any help), the birds
which make mast use of these non-sexual actions in courtship are those
which make least use of the peculiar expressional movements of wings, tail,
and feathers generally, which we saw made up the raw material of courtship
actions in most species. If this should turn out to be generally true, then
we should have to suppose that the emotional tension, since for some reason
it does not in these birds express itself in the originally non-significant
methods of general wing, tail, and feather movements, must find an outlet
elsewhere, in the guise of actions which are already performed by the species
in other connections. In any case, it is clear that these actions are utilized,
like the general non-significant actions, as physical relief or expression for
emotional tension; and that since they are already given in the bird’s
instincts and habits, they form the natural raw material—the natural outlet
for surplus psycho-neural excitement due to emotion temporarily unable to
find its normal expression.

In a sense, therefore, the non-sexual associated actions are mere expressions
of emotion, comparable to the meaningless acts often performed by men
under the influence of great emotional tension. But once they have come to
be associated with sexual emotion, they then become the raw material which
can be worked up into more elaborate courtship ceremonies. Such, for
instance, appears to be the origin of the head-shaking ceremonies in the
Crested Grebe. In diving birds generally, there isan almost universal habit
of shaking the head on emerging after a dive. This is, as we have seen,
slightly associated with sexual emotion in the Diver, but in the Grebe the
head-shaking is a very elaborate part of highly-specialized ceremonies, and
its origin from ordinary non-sexual head-shaking can only be deduced from

indirect evidence. Unless, however, its origin be of this nature, it remains

quite obscure.
The association of diving with courtship in both Grebe and Diver may be

adduced as further proof of our contention. The special “splash-dives” in
the Diver, and the emergence from the water in a vertical attitude which
occurs in both species, are both obviously modifications of the ordinary
diving practised for food, which have now an exclusively emotional
significance, and a sexual function. Under the influence of unsatisfied
sexual emotion the bird indulges in various forms of action habitual with
it ; later, specialization has taken place so that the sexual modes of diving
become quite different in appearance from the original non-sexual mode.

As a matter of fact, when we survey the varieties of courtship, we find
that they are not uncommonly to be regarded as specializations of normal

280 MR. JULIAN 8S. HUXLEY ON

activities of the species. For instance, various hawks (e. g. Kestrel and
Peregrine) have courtships which are based upon the wonderful powers
of flight possessed by the group ; while many water-birds, like Grebes and
Divers, show courtships in which some normal aquatic performance is
sexually utilized. Thus in considering the genesis of courtship ceremonies,
we have to distinguish between the raw materials and the later speciali-
zation. The raw materials are (1) coition-attitudes; (2) originally
non-significant general movements, especially of feathers, wings, and tail ;
(3) actions originally connected with activities in themselves associated
with sexual emotion, such as nest-building; (4) actions originally sub-
servient to some non-sexual function.

Later, specialization consists (1) in the development of striking colour on
parts prominently displayed in the raw actions; (2) in the development
of such parts to form special structures (e.g. tail-coverts of Peacock,
ruff of Crested Grebe).

The difficult question remains as to the reasons for the existence of
dimorphism of courtship in some species, but of mutual ceremonies—
associated, be it remembered, in the majority of species, with special colours
and structures of as definitely sexual (epigamic) significance as in dimorphic
forms—in others. —

In general, it appears that it must have some connection with mode of life,
although in the absence of detailed information on many birds, and of
statistical treatment, our conclusion cannot be wholly satisfactory at present.
In the first place, it is undoubted that polygamous species present the most
extreme cases of male adornment, combined with protective colouring in the
females. Secondly, in monogamous species with marked male adornment,
the females are usually protectively coloured, and usually undertake the
whole of the duties of ineubation—e. g. Anatidee, such of the Pheasants as
are monogamous, &c. In some forms with dimorphic courtship, such as the
Sylviide, the two sexes are often not very distinct, and are both more or less
protectively coloured ; both sexes may incubate here, although the female
does so much more than the male. ‘The Sylviidee may be regarded as
primitive in respect of the evolution of post-mating courtship. The
Fringillide are, on the whole, somewhat similar to the Sylviidee, save that
the colour-dimorphism is usually more marked and the male usually
incubates less. E. Howard (’20) has recently shown that territory plays a great
part in the lives of birds, and that, in both Sylviidee and Fringillide, itis the .
male who, early in the season, before other manifestations of sexual emotion,
occupies a definite territory. Singing then begins, and is associated with the
possession of territory. The process of mating-up consists mainly in the dis-
putes of females for occupied territories with their contained males (PI. 15.
fig.7). Since the whole breeding-life of the birds is connected with the staking-
out of territory at the earliest possible opportunity by the males, it follows that

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 281

these must possess different instincts from the females—instincts which
begin to react to seasonal influences some time before any reaction occurs in
the other sex. Here we have a primary basis for the rest of the dimorphism
observed—a basis given by territory requirements, not by epigamic needs.

In birds with both sexes similar in appearance and with mutual court-
ship, two salient facts are to be noted. In the first place, the birds are
usually not protectively coloured, or at least not to such an extent as in
most dimorphic species. Take for example the Herons and Hegrets, the
Swans, the Gulls, and many of the cliff-dwelling and other sea-birds
(Guillemots, Razorbills, Petrels, &e.). The Grebes and Divers are inter-
mediate—the main coloration is protective, but the head, neck, and under-
surface are conspicuous and used in courtship. In the second place, both
sexes almost always share in nest-building, and often in incubation and
the care of the young. So far as I know, there are no birds with mutual
courtship in which the males do not share some of the duties of parent-
hood, and their share is on the average certainly greater than is that of the
males of birds with dimorphism in courtship. Also the nest-site is, usually
at least, chosen by both birds jointly.

It appears to be definitely the case that several tendencies are at work to
produce the variation in modes of courtship that we see. (1) Where, as in
the Warblers, &c., the whole life-cycle is based on the early staking-out
of territory by the males, there must be a dimorphism of instinct in connection
with this, and this primary dimorphism tends to be continued in other ways,
e.g. courtship. (2) Where protective coloration is of great importance to
a species, it is imperative that one sex at least shall be thus protected ; this,
sex will naturally, save in very exceptional cases, tend to be the female. If
protective coloration is desirable, it will almost always follow that general
inconspicuousness will be desirable; this will mean that any violent mani-
festations of emotion, whether in the form of song or of display, will be
undesirable. Hence there will be a tendency in such species to depress the
level of sexual excitability in the females. This in its turn will, however,
make it necessary for there to exist some mechanism for arousing their
sexual emotion, and this will naturally be found in the courtship of the
male. Thus in general the necessity for inconspicuousness in the female
will have as natural reaction a tendency for courtship activities to be
confined to the male, and to be well developed in him. Inconspicuous
females tend to be associated with dimorphism of courtship, and often of
eolour. The tendencies mentioned in (1) and (2) often interact so that it is
hard to distinguish which is the primary. Where polygamy prevails, and the
males take no share in parental duties, this tendency will naturally produce
its most marked results. (3) In some groups, such as for instance
the Herons, not only is it necessary for both birds to share in feeding
the young, but also to share the duties of incubation (since the feeding-

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 20

282 MR. JULIAN S. HUXLEY ON

grounds are so distant; furthermore, protective coloration, as in most
colony-breeding birds, is unnecessary. Here we find two tendencies acting
in the direction of similarity in appearance and behaviour. In the first
place, there is no need for the female to be inconspicuous, whether in
plumage, voice, or action; thus, ceteris paribus, there will be no tendency to
restrict any variations in the direction of greater conspicuousness—even if
they may have been first acquired by the male sex, which is by no means
necessary—to the males. Secondly, since the male as well as the female
must possess the instinct to incubate and to feed the young, it follows that
there is a tendency to approximate the instincts of the two sexes in these
particulars; and the sex-similarity thus initiated again tends to be con-
tinued in other ways, unless there are reasons to the contrary, which, in birds
to whom inconspicuousness is a matter of indifference, we have just seen do
not exist. In birds of this type, therefore, there exist again two mutually
reinforcing tendencies which tend to bring about a similarity of plumage
and instincts, and so lead to mutual courtship.

There are, it is true, species like certain Penguins (Levick, ’14) in which
the sexes are alike, but much of the courtship is not mutual ; there are other
species, like our Red-throated Diver, in which the sexes are similar and
there exists mutual courtship, but in which the male helps to a comparatively
small extent in incubation. In such a complicated subject we should not
expect to find universal hard-and-fast rules; each case requires to be
investigated on its merits, and with a thoroughness which has so far only
characterised few workers. But that the two general tendencies which
I have mentioned, to dimorphism and to mutualism, do exist, and that they
are definitely correlated with other characteristics of the species’ life-history
in the general way I have sketched, appears to be incontestable.

One other point remains to be considered here. Why is it that among
birds with mutual courtship, some, like Divers and Grebes, perform cere-
monies far from the nest, no emotional expression being seen at or close
to the nest during incubation and the care of the young, whereas in others,
such as Herons, Hgrets, Pelicans, &c., the chief ceremonies observed are
those which take place at the nest or nest-site, especially at the moment when
one bird relieves the other from the duty of incubation ? (P1.14. figs. 5 & 6).

The answer is not, I think, far to seek. The “nest-relief birds,” as we
may call the latter category, are all species in which the nest is not concealed,
and does not appear to be in need of special protection against enemies. In
most eases the birds nest in colonies. The birds with mutual ceremonies
which are unconnected with nest-relief, on the other hand, have nests which
are carefully concealed, or at least are far from conspicuous, and rarely breed
in colonies.

When a sitting bird is rejoined by its mate, especially if the mate’s
presence implies a speedy relief from the tedious business of incubation,

COURYSHIP ACTIVITIES IN THE RED-THROATED DIVER. 283

it is natural that emotion should be aroused. If there is no biological reason
against the expression of the emotion, it will naturally be expressed then and
there in some form of ceremony. On the other hand, if there are biological
reasons for concealing the position of the nest from enemies, then it will be
desirable not to draw attention to the spot by conspicuous sounds and actions,
and selection will see to it that the emotion is not expressed immediately
and naturally at the nest, but at a distance from it. ‘The fact that, in the
Crested Grebe, there exists a special and very elaborate ceremony which
occurs when the members of the pair rejoin each other after having been
separated for some time, indicates that in this species too the same emotions
occur in similar circumstances, but are not allowed expression in the neigh-
bourhood of the nest.

Here again, therefore, we find an influence of the general mode of life upon
the form of the courtship. So that we may really say that, although, given
in the first place the state of general excitement produced by the sexual
hormones, in the second place the stimulative effect of sexual display, and
thirdly the necessity for regulating coition, there will inevitably come into
being displays, &c., which go by the general name of courtship, yet the form
of the courtship, not merely in details but in broad lines as well, will depend
in the main upon other general biological factors affecting the species.

There remains to be considered the physiological and genetic mechanism
which we must suppose responsible for the phenomena; but before passing
to this, I should like to summarize briefly the conclusions just reached. In
few words they are these :-—

1. Polygamy encourages sexual dimorphism. It is only found among
birds with nidifugous young, since nidicolous young require that both parents
should assist in feeding.

2. Need for inconspicuousness encourages dimorphism.

3. Nidifugous habit in the young facilitates the development of dimorphism
otherwise initiated.

4, Harly occupation of territory by one sex encourages dimorphism.

d. Absence of need for inconspicuousness encourages mutualism.

6. Need for incubation by both parents encourages mutualism.

In addition, there are no doubt many special physiological points which
encourage one or the other tendency ; but in general the condition observed
is the result of the interaction of the tendencies we have enumerated.

Recent experimental work is beginning to give us an insight, albeit but a
partial one as yet, into the physiological and genetical mechanism underlying
sex-characters in birds.

It appears that castration of a male, in highly dimorphic species like fowl
and wild duck, produces birds which are neutral in instincts but remain male
in plumage.

In females, on the other hand, castration produces cock-feathering, but not
male instincts.

284. MR. JULIAN S. HUXLEY ON

Recently investigations have been made on breeds of fowls like the Sebright
Bantams, in which the cock is hen-feathered. Two interesting points have
emerged. ‘The first is that the condition is due genetically to a Mendelian
gene, the second that it is due physiologically to the fact that the male gonad
in these animals possesses the same substance as do the gonads of normal
females, which inhibits the development of male plumage. This latter point
is demonstrated by castrating hen-feathered cocks, upon which they become
cock-feathered—a strange but conclusive result (Morgan, °19). Morgan
believes that the condition has its histological basis in the development of
luteal cells in the testes of hen-feathered cocks, similar to those found in the
ovaries of normal hens, but recently Pease (19; g. v. for references to previous
work) has thrown doubt on this.

In any case, the facts are of great interest as showing by how simple a
means, and in how few generations, a dimorphism of plumage could be
converted into a similarity. On the other hand, it also shows that in this
dimorphic species at least, plumage and instincts (as the castration experi-
ments also demonstrate) depend upon different mechanisms for their develop-
ment. The development of male or female instincts is due toa positive effect
exerted by some secretion of tle male or female gonads respectively, acting
upon a basis which is neutral in the absence of the specific secretion, whereas
the male gonad has no positive effect upon* plumage development (the
plumage of capons being, if anything, more male than normal), while the
female gonad exerts in this respect an entirely inhibitory effect.

It must be emphasized, however, that we have no experimental information
with regard to species with similar plumage and courtship (in passing, it
may be remarked that the castration of Grebes or Herons should provide
interesting results). Further, from the general considerations advanced
above, it is clear that we should not expect precisely the same mechanism to
hold good in “mutual” as in dimorphic species. In the latter, as we have
seen, selection must have been at work emphasizing and exaggerating any
sexual differences of instinct which primitively existed; whereas in the
former all the influence would tend in the other direction, of assimilating
instinct in the two sexes.

To what pitch this has as a matter of fact been carried is shown by the
observations of Selous and myself upon the Crested and the Little Grebe
(Dabchick). The Grebes are birds in which mutual courtship attains the
highest development yet described, and in which sexual adornments similar
in the two sexes attain a great specialization. In the two species mentioned,
and probably in others, the instinets of the sexes are so alike that coition
occurs both in the normal or in the reversed position. In these birds, coition
always takes place upon a nest; the “passive” bird extends itself at full
length in a perfectly horizontal position, the “active” bird mounts the other
from behind, and, after coition, walks up its body and off by its shoulder.

bP)

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 285

The differentiation of action between the sexes is thus very considerable ; yet
in spite of this, as I say, either sex adopts either position according to
circumstances. In the Moorhen, Selous (02) has described reversed pairing
as a normal sequence of pairing in the usual position. Mr. HE. Howard tells
me that he has repeatedly had occasion to confirm this himself. In Doves
and Pigeons it may also occur (Whitman & Riddle, 19 ; Marshall, ’22, p. 690).

It should be remembered that sexual dimorphism to any marked extent is
a late development in most groups. Primary and accessory organs are,
naturally, different, but general coloration and body-form is usually similar in
both sexes in primitive animals—e. g. Hchinoderms, many primitive insects ;
Crustacea and other Arthropoda, Cyclostomes and most fish. In any case,
recent work has made it abundantly clear that the genes for the secondary
sexual characters of both sexes are normally carried in all members of the
species, and that either the sex-chromosome mechanism ensures two quite
different types of cellular metabolism, one permitting the development of
male, the other of female characters, as seems to be the case in insects, or
else that special hormones are developed in the gonads which exert specific
growth-promoting effects upon some sexual characters, inhibiting effects
upon others, as in vertebrates (summaries in Marshall, 722, meh 15)
Goldschmidt, ’20 ; Huxley, ’22 a).

If, therefore, the divergence which I have referred to between the sexually
dimorphic and mutualist types of birds originated early, as it seems to have
done (for the distinction characterizes whole groups of the class), we may
suppose that the primitive bird species on which it acted were probably
similar in appearance, without well-marked courtship colours or structures,
whether in one or in both sexes, and with a moderate dimorphism of instinets.
Later development has either accentuated the difference of instincts, as in the
Sylviide, or the divergence of instincts and of appearance, as in most
Gallinaceous birds ; or has allowed bright colours to develop, apparently for
use in courtship, but has not confined them to one sex, although it has not
closely approximated the instincts of the two sexes (as in the Penguins,
Levick, doc. cit.), or finally has encouraged sexual adornments similar in the
two sexes, together with markedly mutual courtship, and has approximated
instincts to a high degree—as in the Grebes.

So far as observation can be a guide in these matters, it appears that
although sex-dimorphism in plumage may depend upon different genetic and
physiological factors from that in instincts and from that in size, yet asa
matter of evolutionary fact, the first two, and probably the third Halse; have
gone hand in hand.

In other words, the divergenee between dimorphic and mutualist species is
primarily dependent on whether the gonadial hormones remain as similar as
is compatible with primary sex-differentiation, or whether they become con-
siderably different in the two sexes. If they adopt this latter alternative,

286 MR. JULIAN §. HUXLEY ON

then there will result, unless special circumstances arise, a simultaneous
dimorphism in both appearance and instincts ; if they remain on the whole
similar, there will be (again unless there are special reasons against it) a
similarity between the sexes, not only as regards appearance, but also as
regards courtship and parental behavionr.

If this turns out to be well founded, and the gonads do normally in birds
determine appearance, behaviour, and size simultaneously, then it is clear that
observation will have contributed important guiding lines for future genetic
and evolutionary research.

In conclusion, I would like to emphasize the fact that, even when both sexes
are similar, there is often no doubt whatever that the coloration and special
structures used by them both in courtship have been evolved in relation to
sexual habits, and in essentially the same way as bright colours and special
structures used in courtship by males alone in dimorphic species. They have
arisen through the necessity of providing an emotional stimulus to the other
member of the pair, if fertilization is to be normally and easily effected,
particularly in the face of adverse environmental conditions; and from the
reverse point of view, have become necessary because it is biologically
desirable to keep the mean sexual excitability of the pair at a level low
enough to prevent excessive coition. They are, to use the useful term of
Poulton’s which I have for some time adopted, always epigamic, although not
always secondary sewual characters. They have also been usually grafted on
to primitive ceremonies which are the direct outcome of sexual excitement,
and indirectly often the result of an “engagement” period which is caused
by non-sexual exigencies of the life-cycle.

Although the direct competition among males, which was one of the main
points of Darwin’s theory of sexual selection, does not appear to occur in the
evolution of most epigamic characters among birds, yet it is noteworthy that
in the polygamous species in which it does occur, the type of coloration and
structure which is evolved is very similar to that evolved in monogamous
species, except that development has usually gone to further lengths of
specialization. This similarity between the end-results in what we may call
“aecessory ’ as well as in “ sexually-selected”’ epigamic characters, is due to
the second salient fact first clearly recognized by Darwin—the fact that the
development of an epigamic character is dependent upon the emotional effect
which it produces upon the mind of a bird of opposite sex. Hpigamic
characters must be emotionally stimulative if they are to perform their
function successfully ; and to be emotionally stimulative, it would appear
that they must be first striking, and secondly not usually yisible, so as to have,
when they are displayed, an element of novelty and unfamiliarity which adds
to their stimulative effect. So far as their striking quality is concerned, they
may be either what seems to us beautiful or else what seems to us bizarre.
So far as unfamiliarity goes, it should be remembered that many epigamic

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER. 287

characters are special structures only fully displayed during courtship (e. g.
seapulars of Hgrets, tail-coverts of Peacock, ruff of Crested Grebe) ; others
are special patches of colour only made prominent by special display actions
(e.g., white on wings of Blackcock and of Crested Grebe, red of legs of Red-
shank, rich purplish-brown of neck of Louisiana Heron); while still others,
although always present, are probably not prominent unless the two birds are
close to each other, and in certain relative positions characteristic of courtship.
This is probably the case with many of the curious patterns characterizing the
front view of birds at short range (e.g. Blue Tit, Turnstone, Redstart, Ringed
Plover) ; that this is the probable explanation is shown by the fact that when
special structures with undoubted epigamie function are developed on the
head, these are often displayed so as to appear most striking (or only so)
when seen in this way, close up and from the front (e.g. the ruff and ear-
tufts of the Crested Grebe, the crest of Louisiana Heron and Little Hgret).

The existence of true sexual selection as found in polygamous species thus
encourages the same tendencies in epigamic characters as does the selection
of those with a purely ‘“‘accessory ’ function ; but, owing to the fact that in
polygamous species the males take no share in incubation or the care of the
young, dimorphism can proceed to its limit, and owing to the fact that there
is a real selection as between different males, and so greater competition in
regard to secondary sexual characters, and that the successful male transmits
his qualities to a greater number of offspring, the process of evolution of
epigamic characters is not only more rapid, but also is generally carried to a
higher pitch than in monogamous species. In other words, polygamy and its
attendant true sexual selection simply accentuate the same processes that are
operative whenever epigamic characters are being evolved, even in the
absence of sexual selection proper.

This development of epigamic characters in relation to the mind of the
opposite sex is a point of very general biological interest, since it is the only
example, in organisms below man, of a secondary effect of the mind of a
species upon the evolution of that species. Mental qualities of course
normally have survival value, but this is quite a different matter ; they thus
determine the survival or extinction of the species, not its modification in new
ways. ‘The mind of one species may play a part in moulding the evolutionary
development of other species, as when acute vision on the part of predaceous
animals renders concealing coloration advantageous, or the visual and olfactory
preferences of flower-visiting insects are reflected in the development of colour
and scent in the flowers visited. Lloyd Morgan (’21) has recently emphasized
the influence of mind upon eyolution by introducing the useful term ‘“ Psychical
Selection.” As indicated above, however, further terms are needed to
distinguish between psychical selection acting upon the evolution of other
species, as in the case of the mental qualities of bees influencing the evolation
of flowers, or upon the evolution of the same species, as in the development

288 MR. JULIAN S. HUXLEY ON

of stimulative display characters. If the barbarism be permitted, the terms
“heterospecific ” and “autospecific” psychical selection might be employed
to make this distinction. But once epigamic characters come to be
advantageous, the mind of the species (in the females in sexually dimorphic
forms, in all individuals in those with mutual courtship) is exerting the
indirect effect we have been describing upon the future development of
colour, structure, and behaviour in the race. This is the most important
fact which Darwin perceived, and this stands firmer than ever in spite of
the rejection of the bulk of the other part of his doctrine.

In concluding, I should like to thank Professor Lloyd Morgan, Mr. Eliot
Howard, and Mr. A. M. Carr-Saunders, all of whom have kindly read the
foregoing article in manuscript, and have helped me with several important
criticisms and suggestions.

Now College, Oxford,
March, 1928.

Postscript 1.—Since the above was written, Dr. J. C. Mottram has been
good enough to write to me on a number of points concerning sexual
coloration in birds, and to allow me to see the MS. of an unpublished paper
on the subject. I would like to take the opportunity of dealing with a few
of the points which he raises in this and in his book (714).

He points out the great importance in many birds of concealing
coloration—a proposition in which every naturalist who has studied
birds in the field would agree with him. I have in the body of the
paper dealt with the ways in which the necessity for concealment or its
absence reacts upon the “courtship ”’ characters and activities of birds.

With reference to the Kingfisher (Alcedo ispida), Dr. Mottram makes out
a strong case for believing that its brilliant colouring, present of course in
both sexes, is aposematic, with the function of warning possible enemies of
the bird’s unpalatability. He points out that records a the Kingfisher being
attacked by birds of prey are extremly rare ; and has found that its flesh is
unpalatable to man and rejected by domestic animals.

However, if the Kingfisher really, as seems probable, presented an example
of warning coloration, it would in no way invalidate my general conclusions ;
it would merely corroborate from a new angle what I have been urging in
this paper—viz., that each species of bird must be worked out on its merits,
and that.coloration and behaviour are always determined not by one single
cause, but by several, of which the two most important are (a) the bird’s
relations with its mate, and (6) its relations with its enemies and its prey.

Dr. Mottram, however, goes farther. He attempts to account for all
“courtship” characters and actions solely in terms of the bird’s relations
with enemies and prey.

(1) By an ingenious train of reasoning he points out that where the hen
alone, or chiefly, incubates, she must be regarded as biologically the more

COURTSHIF ACTIVITIES IN THE RED-THROATED DIVER. 289

valuable of the pair, and that in such species the display and song of the cock
has as its primary function that of drawing the attention of enemies away
from the hen ; so that, if one of the two is sacrificed, it shall generally be the
less ‘‘ valuable ” male.

(2) Where the species is capable of defence, or is unpalatable—z. e. does
not require protective coloration—he postulates that courtship should be not
elaborate, or may even be absent.

(3) Where both sexes are of equal biological value (7. e. where both
incubate), he postulates that both should indulge equally in display.

The reasoning is undoubtedly ingenious, but can be shown not to be
adequate to account for all the facts.

As regards (1) , it should be remembered that the song of male “ territory
birds” is most powerful before any females have arrived. It could be
no possible advantage to the species to have attention attracted to the
males when there were no females present from whom attention could
be drawn off.

Ié is undoubtedly true that where only the female incubates, and where
protection is desirable, dimorphism of sexual characters and behaviour will be
encouraged. .This I have pointed out in the body of the paper. Dr. Mottram
simply restates the problem from a different angle, and with the epigamic
side of the matter left out.

(2) is simply not true in a number of cases. Herons and Egrets are well
protected from enemies, but have an elaborate courtship. So do Swans
(Selous, °13, and my own observations). So do Hawks, though here the
displays are concerned with flight, not with bright coloration (Owen, *16—22,
Sparrow-Hawk ; Kestrel, my own obervations) ; and other examples could
be quoted. ;

(3) Where both sexes are of equal value, it would appear on the face
of it biologically desirable that both should be rendered inconspicuous when
concealment is in any way desirable, if, as Dr. Mottram would have us
believe, the chief function of display is to attract the attention of enemies.
However, in the Grebes, all of which take great pains to conceal their nest,
and are protectively coloured so long as they are in the normal swimming,
feeding, or resting attitudes, elaborate mutual courtship does occur, and at
once makes the pair conspicuous. The same, to a less extent, is seen in Divers.
The vocal duets of Owls which go on in the breeding-season (Huxley 19)
are difficult to account for on any theory which relates them only with
attracting the attention of enemies.

Finally, Dr. Mottram makes the point that since many birds “ get on”
without bright colouring in one or both sexes, and yet reproduce, therefore
when bright colouring is present it cannot be supposed to have any relation
to reproduction. This, however, is an old fallacy, repeatedly exposed since the
early days of the evolution controversy. It would be equally justifiable to say

LINN. JOURN.—ZOOLOGY, VOL. X¥XY. 2A

290 : ‘MR. JULIAN S. HUXLEY ON

that because an earthworm gets on without eyes, and yet reacts to light and
darkness, therefore when eyes are found, they can have no relation to light-
perception ; or that because lower mammals get on with a small brain,
therefore the difference in brain-size between them and man can have no
relation to the functions carried out by the brain of lower mammals ; and
so on, ad lib.

It becomes increasingly clear that to interpret the behaviour and
evolution of a bird, even if apparently in only one regard, it is necessary to
take into account all the circumstances of its life. As regards coloration
and “ courtship ” behaviour, I hope I have shown that relation to enemies
and relation to the mate are two factors of greatest importance. I am
grateful to Dr. Mottram for having brought to my notice a number of points
concerning the relation of birds to their enemies of which.I was not aware ;
and I believe that his contribution to the theory of the subject is of real
value. But so long as cases remain—as they do in large numbers—where
coloration, structure, or behaviour have an obvious function in regard to the
opposite sex, and no discoverable function in relation to the enemies or any
other factor of the bird’s life, 1 maintain that we must take these into account,
and that Dr. Mottram is definitely wrong in attempting to base his theory of
the evolution of courtship solely on one, instead of on both of the two main
factors mentioned above.

On the other hand, his suggestion that the bright colour and general con-
spicuousness of the male may have in some species the function of drawing
the attention of enemies from the biologically more valuable female will,
I believe, prove of importance in certain puzzling cases. I will mention but
two. In the first place we have the well-known fact that many mimetic
species ot butterflies, including some of the most striking examples of mimicry
(e.g. Papilio dardanus), are mimetic in the female sex alone, the male being
conspicuous but non-mimetic. The males are “valuable” only before copu-

lation: the females until after oviposition, and further, have usually a poorer
flight. Ifa certain average toll is taken of the species by enemies, it could
well be a real advantage to have it fall preponderatingly on the male sex.

. The second example concerns birds. Reflection will show that the bright
colours of male birds fall broadly into two categories—thouse which, like’ the
“tail” of the Peacock or Argus Pheasant, are only conspicuous in display,
and those which, like the general coloration of most male ducks, are always
conspicuous. It will be further found that the former are definitely displayed,
the latter usually not. .¢., the display of male ducks is largely a bobbing
of the neck ; the general bright body-colour is not made more conspicuous in
display. The male duck, furthermore, (1.) takes no share in incubation or
rearing the young, (ii.) usually stays near the hen until the young are hatched,
(iii.) as soon as he leaves the hen, moults into eclipse (protective) plumage.
(iv.) The hen is protectively coloured. : (v.) The species is highly palatable and
much attacked by Raptores.

COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER, 291

Thus there is a good prima facie case for applying Dr. Mottram’s ideas to
cases of bright coloration in the males of palatable species in which the
bright colour is not specially associated with display, and especially in forms
in which the female alone looks after eggs and young.

It is, however, becoming clear that many side issues of the problem can
best be attacked by siensineal methods ; and this I propose to do as oppor-
tunity offers.

Postscript 2.—The last number of ‘ British Birds’ (vol. xvi. p. 318) con-
tains an article by Mr, N. Gilroy on the nesting habits of the Black-throated
and Red-throated Divers to which reference should be made.

The Black-throated Diver is mentioned as dipping its bill in the water
when nervous because scared off its eges. This extends my observations of
the beak-dipping of the Red-throated Diver to another species.

Of courtship, the following remark is made—* The pairs may be observed
on the tarns fora conetlerabls time before nesting begins, and one is fre-
quently treated to the extraordinary display which takes place, not only
during courtship and pairing, but even when actual breeding has commenced
aya both birds propel themselves through the water sla vertically.”

This, so far as it goes, confirms my observations. J.S. A.
8th May, 1928.

List or Lrreravrure.

Baur. 07. The Home Life of some Marsh-birds. London, 1907. (Divers.)

Bent. *19. Bull. U.S. Nat. Mus. Washington, no. 107. (Divers.)

Cuance, ’22. The Cuckoo's Secret. London, 1922.

CHapman, 08. Camps and Cruises of an Ornithologist. New York, 1908. (Pelicans, Kc.)

CuHapMman. ’22. Bird Lore, xxiv. p.1. Loon.)

Darwin. 71. Descent of Man. London, 1871. (See esp. p. 277.)

Dewar. 720, Brit. Birds, xiv. p. 89. (Blackbird.)

Groos. 798. The Play of Animals. (Eng. Trans.) New York, 1898.

Gotpscumip?. 20. Meehanismus und Physiologie der Geschlechtsbestimmung. Lerlin,
1920.

Gurney, J. H. 13. The Gannet. London, 1913.

Hays. 718. J. Exp. Zool. xxv. p.571. (R abl bits.)

THlowarp, K. 07-14. The British Warblers. London, 1907-1914.

Howarp. ’20. Territory in Bird Life. Londen, 1920.

Huxtry. 14. Proc. Zool. Soc. 1914, p. 491. (Crested Grebe.)

Huxrry, 716. Auk, xxxiii. p. 152 & p. 256. (General.)

Houxtey. 19. Brit. Birds, xiii. p. 155. (Dabchick; Owls.)

Huxtry. ’21. Nature, cviii. p. 565. (General.)

Huxtny. 22a. J. Roy. Soc. Arts, xx. (Sex-determination.)

Huxtery. 226. Brit. Birds, xvi. p. 99. (Assortative mating.)

Levick. ‘14. Antarctic Penguins. London, Heinemann & Co., 1914.

Itoyp-Jonus & Hays, 718. J. Exp. Zool. xxv. p. 463. (Rabbits.)

Lioyp-More@an. “21. Brit. J. Psychol. xi. p. 206. (Psychical selection.)

MarsHau. 22. The Physiology of Reproduction. London, Longmans, Green & Co., 1922.

Morean. 19. Carnegie Publ. no, 285, (Inheritance of sexual characters; sexual selection.)

292 COURTSHIP ACTIVITIES IN THE RED-THROATED DIVER.

Morrram, 714, Controlled Natural Selection and Value marking. London, Longmans,
Green & Co., 1914.

Owen. 716~22. Brit. Birds, x., xii., xv. (Sparrow-Hawk.)

Pease. ’22, Proc. Cambr. Philos. Soc. xxi. p. 22.

Pixs. 713. Brit. Birds, vii. p. 148. (Crested Grebe.)

Poutton. 08. Essays on Evolution. Oxford, 1908. (pp. 379, 386.)

Pycrart, 713. The Courtship of Animals. London, 1913.

Sepuirz, 7138, Journ. f. Ornith. cxi. p. 179. (Black-throated Diver.)

Srxovus. ’01-’02. Zoologist (4), v. & vi. (Crested Grebe ; Peewit.)

Srxovus. 02. Zoologist, 1902, p. 196. (Reversed pairing in Moorhens.)

Seous. 05a. The bird watcher in the Shetlands. London, 1905. (pp. 122-126.)
(Mutual courtship.)

Sexous. 056. Bird-life Glimpses. London, 1905. (p. 296.) (Dabchick.)

Setovus. 09. Zoologist (4), xiil. & xiv. (Blackcock.)

Seous. 12a. Zoologist (4), xvi. p. 81; xvil. p. 210. (Red-throated Diver.)

Sexous. 120. Zoologist (4), xvi. p. 197. (Origin of Display.)

Sexovs. ’13. Zoologist (4), xvil. (pp. 294, 409). (Swans.)

Setovus. 714. Zoologist (4), xviii. p. 73. (Great Northern Diver.)

Srtovus. 15. Wild Life, vii. (Dabchick.)

Sturtevant. 15. Journ. Anim. Behaviour, v. p. 853. (Drosophila.)

Turner. 713. Brit. Birds, vii. p. 150. (Red-throated Diver.)

Van Oorpr & Huxtry. ’22. Brit. Birds, xvi. p. 34. (Red-throated Diver.)

Wuirman & Rippxe. 119. Carnegie Inst. Pub., Washington, 1919. (Pigeons; Doves.)

Wirnersy. ’00. Ibis, 1900. (Red-throated Diver.)

EXPLANATION OF THE PLATES.

Prater 14,

Fig. 5. Louisiana Heron (Hydranassa tricolor). Twig-bringing ceremony after nest-relief.
The relieved bird is presenting a twig to the bird on the nest. Note raising of
crest, aigrettes, and neck-feathers, and spreading of wings.

Fig. 6. Nest-relief ceremony of Little White Egret (Zgretta candidissima). The bird on the

right is about to step off the nest. Note erection of crests and aigrettes, and
spreading of wings.

PuatrR 15.
Fig. 7. Two female Whitethroats (Sylvia cinerea) fighting for a male ard his territory;
the male watching.

Fig. 8. A pair of Willow-Warblers (Phylloscopus trochilus) ; simultaneous performance of
wing-flapping ceremony.

Avuxiry. JOURN. LINN. SoOC., ZOOL. VOL. XXXV. FL. 14.

Fig. 5.—Louisiana Heron (Hydranassa tricolor). Twig-bringing ceremony
after uest-relief. The relieved bird is presenting a twig to the bird on
the nest. Note raising of crest, aigrettes, and neck-feathers, and
spreading of wings. (Photo by J. S. H.)

Fig. 6.—Nest-relief ceremony of Little White Egret (Egretta candidissima).
The bird on the right is about to step off the nest. Note erection of
erests and aigrettes, and spreading of wings. (Photo by J. 8. H.)

HUuXLry.

Journ. LINN. Soc., ZOOL. VOL. XXXV. PL.

Fie. 7.—T wo female Whitethroats (Sylvia cinerea) fighting for
a male and his territory ; the male watching.
Reproduced by permission from E. Howard, ‘The British Warblers.’

Fic. 8.—A pair of Willow-Warblers (Phylloscopus trochilus) ;
simultaneous performance of wing-flapping ceremony.
Reproduced by permission from E. Howard, ‘ The British Warblers.’

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CONTENTS.
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I. On Cellularine and other Polyzoa. By Sir Srpyny F. Harmer,
K.B.E., Se.D., V.P.R.S., Director of the Natural History
Departments of the British Museum. (Plates 16-19.) ......... 293

II. Notes on Acari, with Descriptions of New Species. By J. N.

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ON OELLULARINE AND OTHER POLYZOA. 293

On Cellularine and other Polyzoa. By Sir Stpney F. Harmer, K.B.H.,
Se.D., V.P.R.S., F.L.S., Director of the Natural History Departments
of the British Museum *.

(PLATES 16-19.)

[Read 1st February, 1923.]

I. IntRopDUCTION.

THE collection of the Polyzoa made by the ‘Siboga’ in Malay waters is
perhaps the most important one which has been obtained in any part of the
world since the ‘Challenger’ Expedition, including as it does representatives
of the shallow and deep water Fauna of a district rich in Polyzoa, but at
present very imperfectly known. A study of the representatives of this
group in the area in question is specially needed, in order to fill a conspicuous
gap in our knowledge. The Polyzoa of parts of Australia have perhaps been
more fully studied than those of any other part of the world, with the
exception of the Atlantic and Arctic areas ; particularly those of Victoria
and New South Wales, thanks to the labours of Busk, Hincks, MacGillivray,
Waters, and others. Those of Japan have been described, to some extent, by
Ortmann, and are receiving attention from Yanagi and Okada. Hincks and
Miss Robertson have described some of the Polyzoa of the Pacific coast of ©
North America, but those of the Pacific in general have been very imperfectly
investigated, although some information on the subject has been given by
Miss Philipps. Waters has published two important papers on the Polyzoa
of the Red Sea and Hast Africa, and Savigny’s admirable figures of Egyptian
species, some of which were obtained in the Red Sea, as well as a recent
paper by Marcus on South African species, must be specially noticed.
Hincks has described Polyzoa from the Indian Ocean, particularly from the
Mergui Archipelago ; and Miss Thornely and Miss Robertson have also made
contributions to our knowledge of the Polyzoa of this Ocean.

In investigating the Cheilostomata of the Malay Archipelago, I have
found it necessary to devote much attention to various questions which do
not form a necessary part of the Report which is in preparation. I have
been obliged in particular to study various genera, in order to attempt to
decide their characters and the names which ought to be assigned to them.

* Published by permission of the Trustees of the British Museum.
LINN. JOURN.—ZOOLOGY, VOL. XXXV. 22

294. SIR SIDNEY F. HARMER ON

In dealing with these questions I have had the great advantage of being able
to make use of the important type-specimens, particularly those of Busk, in
the British Museum. Some of the results thus obtained are given in the
present paper, which is intended to be preliminary to my fuller Report.
The genera considered are those of the ‘ Cellularine ” series, together with
others which have to be discussed in connection with these. For the purposes
of this general survey I am obliged to give diagnoses of several new genera ;
but I restrict myself to those which are strictly necessary, believing as I do
that the practice of publishing preliminary diagnoses which cannot be fully
understood without adequate illustration is one to be avoided as much as
possible. I devote special attention to species which have been included in
the genus Menipea ; and I venture to think that the grouping here suggested,
in this and other genera, will give greater precision to certain questions of
Geographical Distribution. I desire to acknowledge specially the valuable
assistance I have received from my colleagues Mr. C. Davies Sherborn and
Mr. R. Kirkpatrick. Mr. Sherborn has helped me particularly with regard
to the dates of publication of various Memoirs, while his invaluable MS. list,
at present in course of publication (Index Animalium, 1801-1850) by the
Trustees of the British Museum, has been of special assistance in ascertaining
that several genera in current use are preoccupied. Mr. Kirkpatrick, who
is in charge of the Collection of Polyzoa in the British Museum, has given
me much appreciated help in referring to the specimens, and in other ways.
Special attention may be directed to Sect. III. (Internal avicularia) and
Sect. IV. (Methods of bifurcation).

The present paper is divided into the following Sections :—

I. Introduction.
II. Literature and Nomenclature.
III. Internal avicularia.
IV. Methods of bifurcation of the colony.
V. Characters of certain genera and species of Scrupocellaride.

I. Lirerarurr anp NoMENCLATURE.

This Section includes a list of Memoirs to which shortened references are
given throughout the paper; but full references are given, in their proper
places, to many other works not included in the list. The Bibliography is
followed by an alphabetical list of genera, in which I have endeavoured to
establish the respective genotypes, where this has not already been done.
In constructing this list I follow the example which has been given by Lang
(1917, Geol. Mag., Dee. vi, vol. iv. p. 169). I have also indicated, as far as
possible, the names which appear to me synonymous with genera of earlier

CELLULARINE AND OTHER POLYZOA. 295

introduction. The consideration of a number of genera at the same time has
had great advantages, by enabling many cases to be discovered in which the
conclusions drawn from the study of one genus by itself are modified by
taking others into account. I have found it necessary to propose the following
new genera and species in the present Section :—

Camptoplites, n. gen.

Didymozoum, nom. noy., to replace Didymia, pre-occupied.
Euoplozoum, n. gen.

Himantozoum, n. gen.

Onchoporella buskii, nom. nov., for Onchoporella bombycina, Busk.
Stirpariella, nom. nov., to replace Stirparia, pre-occupied.

‘The following new names are proposed in Sect. V. :—
Amastigia kirkpatricki (Levinsen, MSS.), n. sp.
Menipea vectifera, n. sp.

' Notoplites, n. gen., and NV. rostratus, n. sp.

I accept the main divisions of the Cheilostomata which were suggested by
Levinsen (1909, pp. 88-90). The Order is divided into two Sub-Orders :
(1) Anasea, in which the original frontal membrane persists, wholly or in
part, in a membranous condition; (2) Ascophora, in which a compensation-
sac is present (cf. Harmer, Q.J.M.S. xlvi. p. 263). The Anasca are further
subdivided into three Divisions: (1.) Malacostega, with the frontal membrane
for the most part in its primitive, membranous condition ; (ii.) Coilostega,
with a considerable development of a calcareous layer, the cryptocyst,
subjacent to the frontal membrane, and frequently pierced by foramina, the
opesiules, traversed by the tendons of the depressor muscles, which are’
inserted into the frontal membrane; (ili.) Pseudostega, with peculiar,
immersed ovicells and certain other characters, consisting mainly of the
Family Cellariidse. ©

The genera included in the following list are mainly the erect branching
forms of the Anasca. I have included all Levinsen’s Families of recent
Malacostega, with the exception of the Membraniporidee and Cribrilinide ;
the erect, branching genera of the Coilostega, but not the encrusting forms
nor those with an Hscharine habit; and the genera of recent Pseudostega.
The Cellularine assemblage, with which I am principally concerned, consists of
the branching, erect genera of the Malacostega. The branching Ascophorous
‘genera, such as the Catenicellide, provided with a compensation-sac, are
not included in my list, with one or two exceptions, in the case of names
which have some bearing on the nomenclature of other genera under

-consideration.

bo
ite)
lor)

SIR SIDNEY F. HARMER ON

A. LItERATURE.|

Abbreviations :—
A.M. N. H.—Annals and Magazine of Natural History.
A.S. N.—Annales des Sciences Naturelles (Zoologie).
J. L.8.—Journal of the Linnean Society (Zoology).
P. F. T. C.—See under D’Orbigny.
Q. J. M.8.—Quarterly Journal of Microscopical Science.

Avpourn, V., 1826.—* Explication sommaire des planches de Polypes de I’Egypte et
de la Syrie,” in Savigny’s “ Description de l’Egypte,” i. pt. 4.

Avpouin, V., 1828.—Jbid., 2° Edition, xxiii.

Buarnvittr, H. M. D. ps, 1830.—Dictionnaire des Sciences Naturelles, lx., Zoophytes.

Brarnvitte, H. M. D. pn, 1834.—‘* Manuel d’Actinologie.”

Busx, G., 1852*.—Polyzoa and Sertularian Zoophytes, in J. MacGillivray’s Voyage of
the ‘ Rattlesnake,’ Vol. i.

Busk, G., 1852*.—‘‘Catalogue of Marine Polyzoa in the Collection of the British
Museum,” Pt. i.

Bus, G., 1884.—‘* Report on the Polyzoa,” Pt. i., Challenger Reports, Part xxx.
Vol. x.

Exuis, 1755.—‘* An Essay towards a Natural History of the Corallines.”

Eturs, J.. & Sozanper, D., 1786.—%‘ The Natural History of many curious and
uncommon Zoophytes.”

Fromine, J., 1828.—“ A History of British Animals.”

Gray, J. E., 1843.—“ Additional Radiated Animals and Annelides,” in E. Dieffen-
bach’s ‘“‘ Travels in New Zealand,” Vol. 11.

Gray, J. E., 1848.—** List of the Specimens of British Animals in the Collection of
the British Museum,” Pt. i. “‘ Centronize or Radiated Animals.”

Hincxs, T., 1880.—‘ A History of the British Marine Polyzoa.”

Jounston, G., 1847.—<‘ A History of the British Zoophytes,” Second Edition.

Krnues, H., 1914. — Die Bryozoen der deutschen Siidpolar-Expedition 1901-1903,”
pt. i., Deutsch. Siidpolar-Exp. 1901-1903, xv. (Zool., Bd. vii.) pp. 601--678.

Lamarcn, J. B. P. A. pn, 1801.—“Systéme des Animaux sans Vertcbres.”

Lamarcr, J. B. P. A. pv, 1816.—** Histoire Naturelle des Animaux sans Vertébres,”
Vol. i.

Lamovrovx, J. V. F., 1812.—‘ Extrait d’un Mémoire sur la Classification des Poly-
piers Coralligénes non enti¢rement pierreux,” Nouy. Bull. Sci. Soc. Philomat.
il. p. 181.

Lamovrovx, J. V. F., 1816.—* Histoire des Polypiers coralligénes flexibles.”

Lamovrovx, J. V. F., 1821.—* Exposition Méthodique des genres de Ordre des
Polypiers.”

Lamourovux, J. V. F., 1824.—In Encyclopédie Méthodique, xey. Livr., T. ii., 1 Partic,,
Vers.

Lamovroux, J. V. F., 1827.—Jbid. xcvin. Livr., I. i1., 2 Partie, Vers.

Levinsen, G. M. R., 1909.—‘ Morphological and Systematic Studies on the Cheilo-
stomatous Bryozoa,” Copenhagen. ‘

Liyy avs, C., 1758.—“ Systema Nature,” 10th Ed., Vol. i.

CELLULARINE AND OTHER POLYZOA. 297

Linnavs, C., 1767.—“‘ Systema Nature,” 12th Ed., Vol. i. pars 2.
Marcus, E., 1922.—“‘ Siidafrikanische Bryozoen aus der Sammlung des Gothenburger
Museums,” Goteborgs K. Vet.- och Vitt.-Samh. Handl. (4) xxv. 3.

Oxen, L., 1815.—‘‘ Lehrbuch der Naturgeschichte,” 3 Theil, Zool., 1 Abth.
D’Ornieny, A., 1852 (referred to as A. 8. N.).—‘‘ Recherches Zoologiques sur la
Classe des Mollusques Bryozoaires,” Ann. Sci. Nat. (3) Zool. xvi. p. 292.
D’Orsieny, A., 1852 (probably published later: referred toas A. S. N.).—Jbid., Suite,

Ann. Sci. Nat. (3) Zool. xvii. p. 273.
D’Orsieny, A., 1851-1854 (referred to as P. F.T.C.).—* Paléontologie Francaise,
Terrains Crétacés,” T. y.
1851.—pp. 1-188.
1852.—pp. 189-472.
1853.—pp. 473-984.
1854,—pp. 985-1192.
Patras, P. 8., 1766.—* Elenchus Zoophytorum.”
Pattas, P. 8., 1787.—‘* Charakteristik der Thierpflanzen” ... . tibersetzt von C. F.
Wilkens und J. F. W. Herbst, i. Th.
Scuweicerr, A. F., 1819.—‘ Beobachtungen auf Naturhistorischen Reisen,” Berlin.
Scuwerecer, A. F., 1820.—“ Handbuch der Naturgeschichte der skelettlosen ungeglie-
derten Thiere,” Leipzig.
Suirr, F. A., 1868.—“ Kritisk Forteckning ofver Skandinaviens Hafs-Bryozoer,” iii.,
Ofv. K. Vet.-Ak. Férh. xxiv. 1867, p. 279.
Woopwarp, b. B., 1903-1922.—Catalogue of the Books, Manuscripts, Maps and
Drawings in the British Museum (Natural History), vols. iv. and Suppl.
(This work contains much valuable Bibliographical information with regard to
dates of publication.)

B. NoMENCLATURE.

Acamarchis, Lamouroux, 1816, p. 132.—Genosyntypes, A. neritina (L.) and A. den-
tatu, Lamx.,n.sp. Genolectotype, Acamarchis neritina (Sertularia neritina, L.,
1758, p. 815), a selection which may be considered to have been made by
Schweigger, 1819, Tab. 8; 1820, p. 429; with erroneous citation as
neretina. It may be noted that D’Orbigny (1852, P.F.T.C. p. 328) gives
1812 as the date of introduction of Acamurchis; but his synonymy on the
following page shows that he referred to Lamouroux, 1816. I regard
Acamarchis as a synonym of Bugula, Oken, 1815, in accordance with the
practice of most modern authorities.

Actea, Lamouroux, 1812, p. 184.—Genotype (the only species), Aetea anguina (Sertu-
laria anguina, L., 1758, p. 816).

Aeteopsis, Boeck, 1862, Forh. Vid.-Selsk. Christiania, Aar 1861, p. 49 (also given as
Aetiopsis).—Genotype (the only species), Aeteopsis elongata, Boeck, n. sp.,
which is regarded by Smitt, 1868, p. 280, as a synonym of Aetea truncata,
Landsborough. Synonym of Aetea, Lamouroux, 1812.

Aetiopsis.—See Aeteopsis.

298 SIR SIDNEY F. HARMER ON

Alysidium, Busk, 1852, p. 13.—Genosyntypes, A. parasiticum, Busk, n.sp., and
Eucratea lafontii, Audouin, 1826, p. 242; 1828, p. 74. Genolectotype,
Alysidium parasiticum, see Norman, 1909; J.L. 8. xxx. p, 295.

Amastigia, Busk, 1852?, p. 40.—Genotype (the only species), Amastigia nuda, Busk,.
n. sp.

Anderssonia, Kluge, 1914, p. 617.—Genotype (the only species), Anderssonia antare-
ica, Kluge, n. sp. Synonym of Amastigia, Busk, 1852; but pre-occupied by
Andersonia, Boulenger (Pisces), 1900, A.M. N.H. (7) vi. p. 528, and by
Anderssonia, Strebel (Mollusca), 1908, Wiss. Ergebn. Schwed. Siidpolar-Exp..
vi. li, p. 12.

Anguinaria, Lamarck, 1816, p. 142.—Genotype (the only species definitely men—
tioned), Anguinaria. spatulata, Lmk., nom. nov. Lamarck gives Sertularia
anguina, L. (1758, p. 816) as a synonym, from which it appears that the
introduction of a new trivial name was unnecessary. Hincks (1880, p. 2)
states that Angwinaria was mentioned, but not defined, by Lamarck in 1812 ;
but on consulting this work (Extr. Cours Zool. Mus. Paris, p. 24) it is found
that Lamarck did no more than mention the generic name in the French,
form “ Anguinaire.” Synonym of detea, Lamx., 1812.

Angularia, Busk, 1881, Q. J. M.S. xxi. p. 14.—Mentioned as an abyssal genus with:
a web-like expansion at the angle of most of the bifurcations, No species is
indicated, and the genus does not seem to have been referred to by Busk in
any later work.

Avicella, Van Beneden, 1848, Bull. Acad. roy. Belg. xv. 1, p. 74.—Genosyntypes,
5 species, all referable to Bugula, Oken, 1815. As this is the case, there seems
to be no object in selecting a genotype, which it would be difficult to do in
view of uncertainty as to the identification of some of the species described.

Avicularia (Thompson, MSS.), Gray, 1848, pp. 105, 146.—Genotype (the only
species), Avicularia flabellata (Thompson, MSS.), Gray, n. sp. Synonym of
Bugula, Oken, 1815; but pre-oceupied by Avicularia, Lamarck (Arachnida),
1818, An. s. Vert. v. p. 107.

Bactridiwm, Reuss, 1848, Naturwiss. Abhandl. (Haidinger), ii. p. 55.—Genosyntypes,
B. hagenowt, Reuss, n. sp., and 3 other fossil species. Genolectotype, selected
by D’Orbigny, 1852, P. F. T. C. p..363, and A.S. N. (8) xvii. p. 289, Bactridium
hagenowi, This species does not seem to belong to the Cellularine assemblage,
but the other three species described by Reuss were regarded by D’Orbigny as
species of Canda.

Beania, Johnston, 1840, A.M. N.H. v. p. 272.—Genotype (the only species), Beania
mirabilis, Johnst., n. sp. See also Johnston, 1847, p. 371.

Bicellaria, de Blainville, 1830, p. 423; see also 1834, p. 459.—Genosyntypes,
7 species, including “ B. ciliata, Ellis” (Sertularia ciliata, L., 1758, p. 815).
In his revision of genera, Gray (1848, p. 112) includes B. ciliata by itself in

Bicellaria ; and, although he gives no diagnosis, by placing Sertularia ciliata,
L., at the head of his synonymy, he may fairly be considered to have indicated

CELLULARINE AND OTHER POLYZOA. 299

Bicellaria ciliata as the genotype. Pre-occupied by Drcellaria, Macquart
(Diptera), 1823, Rec. Trav. Soc. Amat. Sci. Lille, Années 1819-1522, p. 155,
and replaced by Bicellariella, Ley., 1909 (q.v.). See also Bugula.

Bicellaviella, Levinsen, 1909, p. 431.--Proposed to replace Bicellaria, de Blainville,
1830, pre-oceupied (see Bicellaria), with genotype Bicellariella ciliata (see
p- 110).

Bicellavina, Levinsen, 1909, p. 99.—Genotype (the only species), Bicellarina aldert
(Bicellaria alderi, Busk, 1860, Q. J. M.S. viii. p. 213).

Bifrons, MacGillivray, 1860, Trans. Phil. Inst. Vict. iv. p. 163.—Proposed to replace
Dimetopia, Busk, 1852, at Dr. Mueller’s suggestion, on the ground that
Dimetopia is used in Botany (Umbellifere). The alteration is not required by
the accepted rules.

Brettia, Dyster, 1858, Q.J. M.S. vi. p. 260.—Genotype (the only species), Brettca
pellucida, Dyst., n. sp.

Bugula, Oken, 1815, p. 89.—Genosyntypes, Cellularia neritina, Cell. ciliata, and
Bugula avicularia (authorities not cited). Linneus (1758, p. 809) founded
Sertularia avicularia primarily on the biserial species figured by Ellis (1755,
pl. 20. fig. A), and to this species the Linnean name is now restricted; but he
also included the pluriserial species of the same author (¢. cit. pl. 38.
fig. 7), though wrongly citing the Plate as 28. Gray (1848) made 8. avicu-
laria (s. str.) the genotype of a new genus, Bugulina (p. 114), while (p. 106)
he described the pluriserial species of Ellis, whose Plate is wrongly cited as
58, as Avicularia flabellata (Thompson, MSS8.), A. flabellata being the genotype.
He appears to have overlooked the fact that Oken described C. avicularia as
haying its zocecia in 3 or 5 series, as he includes Oken’s species in his synonyms
of Bugulina avicularia, whereas it should properly have come under Avicularia
flabellata. B. neritina, B. avicularia, and B. flabellata are congeneric, what-
ever the generle name adopted for them.

In 1819 Schweigger made Sertularia neritina, L., the genotype of Acamarchis
(q. v.), to which S. avicularia, L., and Avicularia flabellata (Thompson, MSS.),
Gray, might also be considered to belong. This would necessitate regarding
S. ciliata, L., as the genotype of Bugula, a course which would introduce the
greatest confusion into nomenelature. I regard this as an instance in which
it is essential to disregard the strict application of the Rules of Nomenclature
and to accept Hincks’ selection (1880, p. 75) of Bugula neritina as the geno-
type of Oken’s genus. As this species is also the genotype of Acamarchis, it
becomes necessary to choose between the two genera. I give the preference
to Bugula, partly because it antedates: Acamarchis by a year, and partly
because it has been universally accepted. As a further motive, it may be
pointed out that A. neritina, in its typical form, is devoid of avicularia, and
that certain writers have accepted Acamarchis as a genus differing from
Bugula by the absence of these organs. See also Bicellaria.

Bugularia, Levinsen, 1909, pp. 99, 108.—Genotype (the only species), Bugularia dis-
similis (Carbasea dissimilis, Busk, 1852°, p. 51).

300 SIR SIDNEY F. HARMER ON

Bugulella, Verrill, 1879, Amer. J. Sci. Arts, (3) xvii. p. 472.—Genotype (the only
species), Bugulella fragilis, Verr., n. sp., described as allied to Bicellaria and
perhaps to Brettia.

Bugulina, Gray, 1848, p. 114.—Genotype (the only species), Bugwulina avicularia
(Sertularia avicularia, L., s. str.). The form indicated by Gray is the biserial
species of Ellis (1755, p. 36, pl. 20. figs. a, A), to which the trivial name of
Linneeus is now restricted. See Bugula, of which this genus is a synonym.

Bugulopsis, Verrill, 1880, Proc. U.S. Nat. Mus. ii. (1879), p. 190.—Genotype, selected
by the author, Bugulopsis peachwi (Cellularia peachii, Busk, 1851, A. M. N. H.
(2) vii. p. 82). Synonym of Z’'ricellaria, Fleming, 1828.

Caberea, Lamouroux, 1816, p. 128.—Genosyntypes, C. pinnata and C. dichotoma,
Lamx., n. spp., both from ‘‘ Australasie.” Genolectotype, C. dichotoma, see
Schweigger, 1819, Tab. 9; 1820, p. 480; and Gray, 1848, p. 147.

Caberiella, Levinsen, 1909, pp. 134, 185.—Genotype (the only species), Caberiella
benemunita (Menipea benemunita, Busk, 1884, p. 19). Synonym of Amastigia,
Busk, 1852.

Caberoides, Canu, 1908, Ann. Paldéont. iil. p. 87 (83, sep.).—Genotype, selected by
the author, Caberoides canaliculata, Canu, n. sp. The genus is described as
resembling Caberea and also members of the Lepralioid series (Hippoporine).

Camptoplites*, n. gen.—Genotype, Camptoplites bicornis (Bugula bicornis, Busk,
1884, p. 40). I propose this genus for Sect. 6 of Bugula as given in Busk’s
‘Challenger’ Report, 1884, p. 37, with the following diagnosis :—

Zoarium stalked, the stalk prolonged basally into an attaching tuft of
rootlets. Zocecia biserial or pluriserial, narrow proximally, the opesia
occupying most of the expanded distal portion. Proximal ends of the
zocecia not forked. Avicularia borne on long, flexible stalks, which usually
exceed the head of the avicularium in length. Operculum distinguishable,
ovicells well developed.

The peculiarities of the avicularium and the absence of a forked proximal
end in the zocecia seem to justify generic separation for this assemblage of
forms, which are characteristic of abyssal depths and of the Antarctic area.
In addition to the type-species and B. reticulata, described by Busk in his
original account, the following species are referable to Camptoplites:—
Bugula tricornis, Waters, 1904, “ Belgica” Bryozoa, p. 23; and the following °
species described by Kluge (1914):—Bugula bicornis, vars. (pp. 619-624),
B. areolaia (p. 627), Bugula sp., var. variospinosa, nov. (p. 628), B. multi-
spinosa (p. 628), B. retiformis (p. 629), B. lewaldi (p. 630), B. gigantea (p. 630),
B. angusta (p. 631), B. abyssicola (p. 632), and B. lata (p. 6384), K1., n. spp.

Canda, Lamouroux, 1816, p. 131.—Genotype (the only species), Canda arachnoides,
Lamx., n. sp.

* kapumtos, flexible ; ém\izns, an armed man; in allusion to the long, flexible stalk of the
avicularium. The genus is masculine.

CELLULARINE AND OTHER POLYZOA. 301

Carbasea, Gray, 1848, pp. 105, 146.—The only species is Carbasea papyracea, but
the synonymy includes the following species :—(1) Eschara papyrea, Pallas
(1766, p. 56), from the Mediterranean, still known by Pallas’ trivial name;
(2) Flustra carbasea, Ellis and Solander (1786, p. 14), from Aberdeen and
Edinburgh. By the rule of absolute tautonymy the genotype is /’. carbasea, and
this conclusion is confirmed by Gray’s mention of specimens from Northumber-
land and Scarborough examined by him in the British Museum Collection.
The genotype has been renamed C. solandert by Norman (1903, A, M. N. H.
(7) xi. p. 582), but there appears to be no valid reason for rejecting its original
name, which should accordingly stand as Carbasea carbasea (Hill. and Sol.).

Caulibugula, Verrill, 1900, Trans. Conn. Acad. x. p. 593.—Genotype, selected by
the author, Caulibugula wrmata, Verr., n. sp. Apparently a synonym of
Stirpariella (q. v.).

Cellaria, Ellis and Solander, 1786, p. 18.—Genosyntypes, 18 species, referable to
about 13 modern genera. Thirteen of these species are identical with
species included by Pallas in Cellularia (q.v.), as is shown by the identical
citations, by both authors, of the work of Ellis (1755). It has generally
been assumed that Cellaria was a gratuitous alteration of Cellularia; but
Ellis and Solander do not refer to Pallas, and it seems possible that their
arrangement was an entirely independent one, based on the works of Ellis
and Linneeus.

In 1801 Lamarck (Syst. An. s. Vert. p. 382) defined Cellaria nearly in its
modern sense, including two species only :—(1) C. salicornia, with synonyms
C. farciminoides, Ell. and Sol. and Tubularia fistulosa, L.; (2) C.cirrata, Ell.
and Sol., which later became the genotype of Menipea (q.v.). He placed
O. salicornia alone in the first Section of the genus, distinguished by having
“ Articulations couvertes de cellules dans tous les sens.” Under Cellaria
Lamouroux (1816, p. 125) moreover writes: ‘‘J’ai conservé le nom de Cellaire
au groupe dont Jes Polypiers avoient pour type le Cellaria Salicorma.”

I think this is a case in which the strict Law of Priority should be set on
one side, in view of the undoubted inconyenience of reverting to the earlier
mame. Norman (1903, A.M.N.H. (7) xi. p. 577) suggested a return to
Cellularia, pointing out that Oellaria is to be regarded as an absolute synonym
of that genus. But, on the other hand, Cellaria had already acquired a
definite signification, which it has since retained; and this cannot be said of
Oellularia, which has been used in the most various senses, and should, in
my opinion, be discarded. In this respect I am in agreement with the
conclusions of D’Orbigny (1851, P. F.T.C. p. 27), Smitt (1868, p. 383),
Hincks (1880, p. 104), Waters (1897, J. L. S. xxvi. p. 3), and others. The
genotype is Hschara fistulosa, L., of which C. fareiminoides, included in Ellis
and Solander’s original account, as well as Cellulariu salicornia (pars), Pallas,
selected by Lamouroux (1816) as the genotype, are synonyms.

The identification of the species to which the Linnean name /istulosa
properly belongs requires further consideration, and the conclusion at which
I have arrived is not the one ordinarily accepted. Hschara fistulosa was

302

SIK SIDNEY F. HARMER ON

introduced by Linnzus (1758, p. 804) in his 10th Edition, with references
to Ellis (1755, p. 46, no. 1, pl. 23. fig. A), followed by citations of Bauhin,
Ray, Plukenet, and Barrelierus. In his 12th Edition (1767, p. 1302)
Linneus describes the same species as Zubularia fistulosa, with one or two
additional citations. Pallas (1766, p. 61), in introducing Céllularia
salicornia, gives all the citations of Linneeus (1758), with others, but he
includes two species under one name. Ellis and Solander (1786, p. 26)
describe Cellariu farciminoides, with citations of Ellis (p. 46, pl. 23) and of
Tubularia fistulosa, L., 1767. It is quite clear that the three trivial names),
of Linnzeus, Pallas (in part), and Solander, respectively, refer to one and the
same species.

The examination of the original works cited by Linneus in 1753 shows,.
however, that his synonymy is open to a good deal of criticism. The figures.
of Ellis refer unquestionably to a species of Cellaria as here understood.
Bauhin and Cherler (1651, Hist. Plant. iii. p. 811) describe two forms, a
coarser species, from the Adriatic, and a more slender ‘ varietas,”’ of which
the locality is not given. The coarser species, to which Linnzus expressly
limits his reference (‘Corallina fistulosa fragilis crassior ”), shows projections
on the internodes which suggest the produced peristomes of Tubucellaria, to
which genus I have little doubt that it belonged. The slenderer species was
probably a Cellaria. Ray (1686, Hist. Plant. 1. p. 65) quotes Bauhin and
Cherler verbatim, without giving additional information. Plukenet (1696,
Almagestum Botanicum, p. 118, pl. 26. fig. 2) also cites Bauhin and.
Cherler. His figure might refer to a Cellaria, but his collection is in the.
Sloane Herbarium at the British Museum (Natural History), and his
specimen, preserved in Vol. 95, Fol. 194 of that Collection, is a Coralline
Alga. Barrelierus (1714, Plante per Galliam, Hispaniam et Italiam observatie,
p. 121) quotes Bauhin and Cherler and Ray. He describes and figures a
coarser and a slenderer form, but the coarser species may be an Alga,
although the other is probably a Cellaria. In view of these discrepancies
and uncertainties it is necessary to regard the citation of Ellis, the first on
the list, as the one to which Linneus’ name really refers; and this conclusion
is confirmed by the fact that he consistently made use of Ellis’ admirable
figures in describing branching Polyzoa, many of his species being introduced
with a citation of Ellis and of no other author.

Ellis explicitly states that there are two species of his ‘‘ Bugle Coralline,”
and he is. equally definite in explaining that his figures a, A, B, and C belong
to the “larger Bugle Coralline.” D, the remaining figure on his Pl. 23,
appears to belong to the same species, though this is not stated quite so
definitely. The mere question of size indicates that Ellis’ figured species is
the one usually described as Cellaria sinuosa, and that Elhs’ smaller species is
the ©. jistulosa of Hincks and of other modern authors.

C. sinwosa was introduced, as Marcimia sinuosa, by Hassall (1840, A.M.N.H.
vi. p. 172, pl. 6. figs. 1, 2), who expressly states that it is larger than what
he calls 7. salicornia (Ellis’ smaller species), and that it is distinguished by
having its apertures in the upper third of the cell. This character is clearly

CELLULARINE AND OTHER POLYZOA 303.

shown by Ellis in his figs. B and D. Hassall later (1843, A. M. N. H. xi.
p- 112) altered the name of his species to /”, spathulosa.

In regarding Cellularia fistulosa as the genotype of the genus I am obliged
to accept the following synonymy :—

J. Cernaria Fisrutosa (L.).

Larger Bugle Coralline, Ellis, 1755, p. 46, pl. 23. figs. a, A-D.

Eschara fistulosa, L, 1758, p. 804.

Cellularia salicornia (pars), Pallas, 1766, p. 61.

Tubularia fistulosa, L. 1767, p. 1302.

Cellularia farciminoides, Ellis and Solander, 1786, p. 26.

Tubularia salicornis (fistulosa) (Cellaria salicornea, Pall.), Esper, between
1805 and 1810, Pflanzenth. iii. p. 103, Tubularia, pl. 2. figs. 1-4 (figures
poor, but Ellis cited in synonymy).

Salicornaria dichotoma, Schweigger, 1819, Tab. 8; 1820, p. 428.

Farcimia sinuosa, Hassall, 1840, A. M.N. H. vi. p. 172, pl. 6. figs. 1, 2.

Farcimia spathulosa, Hassall, 1843, A. M.N. H. xi. p. 112.

Cellaria sinuosa, Hincks, 1880, p. 109.

(nee Cellaria fistulosa, Hincks, 1880, p, 106; and of other authors.)

2, CELLARIA SALICORNIA (Pallas).
Cellularia salicornia (pars), Pallas, 1766, p. 61.
? Cellaria salicornioides, Lamouroux, 1816, p. 127.
Salicornaria farciminoides, Johnston, 1847, p. 355.
Cellaria fistulosa, Hincks, 1880, p. 106; et auett.

Pallas divided ©. salicornia into two Sections:—(a) the larger Bugle
Coralline, as shown by his citation of Ellis, Pl. xxiii.; (8) a more slender
form, characterized in his synonymy as “subtilior” and ‘“‘ tenuior,” in contrast
with ‘“crassior” of his first Section. He gives Hschara fistulosa, L., as a
synonym of /3, but this was not admissible, in view of Linnzus’ citations of
Ellis’ figure of the larger Bugle Coralline and of the “Corallina fistulosa
fragilis crassiot” of Bauhin and Cherler, emphasizing the fact that he had
the coarser form in mind. This conclusion is not modified by reference to
Linneus’ “ Fauna Svecica,” 1761, no. 2232, which Pallas wrongly cites as.
2234. Pallas includes Ellis “ Angl. Bugle Coralline” at the end of his
Sect. 3, and this may be taken as the determining factor. As his trivial name
is not applicable to the larger Bugle Coralline, it may be used for the smaller
form. The adoption of @. salicornioides for this species would be open to some
uncertainty, as though Lamouroux also had a slender species in view, the
locality, presumably of specimens in his own collection, is given as Mediter-
ranean. This suggests the possibility that his specimen, if it exists, may
prove to belong to C. (Wellia) johnsoni, Busk (1858, Q. J. M.S. vi. p. 125), the
typical locality of which is Madeira, and not to C. fistulosa, auctt.

Cellarina, Van Beneden, 1848, Bull. Acad. Roy. Belg. xv. 1, p. 70.—Genosyntypes,
C. gracilis and C. scabra, Van Ben., n. spp., and Crisia delilii, Audouin, 1826,
p- 242. Norman (1903, A. M.N.H. (7) xi. p. 579), who had examined a
fragment of Van Beneden’s type-specimen of the first species, described it as.

304 SIR SIDNEY F. HARMER ON

Menipea gracilis, but admitted its identity with Cellularia ternata, forma
gracilis, Smitt (1868, p. 283). Whether C. gracilis is to be regarded as a
species or as a variety, it clearly belongs to Zricellaria, Fleming, 1828; while
the other two species are referable to Scrupocellaria, Van Beneden, 1845.

Cellarina, D’Orbigny, 1851, P. F.'T.C. p. 181; see also 1852, A.S.N. (3) xvi.
p. 336.—Genosyntypes, two fossil species. Pre-occupied by Oellarina, Van
Beneden, 1848.

Cellularia, Pallas, 1766, p. 58.— Genosyntypes, 18 species, referable to about
12 genera. The name has been used by many modern authorities, but in:the
most various senses, and it has been impossible at present to come to any
agreement as to the use which should be made of it. The selection of
C. scruposa as the genotype, by Verrill (1880, Proc. U.S. Nat. Mus. 11. (1379)
p- 190) is-invalidated by the fact that this species was already the genotype
of Scrupocellaria, Van Beneden, 1845. I consider it desirable to suppress
Cellularia in favour of Cellaria (q. v.).

Cercaripora, Fischer, 1866, C. R. Acad. Sci. Paris, xii. p. 987; Nouv. Arch. Mus. Hist.
Nat. ii. p. 312.—Genosyntypes, Angwnaria truncata, Landsborough, 1852,
Hist. Brit. Zooph. p. 288; Aetea ligulata, Busk*, p. 31; and Aetea argillacea,
Smitt, 1866, Ofv. K. Vet.-Akad. Forh. xxii. (1865) p. 29. The genus was
placed in a different Family from that containing detea, which Fischer
restricted to Sertularia anguina, L. (1758, p. 816); but later authors have
regarded it as a synonym of Aetea, of which this species is the genotype.

Chaperia, Julhen, 1881, Bull. Soc. Zool. France, vi. pp. 163, 164.—Although the
author mentions AMembranipora (Steganoporella) magnilabris, Busk (1854,
Brit. Mus. Cat. ii. p. 62), as belonging to Chaperia, the genotype selected by
him on p. 164 is Chaperia australis, nom. nov., to replace M. spinosa,
Q. and G., as quoted by Busk (1879, Phil. Trans. vol. 168, p. 195), in
describing specimens from Kerguelen. Jullien gives a description of
C. australis, from specimens obtained at the Cape of Good Hope. But Busk’s
citation was a mistake, the species described by Quoy and Gaimard (1824,
Zool. Voy. Uranie et Physicienne, p. 605) from the Falkland Is. having been
named by them Flustre épineuse, Mlustra acanthina; see Waters, 1898,
J. L.S. xxvi. p. 673; see also Marcus, 1922, Goteborgs K. Vet.- och Vitt.-
Samh. Handl. (4) xxv. p.6. Jullien’s name australis was proposed on the
ground that spinosa was pre-occupied; but if C. australis is synonymous with
Quoy and Gaimard’s species, the name of the genotype should be Chaperna
acanthina (Q. and G.).

Chartella, Gray, 1848, pp. 104, 145.—Genotype (the only species), Chartella
papyracea (Flustra papyracea, Ellis and Solander, 1786, p. 13).

‘Chaunosia, Busk, 1867, Q.J. M.S. (x.8.) vii. p. 241.—Genotype (the only species),
Chaunosia hirtissima, Busk, n, sp. Although Busk expressly marks his
species as “n. sp.,” he adds that it is not unlikely to be identical with
Diachoris hirtissima, Heller (1867, Verh. Zool.-Bot. Ges. Wien, xvii. Abhandl.
p. 94). Synonym of Beania, Johnston, 1840.

CELLULARINE AND OTHER POLYZOA. 305

Chiidonia, Lamouroux, 1824, p. 192.—Lamouroux states that Savigny’s species (see
below) appears to be the Vorticella polypina “des auteurs,” and the author
he had specially in mind may well have been Esper, who uses this name in
Th. ii. of the Forsetz. d. Pflanzenth., Vorticella, Pl. 1. figs. 1, 2. The text of
this part of the ‘‘ Fortsetzungen” ends with p. 48, and it contains no
description of the plate in question. It was published in two Lieferungen,
9 and 10, in 1798 and 1806, respectively. The plate presumably appeared
between 1798 and 1810, the date of Esper’s death.

The name ‘“Chlidonies” was used by Sayvigny (Description de Egypte), at
the foot of his Pl. 13, in which figs. 3'-3° give admirable representations of
what is almost certainly the same species as Vorticella polypina, Esp-
Audouin, in his ‘‘ Explications” (1826, p. 243) of Savigny’s plates, did not,
however, accept Savigny’s name, as he describes the species figured as
Eucratea cordiert. Wamarck (1816, p. 140) introduced a new name, Cellaria
vesiculosa, with Vorticella polypina, Esp. as a synonym, but with a query; and
Hammer (1829, in Esper, Pflanzenth. i. Lief. 16, p. 255) uses Hucratea
vesiculosa in describing Esper’s plate. Lamarck’s trivial name antedates
Audouin’s, but Bertoloni (1810, Rar. Ital. Plant., Decas Tertia, p. 112; see
also 1819, Amoen. Ital. p. 278) had previously described the same species
as Cellaria pyriformis, in both papers giving a recognizable description of
Ohlidonia, based on actual specimens. His 1810 synonymy refers to Vorticella
polypina, L. (1767, p. 1317), an Infusorian. In 1819 he cited Hsper’s figures,
thereby making his descriptions more intelligible; while by rejecting his
earlier synonymy he furnished a justification for the introduction, otherwise
invalid, of a new trivial name in 1810.

The genotype thus appears to be Chlidonia pyriformis (Bert.), of which the
other names indicated above are synonyms. Chlidonia, Lamx., 1824, ante-
dates both Chiidonia, Hiibner, 1825—-1826* (Verz. bekannt. Schmetterl. p. 393)
and Chlidonia, Herrich-Schiiffer, 1838 (in Panzer, Deutschl. Insecten, Heft
157), two genera of Insects. Chlidonias was introduced by Rafinesque, 1822
(Kentucky Gazette, xxxvi. (8) p. 3), and this name is regarded as valid by
Ornithologists. There seems to be no inconvenience in retaining Chlidonia
for Polyzoa, in spite of its close resemblance to the name of a genus of Birds.

Oinetoscias, von Martens, 1879, Zool. Rec. for 1877, xiv. Molluscoida, p. 94.—An
emendation proposed by von Martens, in recording the introduction of
Kinetoskias (q.v.). Although the name might have been thus spelt it is
undesirable to alter the accepted original form.

Columnaria, Levinsen, 1909, p. 116.—Introduced for Columnaria borealis, n. sp., and
all the species of Furciminaria described by Busk (1884, pp. 48-51), except
F atlantica. No genotype was selected, but this is of small importance, since
the name is pre-occupied, in Anthozoa, by Columnaria, Goldfuss, 1826, Petr.
German. i. pt. 1, p. 72 (for date of publication see Woodward, vol. 11. p. 692).
I prefer not to suggest a new name without making a study of the species
involved.

* For the date of publication see Proc. iy. Int, Congr. Zool. (Cambridge, 1898), 1899, p. 299.

306 SIR SIDNEY F. HARMER ON

Cornucopina, Levinsen, 1909, pp. 98, 109.—Levinsen states that this genus includes
most of the species of Bicellaria, auctt., but the only species he definitely
refers to it (pp. 110, 372) are Bicellaria grandis, Busk (18521, p. 3874; 1852°*,
p. 42) and B. znfundibulata, Busk (1884, p. 33). Of these the latter is
exceptional in certain characters, and I think it more convenient to select
Cornucopina grandis as the genotype.

‘Corynoporella, Hincks, 1888, A. M.N. H. (6) i. p. 215.—Genotype (the only species),
Corynoporella tenuis, Hincks, n. sp., described as allied to Bugula.

Cothurnicella, Wyville Thomson, 1858, Nat. Hist. Rev. v., Proc. of Societies, p. 141.—
Genotype (the only species), Cothurnicella daedala, Wyv. Thoms., n. sp., a
synonym of Cellaria pyriformis, Bert., the genotype of Chiidonia, Lamx.,
1824.

Craspedozoum, MacGillivray, 1886, Trans. Proc. R. Soc. Vict. xxii. p. 131.—Geno-
syntypes, Membranipora roborata, Hincks (1881, A. M. N. H. (5) viii. p. 128);
CO. ligulatum, C. spicatum, MacG., n. spp., and Plustra membraniporides, Busk
(1884, p.54). As I regard this genus as asynonym of Menipea, Lamx., 1812,
it seems unnecessary to select a genotype.

Orepis, Jullien, 1882, Bull. Soc. Zool. France, vii. p. 522.—Genotype (the only species),
Orepis longipes, Jull., n. sp.

‘Orisia, Lamouroux, 1812, p. 183.—-Genosyntypes, Sertularia eburnea, L. (1758,
p. 810), a Cyeclostome, and 5 species of Cheilostomes, including Sertularia
ciliata, L. (1758, p. 815). Under Hucratea I suggest ignoring Schweigger’s
selection of C. ciliata as the genotype. If this proposal is accepted, it will be
possible to consider that Fleming (1828, p. 540) selected Crista eburnea as the
genotype, by including it, with another species not appearing in Lamouroux’s
original account, in Crisia with an amended diagnosis.

Orisularia, Gray, 1848, pp. 111, 147.—Genotype (the only species), Orisularia
fastigiata (Sertularia fastigiata, L. 1758, p. 815 = Cellularia plumosa, Pallas,
1766, p. 66). Synonym of Bugula, Oken, 1815.

Dendrobeania, Levinsen, 1909, pp. 99, 113.—Genotype (the only species), Dendro-
beania murrayana (Flustra murrayana (Bean, MSS.), Johnston, 1847, p. 347.
Synonym of Bugula, Oken, 1815.

Diachoris, Busk, 1852’, p. 381.—Genotype (the only species), Diachoris crotal, Busk,
n. sp. See also Busk, 1852°*, p. 54. Synonym of Beania, Johnston, 1840.

Diachoseris, Ortmann, 1889, Arch. f. Naturg. lvi. i. p. 25.—The name appears to be a
misquotation of Diachoris, Busk, 1852. The following species are included :—
Diachoris magellanica, Busk (1852*, p. 54), Diachoseris discodermiw, and
D. hewvaceras, Ortm., n. spp. Synonym of Beania, Johnston, 1840.

Didymia, Busk, 1852', p. 383.—Genotype (the only species), Didymia simplex,
Bk., n.sp. See also Busk, 1852°, p. 35. The genus being pre-occupied, in
Hymenoptera, by Didymia, Le Peletier and Serville, 1825, Eneyel. Méthod. x.
(Entomologie), p. 574, I propose to replace it by Didymozoum, nom, nov., with
the genotype Didymozoum simplex (Busk).

CELLULARINE AND OTHER POLYZOA. 307

Didymozoum, nom, nov.—See Didymia.

Dimetopia, Busk, 18521, p. 384.—Genosyntypes, D. spicata and D. cornuta, Busk,
n. spp. See also Busk, 18527, p. 35. In 1909 Prof. A. Billard submitted to
me a specimen of Dynamena barbata, Lamx. (1816, p. 178), from Lamouroux’
type-collection at Caen; and I convinced myself that D. spicata, Busk, was a
synonym of this species (see Billard, C.R. Acad. Sci. Paris, 1909, exlviii.
p- 1064). In the apparent absence of an earlier selection I propose Dime-
topia cornuta as the genotype. It appears to me preferable to select a species
of which good figures were published by Busk, rather than to choose D. barbata,
the identification of which with D. spicata rests merely on my own authority.

Dimorphozoum, Levinsen, 1909, pp. 96, 107.—Genotype (the only species),
Dimorphozoum nobile (Flustra nobilis, Hincks, 1891, A.M.N.H. (6) vii.
p. 288).

Diplodidymia, Reuss, 1869, Sitzb. Akad. Wiss. math.-nat. Cl. lix. i. Abth. p. 468.—
Genotype (the only species), Diplodidymia complicata, Reuss, n. sp. Synonym
of Poricellaria, D’Orbigny, 1852.

Dittosaria, Busk, 1866, Geol. Mag. iii. p. 301.—Genotype (the only species),
Dittosaria wetherellii, Busk, n. sp., a fossil, from the London Clay, perhaps
allied to Sertularia loricata, L., the genotype of Hucratea (q. v.).

Emma, Gray, 1848, p. 293.—Genotype (the only species), Hmma crystallina, Gray,
n. sp. Gray gave no generic diagnosis, but this was done by Busk, 1852",
p- 372, and 1852°, p. 27.

Epistomia, Fleming, 1828, p. 541.—Genotype, selected by the author, Hpistomea
bursaria (Sertularia bursaria, L., 1758, p. 814). See Gregory, 1893, Trans.
Zool. Soe. xiii. p. 227. See also Notamia.

Erina, Cana, 1908, Ann. Mus. Nac. Buenos Aires (3) x. p. 273.—Genotype, selected
by the author, Hrina patagonica, Canu, 1. sp., placed in Meliceritide. Pre-
occupied by Hrina, Swainson, 1833, Zool. Tlustr. (2) iii. pl. 134 (Lepidoptera).

Eueratea, Lamouroux, 1812, p. 183.—Genosyntypes, Cellaria cornuta (Sertularia
cornuta, L., 1758, p. 810) and C. loricata (Sertularia loricata, L., 1758, p. 815).
In the same Memoir Lamouroux introduced the genus Crista (q. v.). Hach of
these genera included one species, S. cornuta and S. eburnea respectively,
belonging to Crisia as universally understood at the present time. It is
unfortunate that Schweigger (1819, Tab. 8; 1820, p. 429) indicated
S. cornuta as the genotype of Hucratea, and S. ciliata as the genotype of
Crisia ; both of Lamouroux’ genera being regarded by him as subgenera of
Cellularia. By adopting these selections Bicellaria, auctt., would become a
synonym of Orisia, and Crisia, auctt., of Mucratea. These alterations would
be so inconvenient and confusing that I feel justified in suggesting that
Schweigger’s selections should be ignored. If Crisia is to be maintained, its
genotype must be Orisia eburnea (Linn.), and S. cornuta will fall within the
same genus; Hucratea loricata (Linn.) thus remaining as the genotype of
Eucratea. S. chelata, L. (1758, p. 816), which has usually been regarded as
the genotype of Hucratea, has no claim to this position, as it was not included
in Lucratea by Lamouroux until 1816 (p. 149).

308 SIR SIDNEY F. HARMER ON

Eucratia, Fleming, 1828, p. 541.—Used by mistake for Hucratea (q. v.).

Euoplozoum*, n. gen.—Genotype, Huoplozoum cirratum (Cellularia cirrata, Busk,.
1884, p. 17).

Zoarium erect, attached by a basal tuft of rootlets. Branches biserial, the
zocecia large and obliquely alternating, all facing in the same direction, their
proximal portions narrow, their distal portions expanded, the frontal membrane:
occupying the entire surface of the expanded portion. Basal walls of zocecia
strongly convex. Branches traversed by oblique joints, each zocecium being
crossed by one joint near its proximal end and by another at the commence-
ment of its expanded portion. Avicularia of two kinds, the smaller at the
distal outer border of the zocecium, and a very large kind (present on only a
few of the zoccia) arising from the inner border. Strong flexor zocecii
muscles present, by which the branches are inflected. Ovicells very large..
Bifurcation of type 6 (Pl. 16. fig. 6).

It can hardly be doubted that this very remarkable species, which was:
placed by Busk in a heterogeneous assemblage referred to Cellularia, deserves
recognition as the type of a new genus. It was described recently, by the
late Miss Alice Robertson (1921, Rec. Ind. Mus. xxii. p. 39), as Kinetoskias
arabianensis, Roberts., n. sp.

Euthyris, Hincks, 1882, A. M. N. H. (5) x. p. 164.—Genosyntypes, Z. obtecta, Hincks,,
n. sp., Hlustra bombycina, Ellis and Solander (1786, p. 14) and Carbasea
episcopalis, Busk (18522, p.52). C. epzscopalisis the genotype of Huthyroides,.
Harmer, 1902. F. bombycina Ell. and Sol. (nec Busk; see Onchoporella),.
which appears to be at present unrecognizable, is the genotype of Semaflustia,
D’Orbigny, 1852. Huthyris obtecta is thus left as the genotype of Luthyris,
but if 7. bombycina could be recognized it would probably become necessary to-
regard Huthyris as a synonym of Semiflustra (q. v.).

Euthyroides, Harmer, 1902, Q. J.M.S. xlvi. p. 280.—Genotype, selected by the
author, Euthyroides episcopalis (Carbasea episcopalis, Busk, 1852°, p. 52).

Falcaria, Oken, 1815, p. 91.—Genosyntypes, Cellularia falcata (Pallas, 1766, p. 76),
s. cornuta (Sertularia cornuta, L., 1758, p. 810), C. eburnea (S. eburnea, L.,
1758, p. 810) and C. anguina (S. anguina, L., 1758, p. 816); the first two
referable to Crista, and the third to dAetea, two genera introduced by
Lamouroux in 1812. Gray (1848, p. 136) selected Halcaria cornuta as the
genotype. Synonym of Crisia, Lamx., 1812, and pre-occupied in Lepidoptera
by Falearia, Haworth, 1809, Lepid. Britann. p. 152; for date of publication
of which see Woodward, vol. 1i. p. 804.

Farcimia, Fleming, 1828, p. 534.—Genotype (the only species), Furcimia fistulosa
(Eschara fistulosa, L., 1758, p. 804). There is no justification for including
Nellia oculata, Busk (see Nellia) in this genus, as has been done by Waters
(1887, A. M. N. H. (5) xx. p. 92) and others. Synonym of Cellaria, Ell. and
Sol. 1786.

* evomAos, well armed.

CELLULARINE AND OTHER POLYZOA. 309

Farciminariu, Busk, 18527, p. 32.—Genotype (the only species), Farciminaria aculeata,
Busk, n. sp.

Filicella, Searles Wood, 1844, A. M.N.H. xiii. p. 15.—Genotype (the only species),
Filicella anguinea, S. Wood, n. sp., a Crag species, possibly the aduate part of
a species of Aetea.

Fistulana, O. F. Miller, 1776, Zool. Dan. Prodr. p. 282.—In his Corrigenda (p. 282),
Miiller states that Fistularia, introduced in the same work, is to be changed to
Fistulana, a course which was probably adopted because Fistularia was pre-
occupied, in Pisces, by a Linnean genus (1758, p. 312) of the same name.
Fistularia was introduced by Miller, without figures, on p. 254, with the
genosyntypes /. ramosa (T'ubularia ramosa, L., 1758, p. 804; 1767, p. 1302),
Ff. muscoides (I. muscoides, L., 1767, p. 1302), F. simplex, F. longicornis and
F. multicornis, Mill.,n. spp. In 1780, Fabricius (Fauna Groenl. p. 441) uses
Fistulana for F. ramosa and Ff. muscoides, quoting the diagnoses of Miller
verbatim, in words identical with those of Linneus, 1767. In 1789,
Abildgaard (in Miller, Zool. Dan. i. p. 15) describes and figures F. multi-
cornis, quoting Miiller’s diagnosis, with additions.

Of Miiller’s species, the first two are Hydroids, while the diagnosis of
F. longicornis suggests a Polychsete (Spionid). F. simplex may be a Ctenostome
Polyzoon, while it seems probable, from Abildgaard’s description and figures
(pl. 90. figs. 1-3) that #. multicorms was also one of the Polyzoa. Smitt
(1868, pp. 279, 280) suggests that it might be identical with Aetea truncata,
Landsborough (see Cercaripora), but he points out, as an objection to this
conclusion, that Miller had described the tentacles as 30 in number. This is
not quite correct, as Miiller described them as being “ultra viginti,” and it
was Abildgaard who stated that they were “ad triginta.” In an earlier paper
Smitt (1865, Ofv. K. Vet.-Akad. Férh. xxii. (1865), p. 13, pl. 3. fig. 4) had
shown that Aetea truncata has 12 tentacles, and on this ground I think that
F. multicornis cannot have been that species. Abildgaard’s figures seem more
likely to have been taken from a species of Wolella, Gosse (Cylindrecium,
auctt.; see Harmer, 1915, Siboga Rep. Polyzoa.i. p. 52). Fistulana should
be disregarded for Polyzoa. The same name was introduced in other groups.
by Bruguiére (1789) and Lamarck (1799); see Sherborn, Index Animalium, i.
1902.

Fistularia, O. F. Miller, 1776, p. 254.—See Mistulana.

Flabellaria, Gray, 1848, pp. 106, 146.—Genosyntypes, Sertularia spiralis, Olivi,
1792 (Zool. Adriat. p. 291, pl. 6. figs. 2, a, a) and Flustra setacea, Fleming,
1828 (p. 586), referable to Caberea. Bugula murrayana was given as a
synonym of Olivi’s species ; but the occurrence of this northern form in the
Adriatic seems to be very unlikely, and Oliyi’s species ought probably to be
placed elsewhere in Bugula. Gray’s genus thus appears to be unnecessary.
It may be remarked that Mlabellaria, Lamarck, 1816, p. 342, is used for
certain calcareous Alge.

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 23

310 SIR SIDNEY F. HARMER ON

Flabellaris, Waters, 1898, J. L.S. xxvi. p. 672.—Based on several recent species, but
with no clear indication which of them are to be included. Alenipea flabellum
(on which the generic name is obviously founded) is specially mentioned ; and
a description is given of Membranipora roborata, Hincks (1881, A.M. N. H.
(5) viii. p. 128), which is referred to the genus. One of these two species
might be selected as the genotype, but it seems unnecessary to do so, since the
species mentioned on pp. 672, 673 of Waters’ paper, in addition to those
referred to by him, as given in one of his earlier papers (1897, J. L.S. xxvi.
p- 2), can all be placed in other genera which antedate /abellaris.

Flabellina, Levinsen, 1902, Vid. Medd. naturh. Foren. Copenhagen, p. 21.—An
emendation of Jlabellaris, Waters, perhaps suggested because of the
resemblance of this name to Flabellaria, Gray, 1848. The only species
mentioned is labellina roborata (Hincks), 1881 (see Flubellaris); but the name
‘is pre-occupied by /lubellina, Voigt, 1834, Das Thierreich, iii. p. 124, used for
a Nudibranchiate Mollusc.

Flustra, Linneus, 1761, Fauna Svecica, p. 539.—The history of this name has been
given by Lang (1917, Geol. Mag., Dec. vi, vol. iv. p. 170), who shows that
Linneus deliberately altered his own genus Eschara (1758, p. 804) to Flustra,
and that Flustra foliacea, Linn. (Eschara foliacea, 1758, p. 804) is the genotype
of both genera. Although this course is not admissible under the Rules, I
fully agree with Dr. Lang that it is in the highest degree desirable to suppress
Eschara and to use Flustra in its accepted sense. It may be noted that
Lamarck (1801, Syst. An.s. Vert. p.383) accepted /Justra, with the genotype
Flustra foliacea, L.

Gemellaria, Van Beneden, 1845, Nouv. Mém. Acad. Roy. Brux. xviii. p. 9.—Based on
Savigny’s name ‘“ Gémellaires,” appearing at the foot of pl. xiii (referring to
figs. 47-4°) of the “Description de I’Egypte.” Savigny’s species was
described by Audouin (1826, p. 243) as Loricaria egyptiaca, without reference
to the fact that Zoricaria was used for a Fish hy Linneus (1758, p. 307), as
pointed out by Fleming (1828, p. 541). Lamouroux (1827, p. 434) mentions
Gemellaria, without any associated trivial name, but only to state that it is
referable to Loricaria. Gregory (1893, Trans. Zool. Soc. xiil. p. 227) discusses
the generic name, but de Blainville (1830, p. 425; see also 1834, p. 461),
whom he quotes as the first author to mention the genus in a correct form,
merely places Gemellaria loriculata in his synonymy of Gemicellaria loriculata,
ascribing the combination wrongly to Savigny. The first use of the generic
name which is completely in order appears to be that of Van Beneden,
1845, who describes Gemellaria loriculata (Cellularia loriculata, Pallas, 1766,
p. 64 = Sertularia loricata, L., 1758, p. 815); and in this sense the genus has
been used by the majority of recent writers. Gemellaria thus becomes a
synonym of Hucratea (q. v.). It is in any case desirable that it should drop

- out of use, since it is based on Savigny’s ‘‘ Gémellaires,” and the species
figured by this author is not congeneric with S. loricata, L. (See Synnotum.)

Gemicellaria, de Blainville, 1830, p. 425; see also 1834, p. 460.—Genosyntypes,
Gemicellaria loriculata (see Gemellaria), among the synonyms of which

CELLULARINE AND OTHER POLYZOA. 311

’

appears ‘“ Gemellaria loriculata, Savigny,” a combination which was used
neither by Savigny nor by Audouin; and G. bursaria (Sertularia bursarca, L.,
1758, p. 814). These two species are respectively the genotypes of
Eucratea, Lamx. 1812, and Epistomia, Fleming, 1828. G'emicellaria, which
was an alteration of the genus proposed by Savigny, ‘‘sous le nom de
Gemellaria” (really ‘‘Gémellaires”’), thus becomes synonymous with two
genera of earlier introduction.

Halophila, Gray, 1843, p. 292.—Genotype (the only species), Halophila johnston,
Gray, n. sp. (thus written, although frequently modified later, even by Gray
himself, to johnstonie). If this species is referred to Bugula, as I think is
proper, Halophila becomes a synonym of that genus.

Heterocella, Canu, 1907, Ann. Paléont. ii. p. 70 (sep. p. 14).—Genotype, selected by
the author, Heterocella fragilis (Vincularia fragilis, Defrance, 1829, Dict. Sci.
Nat. lyiti. p. 214). See also de Blainville, 1834, p. 454.

Heteroflustra, Levinsen, 1909, pp. 124, 125.—A very unsatisfactory genus, proposed
for those species of Flustra, auctt., which have not been placed in other genera,
and ‘must provisionally be characterized mainly in a negative way.” This
appears to mean that all species not belonging to Mlustra (s. str.), Sarsiflustra,
Kenella, Retiflustra, and Spiralaria (genera recognized by Levinsen, p. 88),
together with other genera such as Carbasea, Chartella, ete. (not considered
by him), are to be placed in this provisional genus, which has no validity
until it is more definitely characterized.

Hiantopora, MacGillivray, 1887, Tr. Proce. R. Soc. Vict. xxiii. p. 208.—Genotype
(the only species), Hiantopora ferox (Lepralia ferow, MacGillivray, 1869,
Ibid. ix. p. 132).

Himantozoum *, n. gen.—Genotype, Himantozoum mirabile (Bugula mirabilis, Busk, |
1881, Q. J. M.S. xxi. p. 12; 1884, p. 39). :

I propose this genus for Sect. a of Bugula, as given by Busk, 1884, p. 37,
with the following diagnosis :—

Zoarium stalked, the stalk composed of rootlets and prolonged into an
attaching tuft of rootlets. Zocecia biserial to multiserial, the biserial branches
composed of asymmetrical zocecia, between which are intercalated, in the
pluriserial branches, one or more rows of median zocecia, which are sym-
metrical and some of which produce eggs. The oyicells are vestigial, and the
ege, which is of large size, develops in the body-cavity of the fertile zocecium.
Opesia occupying all or most of the front, an operculum being distinguishable.
Zocecia overlapping their predecessors on the basal surface, their proximal
ends strongly forked. Avicularia unstalked, attached to the proximal ends of
the zocecia, those of the lateral and median rows more or less unlike.

The differentiation of median, symmetrical, fertile zocecia, as well as the
characters of the avicularia, are striking features of this genus. In addition
to the genotype, the following species may be included :—Bugula leontodon,
Busk (1884, p. 39), B. sinuwosa, Busk (1884, p. 39), B. margaritefera, Busk

* imas, a strap, in allusion to the strap-shaped branches.
QQ*

312 SIR SIDNEY F. HARMER ON

(1884, p. 41) and B. sinuosa, Busk, var. variabilis, Kluge (1914, p. 632).
Most of the species are from abyssal depths, the shallowest record being that
of Busk, 80-150 fathoms, for B. sinuosa.

The statements in the diagnosis referring to the eggs are based on the
examination of Siboga material. B. versicolor, a mémber of Busk’s Sect. a
(1884, pp. 37, 38), differs from the other three species in certain points which
appear to be important, and I do not feel justified in placing it in Himantozowm,
although I am unable to make any other suggestion. The most important of
its characters are the large endozowcial ovicells, the unforked proximal
ends of the zocecia, which barely overlap their predecessors, and the absence
of avicularia.

Hoplitella, Levinsen, 1909, pp. 185, 136.—Genotype (the only species), Hoplitella
armata (Carbasea armata, Busk, 18522, p. 50).

Huzleya, Dyster, 1858, Q. J. M. 8. vi. p. 260.—Genotype (the only species), Hualeya
fragilis, Dyst., n. sp.

Jubella, Jullien, 1882, Bull. Soc. Zool. France, vil. p. 519.—Genotype (the only
species), Jubella enucleata, Jull., n. sp., described as being near Cuberea, but
differing from it by the absence of vibracula.

Kenella, Levinsen, 1909, p. 124.—Genotype (the only species), Kenella biseriata
(Flustra biseriata, Busk, 1884, p. 54).

Kinetoskias, Danielssen, 1868, Forh. Vid.-Selsk. Christiania, Aar 1867, p. 23.—
Genosyntypes, K. arborescens and K. smithi, Dan.,n. spp. In their detailed
account, Koren and Danielssen (1877, Faun. Litt. Norv. Pt. 3, pp. 104, 109)
describe the same two species, the latter as K. smittii, but without comment
on the altered spelling. There can be no doubt that this species was named
after Prof. Smitt, and the form smzthz should be considered a printer’s error.
K. arborescens is not a completely typical member of the genus, as the stalk is
very short and the avicularia are more Bugula-like than usual. I propose
accordingly to regard Kinetoskias smittii as the genotype.

Loricaria, Lamouroux, 1821, p. 7.—Genosyntypes, L. europea and L. americana,
Lamx., n. spp., both synonymous with Hucratea loricata, L. (q. v.). Pre-
occupied by Loricaria, L. (Pisces), as pointed out by Fleming, 1828, p. 541.
See Gemellaria and Hucratea.

Loricula, Templeton, 1836, Loudon’s Mag. Nat. Hist. ix. p. 469.—This genus is
usually attributed to Cuvier, who used it (1830, Régne An. Nouv. Ed. iii.
* p. 303) merely as “ Les Loricules,” proposing it for Sertularia loricata, L.
(1758, p. 815), on the ground that ‘“ Loricaires ” Lamx. (see Zoricama) was
pre-oceupied for Fishes. The genus was used in a correct form by Templeton,
for Loricula loricata; and, later, by Voigt, 1848, Das Thierreich, vi. p. 248, in
the same combination. It is synonymous with Hucratea, Lamx. 1812, with
the same genotype, but it is pre-occupied by Loricula, Curtis, 1833, Ent. Mag.
i. p. 197, for Hemiptera.
Maplestonia, MacGillivray, 1885, Tr. Proc. R. Soc. Vict. xxi. p. 92.—Genotype (the
only species), Maplestonia cirrata, MacG., n. sp.

CELLULARINE AND OTHER POLYZOA. 313

Melicerita.—The introduction of this genus by Milne Edwards (1836, A. 8. N. Zool.
(2) vi. p. 847), with only one species given merely as “‘ Mélicérite de
Charlesworth,” was not strictly in order. Searles Wood (1844; see Ulidiwm)
quotes this as Melicerita charlesworthi, which was thus regularized, and became
the genotype. His simultaneous introduction of Ulidiwm, with the same
genotype, was unnecessary. Inthe A.S. N. (3) xvil., dated 1852, but probably
published later, D’Orbigny refers to the species as M/. charlesvorthir.

Melicertina, Ehrenberg, 1839, Phys. Abh. Akad. Wiss. Berlin, J. 1838, Tab. 11.—
Proposed to replace Melicerita, which “ist nicht sprachgemiiss.” This
emendation need not be accepted.

Membranicellaria, Levinsen, 1902, Vid. Medd. naturh. Foren. Copenhagen, 1902,
p- 22 n.—Introduced for Melicerita dubia, Busk (1884, p. 97) and a number
of Cretaceous species, which are indicated. Genotype, now selected, Membrani-
cellaria dubia (Melicerita dubia, Busk). Leyinsen subsequently (1909, p. 207)
described M. dubia, from part of Busk’s original material, but he gives
M. dubia, Busk, as a doubtful synonym. His reason for inserting a query is
not apparent, but if it should prove that two species were included in Busk’s
material the form described by Busk would be the genotype, as no other was
indicated in Levinsen’s original account.

Menipea, Lamouroux, 1812, p. 183.—Genosyntypes, Cellaria cirrata, Ellis and
Solander (1786, p. 29), quoted by Lamouroux as cirrhata, and C. flabellum,
Ell. and Sol. (1786, p. 28). Genolectotype, Menipea cirrata: see Schweigger,
1819, Tab. 8; 1820, p. 428. The reasons given by Marcus (1922,
p. 11) for regarding M. cirrata as a synonym of Cellularia crispa, Pallas
(1766, p. 71) seem to me sufficient. Pallas gave the locality as “ Oceanus
Orientalis,” but on p. 72 he states that it was found with a Fucus qualified
by the adjective ‘‘ capensis,” which presumably meant the Cape of Good Hope,
as explicitly stated in the German Edition (1787, 1. p. 107) of Pallas’ work.
Seba’s figure (Thesaurus, iii. Pl. 101, No. 8) is hardly demonstrative, as it is
stated to be by Marcus, but it may have referred to the species which was
later described by Pallas, who gives this reference. The correct name of the |
genotype appears to be Menipea crispa (Pall.). Waters’ selection (1897,
J. L. 8. xxvi. p. 2) of WM. buskit, Wyv. Thoms. (see under- Hmma, in Sect. V.)
is obviously invalidated by the fact that this species was not included by
Lamouroux.

Mononota, Pieper, 1881.—See Synnotwm.

Monsella, Canu, 1900, Bull. Soc. Géol. France (8) xxviii. p. 437.—Genotype, selected
by the author, Monsella eocena (Planicellaria eocena, Meunier and Pergens,
1886, ‘‘Les Bryozoaires du Syst¢me Montien (Eocéne Inférieur),” Louyain,
privately printed, p. 7). Canu gives figures of this species, copied from
Meunier and Pergens.

Naresia, Wyville Thomson, 1873, Nature, vii. p. 388; see also Humbert, 1874,
J. de Zool. iii. p. 184 and Wyville Thomson, 1877, “The Voyage of the

314 SIR SIDNEY F. HARMER ON

Challenger,” “The Atlantic,” i. pp. 144, 142, 193, fig. on p. 143.—Genotype
(the only species), Maresia cyathus, Wyv. Thoms., n. sp. Synonym of
Kinetoskias, Danielssen, 1868,

Nelha, Busk, 18522, p. 18.—Genosyntypes, Nellia oculata, Busk, nom. noy.(Salicornaria
dichotoma, Busk, 1852", p. 367, nec S. dichotoma, Schweigger, 1819, Tab. 8 ;
1820, p. 428) and WV. simplex, Busk, nom. noy. 1852”, p. 19 (S. marginata,
Busk, 18521, p. 367). Genolectotype, Nellia oculata; see Canu, 1900, Bull.
Soc. Géol. France (3) xxviii. p. 382. It seems moderately certain that
NV. oculata is a synonym of Cellaria tenella, Lamarck, 1816, p. 185; and the
correct name of the genotype would thus be Wellia tenella (Lamk.).

Notamia, Fleming, 1828, p. 541.—Proposed, to replace Loricaria (q. v.), pre-occupied,
for Cellularia loriculata, Pallas (1766, p. 64) (Sertularia loricata, L., 1758,
p. 815) and S, bursaria, L. (1758, p. 814), but immediately qualified by the
statement that NV. bursaria is the type of a new genus, Hpistomia (q. v.). The
genotype is thus Notamia loricata, and the genus becomes a synonym of
Eucratea, Lamx. 1812 (q. v.). The loss of Notamia, in Polyzoa, would in any
case have been inevitable, as the name was pre-occupied by Notamia, Rafinesque
(1819, J. de Physique, etc., Ixxxix. p. 153), introduced for a species described
as a ‘‘ Polype” and also as belonging to the Fam. Sipunculide, but with a
terminal anus. See also Gemellaria.

Notoplites, n. gen.—See p. 348.

Onchoporella, Busk, 1884, p. 103.—Genotype, Carbasea bombycina, Busk, 18522,
p. 52. Busk’s species was not identical with Plustra bombycina, Ell. and Sol.
(see Huthyris), and it thus appears necessary to propose a new name for
C. bombycina, Busk. I suggest, therefore, that the genotype should be known
as Onchoporella buski, n. sp. I consider that Busk indicated his own
C. bombycina as the genotype, by placing it alone under the generic heading,
although he stated in a footnote (p. 104) that Sceruparia diaphana, Busk, is a
second species of Onchoporella. He here made a curious mistake, as it appears,
on consulting the reference he gives (1860, Q. J. M.S. viii. p. 281, pl. 31.
fig. 1), that he was deceived by the fact that fig. 1 does not occur as the first
species on the Plate. It is clear, from the description and figures, that Busk
meant to refer to fig. 2, which comes first on the Plate, and that the second
species he intended to place in Onchoporella was Carbasea ligulata, described
by him on p. 282 of his 1860 paper.

Ornithopora, D’Orbigny, 1852, A. 8. N. (8) xvi. p. 312; 1852, P.F.T.C. p. 321.—
Genotype (the only species), Ornithopora avicularia (Sertularia avicularia
(pars), L., 1758, p. 809; Cellularia avicularia, Pallas, 1766, p. 68). This
species is also the genotype of Bugulina, Gray, 1848; and the genus is a
synonym of Bugula, Oken, 1815.

Ornithoporina, D’Orbigny, 1852, A. 8. N. (3) xvi. pp. 312, 313 (Ornithoorina, errorim,
on p. 312); 1852, Pp. F. T. C. p. 822.—In the A. 8. N. D’Orbigny mentions
only one species, O. avicularia, which should accordingly be regarded as the
genotype. His citation of Ellis (1756, French Ed., p. 119, pl. 38.
fig. 7) shows, however, that the species intended was probably Avicularia

CELLULARINE AND OTHER POLYZOA. 315

flabellata, Gray, 1848, p. 106 (see Avicularia); and the correct name of the
genotype is thus Ornithoporina flabellata (Thomps., MSS.) (Gray). In the
second work cited D’Orbigny includes two other species as well. Synonym of
Avicularia, Gray, 1848 (with the same genotype) and of Bugula, Oken, 1815.

Petalostegus, Levinsen, 1909, pp. 97, 114.—Genotype (the only species), Petalostegus
bicornis (Catenaria bicornis, Busk, 1884, p. 14).

Planicellaria, D’Orbigny, 1852, A.S. N. (8) xvi. pp. 333, 338; P.F.T.C. pp. 26,
36, 181.—Genosyntypes, P. oculata and P. fenestrata, two fossil species,
apparently belonging to different genera.

Poricellaria, D’Orbigny, 1852, A.S. N. (8) xvi. p. 338; 1854, P. F. T.C. p. 1106.—
Genotype, selected by the author, Poricellaria alata, D’Orb., n. sp. Canu,
1907, Ann. Paldéont. ii. p. 142 (sep. p. 38) regards Diplodidynua, Reuss,
1869, as a synonym of this genus; and he describes and figures a specimen
which he refers to D. alata (D’Orb.).

Retiflustra, Levinson, 1909, pp. 124, 125, 414.—Genosyntypes, 2. schdnaut, Lev., n.sp.,
Carbasea cribriformis, Busk, 1852*,p. 51 (Retepora cornea, Busk, 1852", p. 380)
and Mlustra reticulum, Hincks, 1882, A.M.N.H. (5) x. p. 163. Genotype,
now selected, Retiflustra cornea (Busk).

Rhabdozowm, Hincks, 1882, A. M. N. H. (5) x. p. 160.—Genotype (the only species),
Lhabdozoum wilsoni, Hincks, n. sp.

Saccohydra, Billard, 1914, 2° Exp. Antarct. Franc., Hydroides, p. 5—Genotype (the
only species), Saccohydra problematica, Billard, n. sp. Prof. Billard recently
wrote to me informing me that he had ascertained this genus, described as a
Hydroid, to be a synonym of Burentsia. There is no doubt, from his figures,
that this conclusion is correct, and that Saccohydra is to be regarded as oue of
the Entoprocta, and asa synonym of Barentsia, Hincks, 1880, A.M. N. H. (5)
vi. p. 285. I publish this note at Prof. Billard’s request.

Salicornaria, Schweigger, 1819, Tab. 8; 1820, p. 428.—This genus is usually
attributed to Cuvier, 1817 (Régne An. iv. p. 75), who introduced it, in the
form “ Salicorniaires,’ for Cellularia salicornia (Pallas, 1866, p. 61), and
three other species referable to Tubucellaria and Menipea. Schweigger seems
to have been the first to use the form Salicornaria, while he may be considered
to have selected the genotype by including only one species, S. dichotoma
(Cellularia salicornia, Pall.). If Cellaria (q. v.) is accepted, Salicornarra
becomes a synonym of that genus.

Salicornia, de Blainyille, 1830, p. 419 ; see also 1834, p. 455.—De Blainville gives
this as Salicornia, Cuy., although Cuvier did not use it in this form, and he
includes two species, Cellularia salicornia, Pall. (see Salicornaria) and Cellaria
salicornioides, Lamouroux (1816, p. 127, but not in Cuvier’s list). Salicornia
is merely another form of Salicornaria.

Salicorniaria, Schinz, 1825, Das Thierreich, iv. p. 155.—Another variant of the same
name, the species included being those of Cuvier’s original list (see Sal-
cornaria), with some additional synonyms. The genus is used in the same
form by Templeton (1836, Loudon’s Mag. Nat. Hist. ix. p. 469).

316 SIR SIDNEY F. HARMER ON

Salpingia, Coppin, 1848, A. M.N.H. (2) ii. p. 273; see also Gray, 1848, p. 149.—
Genotype (the only species), Salpingia hassallii, Copp., n. sp. Synonym of
Aetea, Lamouroux, 1812.

Sarsiflustra, Jullien and Calvet, 1903, Rés. Camp. Sci. Prince de Monaco, xxiii.
pp. 48, 126.—Genotype (the only species), Sarsiflustra abyssicola (Flustra
abyssicola (M. Sars, MSS.), G. O. Sars, 1872, Christiania Uniy. Progr. Ist
half-year, 1869, p. 19). ;

Scruparia, Oken, 1815, p. 90.--Based on 9 species, one of which (Cellularia repens,
presumably =Sertularia repens, Ellis and Solander, 1786, p. 52) is a Hydroid.
The others are Polyzoa, corresponding with four modern genera. The first
species mentioned, and the only one in which a trivial name is definitely
associated with Scruparia, is Scruparia chelata (Sertularia chelata, L., 1758,
p- 816). This was the only species included in Scuparia (errore), Gray,
1848, p. 132, and in Scruparia, Busk, 1852°, p. 28; in the latter case with a
new diagnosis. Scruparia chelata should thus be regarded as the genotype.
Hincks’ later proposal (1880, p. 21) to make his own S. clavata the genotype
is inadmissible, as this species is not included in Oken’s original list. Verrill,
1880 (Proc. U. 8. Nat. Mus. ii. 1879, p. 190) chose S. reptans [Scrupocellaria]
as the genotype, but although in Oken’s list, this species has no claims if those
of S. chelata are established.

Scrupocellaria, Van Beneden, 1845, Nouv. Mém. Acad. Roy. Brux. xviii. p. 26.—
Genotype (the only species), Scrupocellaria scruposa (Sertularia scruposa, L.,
1758, p. 815).

Scuparia, Gray, 1848, p. 132.-This seems to have been a misquotation of Scruparia
(q. v.). ;

Scupocellaria, Gray, 1848, p. 111.—A similar mistake for Scrupocellaria (q. v.).

Selbia, Gray, 1843, p. 292.—Genotype (the only species), Selbia zelandica, Gray,
n. sp. Busk (1852°, pp. 37, 38) cites Gray’s species as a synonym of
Caberea (Crisia) boryz, Audouin (1826, p. 242); and even if not identical
with Audouin’s species, Selbia, of which no generic diagnosis was given,
must be regarded as a synonym of Caberea (q. u.).

Semiflustra, D’Orbigny, 1852, A. 8. N. (8) xvi. p. 317; 1852, P. F. T.C. p. 326.—
Genotype, Semiflustra bombycina (Llustra bombycina, Ellis and Solander, 1786,
p- 14), which is the only species mentioned in the “Annales des Sciences
Naturelles.” In the second work cited two other species are also included.
Carbasea bombycina, Busk, 1852°, p. 52, is a distinct species (see Onchoporella).
Euthyris, Hincks, 1882, A. M. N. H. (5) x. p. 164, may perhaps be regarded
as a synonym of Semiflustra (see Huthyris).

Spiralaria, Busk, 1861, Q. J. M.S. (a.s.) i. p. 153.—Genotype, Spiralaria florea,
Busk, n. sp.

Spiralis, Levinsen, 1909, p. 408 (Explanation of Pl. 19. fig. 10a).—A mistake for
Spiralaria, as shown by p. 125.

CELLULARINE AND OTHER POLYZOA. 317

Stirparia, Goldstein, 1880, Q. J. Micr. Soc. Vict. Proc. i. p. 75.—Genotype (the only
species), Stirparia annulata (Bicellaria annulata, Maplestone, 1879, t. ct.,
p. 19). The generic name is pre-occupied, for Pennatulacea, by Leuckart,
1841, Zool. Bruchstiicke, ii. p. 122; and I propose that it be replaced by
Stirpariella (nom. noy.), with the genotype Stirpariella annulata (Mapl.).

Stirpariella, nom. nov.—See Stirparia.

Stolonella, Hincks, 1883, A. M. N. H. (5) xi. p. 197.—Genotype (the only species),
Stolonella clausa, Hincks, n. sp.

Synnota, Pieper, 1881.—See Synnotum.

Synnotum, Pieper, 1881, Neunte Jahresb. Westfil. Provinzial-Ver. pro 1880, p. 47.—
Pieper suggested the alternative names Mononota and Synnota for a single
new species, Giemellaria (?) avicularis. Synnotw was adopted, but amended to
Synnotum, presumably on etymological grounds, by Hincks, 1886 (A. M. N. H.
(5) xvii. p. 255); and I think this change in spelling may be accepted.
Waters (1897, J. L. S. xxvi. p. 15) pointed out that S. aviculare is probably
a synonym of Loricaria cegyptiaca, Audouin, 1826, p. 243; and I have
satisfied myself that this conclusion is correct. ‘The name of the genotype
should thus be Synnotum egyptiacum (Aud.). It does not seem to have
occurred to other authors that this species cannot be placed in the same genus
as Hucratea loricata (Linn.). See Hucratea and Gemellaria.

Ternicellaria, D’Orbigny, 1851, P. F. T. C. pp. 40, 47; see also 1852, A. 8. N. (3)
Xvi. p. 330.—Genotype (the only species), Zernicellaria aculeata (Bicellaria
(Lricellaria on pl. 2) aculeata, D’Orb., 1839-1846, Voy. Amér. Meérid.

vy. 4, p.8). Synonym of Zricellaria.

Tricellaria, Fleming, 1828, p. 540.—-Genotype (the only species), Dricellaria ternata
(Cellaria ternata, Ellis and Solander, 1786, p. 30).

Ulidium, Searles Wood, 1844, A. M. N. H. xiii. p. 17.—Genotype (the only species),
Ulidium charlesworthii (Melicerita charlesworthii, Milne Edwards). ‘The prin-
cipal reasons given for its introduction are that Jelicerita is etymologically
incorrect, Melicertina is objectionable, and names similar to Melicerita have
been used in other groups. Synonym of MJelicerita; see also Melicertina.

Unicellaria, de Blainville, 1830, p. 425; see also 1834, p. 461.—Genosyntypes,
Sertularia chelata, L. (as described by Ellis, 1756, Hist. Nat. Cor. French Kd.
p. 57), S. cornuta, L., 1758 (p. 810), Hucratea appendiculata, Lamouroux, 1821
(p. 8), and Lafoea lafoyi, de Blainville, nom. nov. for Lafoew cornuta,
Lamouroux, 1821 (p. 8). The first of these species is referable to Scruparia,
Oken, 1815; the second and third to Crista, Lamouroux, 1812; and the
fourth appears to be a Hydroid.

Watersia, Levinsen, 1909, pp. 94, 99.—Genotype (the only species), Watersia
militaris (Flustra militaris, Waters, 1887, A. M.N. H. (5) xx. p. 93).

318 SIR SIDNEY F. HARMER ON

III. Invernan Avicunaria.
Levinsen (1909, p. 139, pl. 2. figs. 7b, d, e, g, j, &) has described the

occasional replacement of an ordinary frontal avicularium, in Menipea
roborata, by an avicularium which grows into the body-cavity ; and in one
of his letters quoted in Sect. V. (p. 336) he refers to the presence of internal
avicularia in J/. marionensis. He makes no comment on this extraordinary
occurrence ; but however improbable it may seem, both on morphological
and on physiological grounds, there is no doubt of the accuracy of the
description. I have found these internal avicularia, not only in the two
species indicated by Levinsen, but also in certain other species of the same
genus.

The internal avicularia of AZ. marionensis are represented in three of my
figures. Pl. 19. fig. 44, a basal view, shows one of these structures (7. av.)
in each of the zocecia D and H. The avicularium is a more or less cylindrical
structure situated on the inner side of the frontal wall, and projecting basally
into the body-cavity. The palatal surface is on the basal side, there is a
normal beaked rostram, with which is connected an acute, triangular
mandible. The material is not in the best condition, and it has the
appearance of having been allowed to dry. The polypides are, however,
perfectly recognisable, and there seems to be clear evidence that the mandible
opens and closes in the body-cavity, and in actual contact with the tissues of
the polypide. The arrangement seems an inconvenient one, and I can suggest
no explanation of its purpose. There is no evidence whatever that the
avicularium lies in a cavity invaginated from the frontal side, and I cannot
escape from the conclusion that the mandible actually moves in the body-
cavity. Four of these internal avicularia are shown, in basal view, in Pl. 17.
fig. 22, in the zowecia C, D, H, and K. The base of the cylindrical avicularium,
i.av. (PI. 19. fig. 43), is not raised above the general level of the frontal wall,
and from it originate the occlusor muscles, which are seen in frontal view, of
the mandible. The avicularium has in fact an inverted position, and has been
developed on the inner side of the wall of the zocecium, instead of on its
outer side.

The account given by Levinsen of the internal avicularia of Menipea
roborata is not easy to understand ; and a renewed study of these remarkable
structures was required. There is no doubt that, as stated by Levinsen, the
internal avicularium replaces an ordinary frontal avicularium. I do not find
them always present, as he states, when an external avicularium is absent,
and I have not found them in the marginal rows. They may usually be
found without difficulty in some of the zocecia of the submarginal rows, and
oceasionally in other zocecia further removed from the margin. They seem
to be always wanting on the distal side of an ovicell, where two external
avicularia are constantly present, directed obliquely distally, and thus reversed

CELLULARINE AND OTHER POLYZOA. 319

in their direction as compared with the avicularia not related to an ovicell ;—
again in accordance with Levinsen’s description.

The internal avicularium is longer than would be inferred from Leyinsen’s
account. Its proximal end is in the same position as that of an external
avicularium. The side view (PI. 18. fig. 29) shows, in each of two zoeecia, a
pair of avicularia lying side by side. While one of them (f.av.) rises
normally on the frontal side of the zocecium, the other (i.av.) runs distally
and basally below the cryptoeyst, the avicularium being long and sub-
eylindrical, and terminating in a beaked rostrum, with an opesia subdivided
by the projection into it of two calcareous teeth (see Levinsen, pl. 2. fig. 7h).
In a frontal view (Pl. 18. fig. 30) the distal end of the internal avicularium
(t.av.) is seen through the opesia, and its proximal end (p.) forms a slightly
convex calcareous film, which is partly crossed by the triangular proximal
end of the cryptocyst (er.). This overgrowth by the cryptocyst does not
occur in the external avicularia, which grow frontally from their base.

Levinsen did not discover internal avicularia in M/. ligulataand WM. spicata,
two species which were associated by MacGillivray with J. roborata in his
genus Craspedozoum (see Sect. II.). I have found them in both these species,
as well as in MV. vectifera, n. sp. (see Sect. V.) and in WV. triseriata, Busk.
Their occurrence in undoubted species of Menipea is of special interest, as
indicating that Craspedozoum should probably be merged in that genus.

The internal avicularia do not seem to be common in MV. ligulata, but I
have found them, as shown in Pl. 18. fig. 33, in one or two zowcia. They
resemble those of M/. roborata, but they appear to be less asymmetrical in
position.

In M. spicata (figs. 32, 35) the internal avicularia are considerably smaller
than in the other two species. They lie closely attached to one of the lateral
walls of the zoccium, and do not reach its opesia (fig. 35). In side view
(fig. 32) they are seen to pass nearly vertically downwards into the body-
cavity. The internal avicularia of M/. vectifera (fig. 36) resemble those of
M. spicata.

In MW. trisertata (Pl. 19. figs. 41, 42) the internal avicularium is broad and
short, its distal end being just visible, through the opesia, in frontal view
(fig. 42). It will be seen from these figures, as well as from one or two of
those illustrating other species, that a considerable proportion of the internal
avicularia found occur in the more proximal of the two zocecia formed when
a longitudinal row is doubled.

The species in which I have found internal avicularia may all be placed in
Menipea, s. str., as understood in Sect. V.; and they constitute a definite
evidence of affinity in a group of species which can be placed together for
other reasons. It seems probable that these remarkable structures will be
found in other species of the same genus, though I have failed to find them
at present in any but the ones which have been mentioned above.

320 SIR SIDNEY F. HARMER ON

IV. Mernops or BiruRcATIoN OF THE COLONY.

The ‘ Cellularine” series of Cheilostomatous Polyzoa, which takes its
name from the genus Cellularia (see Sect. II.), consists of numerous genera
and species in which the zocecium has a membranous frontal wall and the
colony has an erect, branching habit. The character of the frontal wall
places them in Levinsen’s Sub-order Anasca (1909, p. 91). The Cellularine
habit of growth is closely parallelled, however, in members of the Sub-order
Ascophora, in which the frontal wall is caleareous and a compensation-sac is
present (cf. Harmer, 1902, ref. on p. 295). The genus Catenicella and its
allies may be mentioned in this connection ; but none of the Ascophorous
genera are here considered.

The Anascous branching forms are usually placed, in systematic treatises,
near the commencement of the Cheilostomatous series; and it has often been
at least tacitly assumed that they represent a low stage in the evolution of
the Cheilostomata. This view seems to me erroneous; and the highly
evolved nature of this assemblage is indicated by the characters of their
heterozocecia, a term introduced by Levinsen (Vid. Medd. Naturh. Foren.
Copenhagen, 1902, p. 3) to include the avicularia and vibracula. The avi-
ccularium reaches the summit of its development in such Cellularine genera
as Bugula and Cornucopina, while the vibraculum is highly specialized in
Caberea, belonging to the same assemblage. The assumption that the
branching habit is in any sense primitive appears to me fundamentally
wrong. It is no doubt true that the Cellularine species are less adapted for
preservation as fossils than the encrusting forms ; but, making every allowance
for this consideration, the Paleeontological evidence points to the encrusting
habit as the more primitive ; and already in the Cretaceous Period, large
numbers of encrusting Cheilostomes, of a primitive type in other respects,
are known.

Not only are there these reasons for viewing with suspicion the claims of
the Cellularine genera to be regarded as representing an early stage in
evolution, but the study of their mode of branching leads readily to the
conclusion that the erect colony is a lamina which has been more or less
subdivided. It may be noticed that the majority of Cellularine species
consist entirely of branches having two surfaces sharply differentiated. The
basal surface of the branch shows merely the “backs” of the individual
zocecia, all of which have their orifices on the opposite or frontal surface.
There is thus no difficulty in regarding the typical Cellularine colony as a
unilaminar sheet of zocecia, divided by more or less radial slits into narrow
branches, which in the majority of species are built up of two longitudinal
series of zooecia, alternating on the two sides of the branch. It might appear
logical to regard the biserial condition as representing the last stage in this
process, and thus to suppose that when biserial and multiserial branches
occur in closely related species, the multiserial condition should in all cases

CELLULARINE AND OTHER POLYZOA. 321

be regarded as the more primitive. Iam not sure that this is always the case,,
and I think it is at least possible that a multiserial branch may in some cases
have been derived secondarily from a preceding biserial condition. I do not
consider it necessary to discuss this question in the present paper, in which
I desire to point out, however, that the extent to which the splitting of the
original lamina has taken place differs in various genera and species; and
that the study of the actual facts of the bifurcation of the branches may be
of great assistance in systematic work. Sofar as I am aware, there has been
no previous comparative account of this subject, although the importance of
the mode of branching has not been overlooked by other writers. I may
reter especially to the account given by Davenport (1891, Bull. Mus. Comp.
Zool. Harvard, xxii. p. 41) of the branching of Bugula, and to two papers by
Waters (1897, J. Linn. Soc. xxvi. p. 2; 1913, Proc. Zool. Soc. p. 473),
dealing with the mode of bifurcation in Scrupocellaria and Menipea. These
papers record important facts, but they do not exhaust the subject.
Davenport is concerned with the general laws of branching, and he does
not attempt to show how his results can be applied to classification. In his
1913 paper Waters divides Menzpea into several groups, based on differences
in the mode of branching ; but I think he has not been altogether successful
in his conclusions. Waters (1913) defines a group “ O,” for instance, which
he supposes to include species having a common type of bifurcation ; but he
places in it Menipea patagonica and Bugulopsis peachii, two species which,
according to my own results, differ essentially in their mode of bifurcation.
In studying the bifurcation, particularly in the Family Scrupocellariide,
it is necessary to devote special attention to the formation of the chitinous
joints with which many species are provided. In the majority of cases, each
of the branches is jointed at its base, in such a way that the actual bifur-
cating point is immediately succeeded by a joint at the origin of each branch.
In some cases, only one of the branches is thus jointed ; an arrangement
which may give rise to a sympodial form of stem, unilateral when the joints
all occur on the same side, or bilateral when they are formed alternately on
the two sides. The joint is developed in a manner which is remarkably con-
stant in the most diverse members of the calcareous Polyzoa ; occurring in
fundamentally the same way in Cyclostomata (Crisia) and in a variety of
Cheilostomata,; which seem to have ne close connection with one another.
The process has been described by various authors, among whom I may
mention Waters (1881, Q. J. Geol. Soc. xxxvil. p. 320; 1887, Ann. Mag.
Nat. Hist. (5) xx. pp. 89, 92; 1897, J. Linn. Soc. xxvi. p. 2; 1913, Proc.
Zool. Soc. p. 472) ; Pergens (1890, Bull. Soc. Belge Géol. iii. p. 313) ; and
Lomas (1889, Proc. Liverpool Biol. Soc. iil. p. 219). The joint is formed
across one or more zocecia, the calcareous walls of which are at first complete
and continuous. A chitinous lining is developed on the inner side of the
caleareous wall and in close contact with it (ef. Pergens, text-fig. 8, on

322 SIR SIDNEY F. HARMER ON

p. 314), in the form of a cylindrical tube, rather longer than the future joint,
and open at both ends. An annular zone of the calcareous wall is then
absorbed, at the middle of the chitinous tube, which is left as the sole con-
nection between the distal part of the zocecium and the parent-internode.
Tt thus follows that the jointed zocecium partakes in the formation of two
internodes; the greater part of it usually lying in the proximal end of the
daughter-internode, while the part on the proximal side of the chitinous
joint is immersed in the parent-internode. The polypide commonly passes
through the jointed region of the zowcium, not only in the early stage, before
the absorption of the caleareous annulus, but through the chitinous tube even
after the completion of the joint, and in fact throughout the whole period of
its own life. The proximal segment of the jointed zocecium has often been
described as a “special chamber” of the parent-internode, by authors who
haye not fully appreciated its morphological significance.

In certain cases, as in species of Scrupocellaria and in Poricellaria (Diplo-
didymia), further strength is afforded to the joint by the development of
additional tubes of chitin, each formed on the inner side of its predecessor
and somewhat longer than it. A considerable number of these tubes may be
formed, and the whole chitinous complex thus acquires a considerable thick-
ness, showing at each end aseries of rings of diminishing diameter in passing
from the outer to the inner surface of the tube. ‘This arrangement, which
may be described as a system of tubes ‘Sen échelon,’ is indicated by
Claparéde (1870, Zeitschr. wiss. Zool. xxi. pl. 9. fig. 1 C), and his figure also
shows the circular, transverse furrows of the annular thickenings of the
calcareous wall with which the edges of the successive chitinous tubes are
firmly united. This arrangement, which is most noticeable in the more
robust species of Serupocellaria, appears to be a mechanical arrangement
admirably adapted to give the greatest strength at the middle of the joint,
and to allow the entire stem the flexibility which is required in order to obviate
fracture of the delicate branches.

The bifurcation of a biserial branch takes place by the doubling of the
number of zocecia, and generally in such a way that two successive, alter-
nating zocecia of opposite side of the parent-internode are each followed by
two distal successors, instead of by a single successor as in parts where the
internode is merely elongating without dividing. For the purpose of com-
parison I have adopted a uniform notation for the zocecia concerned in the
bifureation, as may be seen by reference to figs. 1-18. The more proximal
of the two zocecia which prepare the way for.the bifurcation is distinguished
as A, and the more distal zowcium, on the other side of the branch, as B.
C and D, on the outer sides of the arms of the Y-shaped bifurcation, may be
regarded as the direct successors of A and B respectively ; and it will be
seen that they agree closely with their predecessors in form, differing from
them only by diverging from one another to an extent sufficient to allow

CELLULARINE AND OTHER POLYZOA. 323

two other zocecia to be intercalated between them. These two zoccia are
denoted EH and F'; Bi being derived from A and lying on the inner side of
C; and F being derived from B and lying on the inner side of D. E is
usually in close relation with the axil of the bifurcation, and for this reason
I term it the axillary zocecium. It may be the proximal inner zocecium of
one of the branches and it is succeeded by G, the second inner zoccium.
F and H are the corresponding members of the other branch.

A careful study of various Cellularine genera, with particular reference to
the relations of the zocecia above enumerated (and in some cases of one or
two additional zocecia) has led me to the conclusion that the mode of bifur-
cation may be used as an important generic or specific character. I do not
overlook the danger of relying exclusively on a single character, but the use
I make of the bifurcation seems to be justified by finding that species thus
assorted appear, on the evidence of other characters as well, to form natural
groups. It appears to me probable that during the evolution of these genera
particular methods of bifurcation were adopted at an early stage, and can be
recognized, with modifications, in most if not in all of the species which
constitute the genera. It must be added that in most of the species here
considered the proximal end of the distal zocecium of a longitudinal row over-
laps its predecessor on its basal side. The diagrammatic figures are all repre-
sentations of basal views, and the distal ends of the zocecia are thus hidden by
the proximal ends of their successors. The zocecium A always lies to the
right of the figure ; and, in species in which A and B alternate, A typically
lies on that side of the parent-internode which is external in relation to the
preceding bifurcation. It thus follows that if the branches ECG and
F DH were followed further to the points where new bifurcations occurred,
the “A” zocecium of each of these would be found on the outer side. ‘The -
right band bifureation would be a repetition of the bifurcation actually
figured, and the left hand bifurcation would be its looking-glass image,
A lying on the left or outer side. Didymozoum forms an exception to this
rule, as the intercalation of a median ovicell-bearing zoccium takes place by
a division of the inner row of a biserial branch into two rows.

Before bifurcation occurs, the zocecia of opposite sides of the branch are
in lateral contact with one another, and are connected by rosette-plates or
communication-pores. In Bugula and some of its allies a peculiar modifi-
cation of this relation is found at the bifurcations (P1.16. figs. 3-5). Before
becoming completely disjoined, the two branches remain as a rule united with
one another by a special communication, formed in the axil by two of the
lower members of the branches. The connecting process (c. p.) is formed in
several different ways, which are remarkably constant within the limits of a
species, which may thus be distinguished from others with which it might be
confused. In only one or two species I have found, however, that the more
proximal bifurcations of a colony may belong to one type, and the more

324 SIR SIDNEY F. HARMER ON

distal ones to another. This is the case, for instance, in a colony of Bicel-
larina alderi, in which two successive types are represented ; but, so far as
my observations go, this is exceptional. It is well known, however, that the
proximal end of a colony may show characters regarded as juvenile, as com-
pared with the more differentiated distal parts; and even if it should be
found that differences in the mode of bifurcation occur at opposite ends
of the colony, in other species, it will not, I think, affect the correctness of
the statement that each species of Bugula has a practically constant method
of bifurcation, in its fully developed condition. It may be added, finally,
that the arrangement assumed is dependent, in the main, on the extent to
which the inner zocecia remain in lateral contact with one another, or, in
other words, on the distance to which the split forming the bifurcation
extends towards the zowcia A and B.

The position of the rootlets, with regard to the bifurcations, has previously
been recognized as a character of importance, particularly by Waters (1897,
1913, cited on p. 321). In the series including Menipea and its allies, these
structures are given off from a pore-chamber which projects into the body-
cavity, sometimes on the distal side of the jointed region (fig. 15), and some-
times on its proximal side (figs. 9-11). The difference may appear a slight
one, but the general flexibility of the colony must be affected by the position
of the rootlets. It seems probable that, when a joint has been evolved, the
relation of the rootlets to it would not be easily altered during subsequent
modifications of the species. Thus one series of species, represented by
Menipea and Notoplites, may be supposed to have started with their rootlets
on the distal sides of the joints, and at the proximal ends of the internodes ;
and to have retained these relations during their later evolution. Tricellaria
may be similarly supposed to have originated from a condition in which the
rootlets were on the proximal sides of the joints and at the distal ends of
the internodes.

Levinsen (1909, p. 132 n.) has stated that vestigial vibracula may be recog—
nized in the pore-chambers of the rootlets of Zricellaria ternata. I think
this view is correct, and in fig. 11 I have shown structures which admit of
this interpretation in 7’. peachii, on the proximal segments of the zocecia C
and D. In Serupocellaria each zocecium is typically provided with a vibra-
culum, which lies on the basal surtace of the branch, usually near its external
border. The vibraculum belongs to the proximal end of the zocecium,
although it is in close relation with. the external or marginal avicularium of
the preceding zoccium of the same longitudinal row. The vibraculum is
constantly provided, in this genus, with a chamber, cut off by calcareous walls.
from the cavity which contains its muscles, and this chamber gives origin to
a rootlet, in the more proximal members of the colony at least ; although in
many of the other vibracula the rootlet is merely represented by an oval
fenestra in the outer wall of the rootlet-chamber, corresponding exactly with

CELLULARINE AND OTHER POLYZOA. 325

the fenestra occurring in a vibraculum which actually develops a rootlet.
T cannot agree with Levinsen’s statement (1909, p. 134) that the rootlet-
chamber is not thus cut off in Serupocellaria. In Tricellaria peachii (fig. 11)
the structure from which the rootlet originates is a chamber cut off by a wall
from a more distally placed portion which shows a slight longitudinal groove,
apparently representing the rostral groove which receives the seta of a normal
vibraculum. I regard the structure in question as a vestigial vibraculum,
situated in its proper place at the proximal end of a zocecium and on its basal
surface. The rootlet-chambers thus situated are doubtless the rootlets pro-
duced by “a definite dorso-lateral chamber situated just above the lateral
avicularium,” mentioned by Miss Robertson (1905, Univ. California Publ.,
Zool. ii. p. 250) in “ Menipea” (Tricellaria). As pointed out by the same
author, the rootlets given off by these chambers may pass distally and form
the tendril-like structures known to occur in Tvicellaria. The tendrils are
not always formed in this way, however ; as I find that in Hmma tricellata,
Busk, one of these structures may replace a branch at a bifurcation. I believe
this to occur by the suppression of the proximal segment of F or G at a
bifurcation, as in fig. 13, and by the hypertrophy of the chitinous tube which
would otherwise have formed a joint, unaccompanied by the production of
the calcareous parts constituting a zocecium. In other cases, as in fig. 12,
a rootlet is developed from the outer side of a zocecium which is not con-
cerned in a bifurcation. The pore-chamber of the rootlet is still in the
proper place for a vibraculum, at the proximal end of a zocecium ; and
although greatly reduced may still represent that structure.

The rootlets which do not become tendrils pass proximally, sometimes
along the basal surface, but in many cases along the lateral margins of the
branches. The formation of lateral bundles of parallel rootlets is specially
characteristic of Amastigia, Notoplites, and Menipea.

Type 1 (Pl. 16. fig. 1).—The zocecia occur in pairs, and not alternately on
the two sides of the branch. A and B are thus at the same level, and the
split separating the two branches reaches their distal ends. H, C and F, D
thus form symmetrical pairs, H and F remaining in contact with one another,
at their proximal ends, on the basal.side of A and B.—Didymozoum. (The
arrangement is slightly modified in an internode which has become triserial
distally by the development of an ovicell.)

Type 2 (fig. 2)—Bifureation oecurs before a doubling of the number
of zocecia takes place, the proximal end of each branch being thus uniserial.
C remains in contact with the inner lobe of the proximal fork of D; and the
doubling of the number of zocecia takes place at the distal ends of C and D,
or of one of their successors if more than a single uniserial zocecium occurs
at the proximal end of the branch.—Stirpariella, sp. (‘ Siboga’ Collection).

LINN. JOURN.—ZOOLOGY, VOL. XXXY. 4

326 SIR SIDNEY F. HARMER ON

Type 3 (fig. 3).—The split extends nearly to the proximal end of the
axillary zocecium, H, which is accordingly free, on its inner side, for the
greater part of its length. E gives off a connecting process (¢.p.) to meet
the proximal end of F. In the species figured, each of the pairs H, C and
F, D has a single fork, at the proximal end, divided by the common septal
wall in such a way that each of the four zocecia has only a half fork, instead
of a complete fork as in the other zowcia.—Bugula johnstone (Gray), 1843,
and other species of this genus.

Type 4 (fig. 4).—The split does not extend proximally so far as in type 3,
not more than half the axillary zocecium (E) being free on its inner side.
The connecting process (c.p.) is given off by F, and joins the inner lobe of
the fork of G, which crosses the basal side of E.—Species of Bugula, e. g.,
B. scaphoides, Kirkp., 1890, A. M. N.H. (6) v. p. 18, and Bicellariella, and
most of the species of Stirpariella, e.g., S. zanzibariensis, Waters, 1913,
P.Z.S. p. 469.

Type 5 (fig. 5).—The axillary zocecium is almost completely immersed in
the parent-internode, the split only reaching its distal end. The connecting
process (c.p.) is given off by G and joins the inner lobe of the fork of H.—
Species of Bugula, e. g., B. dentata (Lamx.), 1816, p. 135, and of Stirparielia,
e.g., S. caraibica, Levinsen, 1909, p. 104.

Type 6 (fig. 6).—Although F is in contact with B, it appears to be derived
from H, which it meets near the middle of the length of the latter. The
connection between the proximal ends of the inner zocecia of the branches is
formed by the independent junction of F and G with E. The proximal ends
of the zoccia are not bifureate. Oblique joints, indicated by dotted lines
and of a type common in Bugula, traverse the zocecia.— Huoplozoum cirratum
(Busk), 1884, p. 17.

Type 7 (fig. 7).—Resembles type 6 in the relation of G and F to H, but is
peculiar in other respects. The axillary zocecium (HE), which gives rise to
and G, passes into the proximal end of the branch on the side opposite to A,
a relation which has not been found in any other type. B takes no part in
the doubling of the number of zoccia, the proximal ends of which are not
bifureate.—Kinetoskias cyathus and all the other species (4) examined.

In the following types (8-18) the position of the chitinous joints is of
importance.

Type 8 (fig. 8).—The proximal segments of F and G are in contact on the
basal side of EH, and are shorter than the corresponding parts of C and D;
the joints traversing F and G near their proximal ends, and passing much |
more distally through C and D. In the species figured (Scrupocellaria ferow,
Busk, 1852), the joint traverses the middle of the opesia, which is represented
by dotted lines, of each of the outer zocecia C and D. In certain other

CELLULARINE AND OTHER POLYZOA. 327

species of the genus each joint passes across the proximal end of the opesia
of the outer zocecium ; and in others, particularly in those with elongated
zoccia, it passes entirely on the proximal side of the opesia of C and D.—
Serupocellaria, all species. Himantozowum may be considered to belong to
this type, with which it agrees exeept that it is unjointed.

Type 9 (fig. 9).—The proximal segments of F and G are not in contact,
and are shorter than the corresponding parts of C and D; the joints
traversing CG and DF respectively. A rootlet-chamber occurs on the
proximal segment of C or D, or of both, and the rootlet is thus given off on
the proximal side of the joint.— 77icellaria, spp.

Type 10 (fig. 10).—The proximal segments of F and G are separated by
the whole width of E, and the two branches come off at different levels, that
on the side of A being the more proximally situated. Internodes commonly
composed of three zocecia, C and D then taking on the character of A and B
zocwcia in the next internode.—Tricellaria, spp. (Figured specimen, 7. occi-
dentalis var. dilatata, Ortmann, 1889, p. 21.)

Type 11 (fig. 11).—Hxcept in the fact that the proximal segments of F
and G are in contact, this agrees with type 9.—Tricellaria peachii (Busk).

Type 12 (fig. 12).—Readily derivable from type 10 by the suppression of
the proximal segments of F and G, the internode thus consisting of a single
zocecium at its proximal end, connected with its predecessor by a single
chitinous joint.— T’ricellaria aculeata, D’Orb.

Type 13 (fig. 13).—A modification agreeing with type 12, occurring in
Emma cyathus. The zocecia C and D take on the character of A and B
zocecia at their distal ends.

Type 14 (fig. 14).—Internodes composed of three zocecia and, like the
zooecia, very short. A slight modification of type 9.—mma, spp.

Type 15 (fig. 15).—Proximal segments of F and G in contact, much
longer than the corresponding parts of J and K. The joints are on the distal
‘sides of C and D, and traverse the zoecia FJ and GK. The joints are
‘situated at some distance from the axil, instead of being on the same level
with it, and do not traverse the opesia of any zocecium., Rootlets given off
on the distal sides of the joints.— Notoplites. A modification represented, in
the same genus, in the * Siboga’ Collection results from the suppression of one
of the joints of the bifurcation.

Type 16 (Pl. 1%. fig. 16).—The axillary zocecium is free on both sides
distally. The zocecia F and G are not in contact with one another, but each
is connected with one side of H at a point marked by the presence of a con-
spicuous rosette-plate (7.p.).—Cornucopina, all species examined. The
zocecia are very long, and a constriction partially separates a small proximal

section from the main part of the zocecium.
24*

328 SIR SIDNEY F. HARMER ON

Type 17 (fig. 17).—Proximal segments of F and G in contact (rarely
separated), longer than the corresponding parts of J and K. The joints are
on the distal sides of C and D, lying hardly in advance of the axil, and they
traverse the zocecia FJ and GK, passing through the opesia of F and G, the
two inner zowcia. Rootlets given off on the distal sides of the joints.—
Menipea. :

Type 18 (fig. 18).—The internode becomes triserial by the development of
a median row of two or more zocecia. The distal member of the median row
(Hi? in the figured specimen) behaves as an axillary zocecium. Bifureation
in other respects as in type 17.—Menipea (triserial species).

Type 19 (fig. 19).—Part of a sympodial colony of Menipea spicata, only
one of the branches being jointed at each bifurcation.

Type 20 (fig. 20).—Agreeing with type 8 except for the intercalation of a
median series of zocecia in the internode.—Amastigia kirkpatricki (Lev.,

MSS.), n. sp.

V. CHARACTERS OF CERTAIN GENERA AND SPECIES OF
ScRUPOCELLARIID&.

Family SCRUPOCELLARIIDA.
Cellulariidee, Hincks, 1880, p. cxxxvii; Busk, 1884, p. xxii; MacGillivray, 1887,
Trans. Proc. R. Soc. Vict. xxiii. p. 199; et auett.
Serupocellariide, Levinsen, 1909, pp. 89, 180; Canu & Bassler, 1920, U.S. Nat.
Mus., Bull. 106, p. 180.

A diagnosis, based on Levinsen’s study of the Family, has been given by
Canu and Bassler, and this may be accepted with but few modifications.
The Family consists of Anascous Cheilostomata, of erect habit, usually much
branched, and attached by rootlets. With rare exceptions the zoarium is
unilaminar, and the branches are biserial in the majority of species. The
branches are nearly always flexible, well-developed chitinous joints occurring
in most cases. The opesia is large, and spines usually occur distally or at the
sides of the orifice. A specially modified spine, the scutum or fornix, jointed
at its base, projects horizontally over the opesia, being branched or broadened
except in the region of its narrow stalk ; but it may be completely wanting.
The full equipment of heterozocecia consists of (1) an avicularium, or a pair
of avicularia, on the frontal surface, typically on the proximal side of the
opesia, and with the rostra directed proximally. The frontal avicularia on
the distal side of an ovicell are commonly paired and directed distally ;
(2) a lateral or marginal avicularium at the outer distal corner of the
zocecium ; (3) a basal heterozocecium, which may be either an avicularium
or a vibraculum, on the basal surface, at the proximal end of the zocecium,
usually in contact with the lateral avicularium of the preceding zocecium.
The heterozocecia are not jointed at their base, and any or all of them may

CELLULARINE AND OTHER POLYZOA. 329

be wanting. The mandibles of the avicularia are invariably of the poimted
type. The terminal wall of the zocecium is at first vertical, its origin from
the basal wall straight or slightly curved, but then bends distally, in
approaching the frontal surface ; the proximal portion of each zocecium thus
overlapping its predecessor on its basal side. The ovicells are hyperstomial,
frequently completely contained in a depression on the frontal wall of the
zocecium which succeeds the fertile zocecium, and not projecting beyond the
general frontal level of the zocecia.

The scutum and the heterozocecia appear to be very capricious in their
occurrence. In several of the genera one or more of these structures
are present, on most of the zocecia, in certain species ; while they are absent
in other species which appear to be closely allied to those of the first series.
I think the only interpretation which can be given of these facts is that all
these structures are to be regarded as original possessions of the Family,
and that their absence is to be attributed to loss.

Dr. W. D. Lang has come to a similar conclusion with regard to Creta-
ceous Cribrimorphs (1922, Cat. Foss. Bry. (Pol.) Brit. Mus. iv. pp. 3, 196,
and elsewhere).

The extreme amount of loss is found in 7ricellaria peachii, in which scutum
and heterozocecia are completely wanting, if exception be made of the
possibly vestigial vibraculum which has been described on p. 324. On the
assumption that the heterozoccia are structures which give their possessors
some advantage in the struggle for existence, it is very difficult to understand
the frequency with which they disappear entirely ; the species in which this
takes place continuing to exist, apparently with undiminished efficiency.

Genera here considered :-—
1. Amastigia, Busk.
2. Menipea, Lamouroux.
3. Wotoplites, n. gen.
4. Tricellaria, Fleming.
5. Emma, Gray.
6. Serupocellaria, Van Beneden.
[The genera Canda and Caberea are omitted. |

1. AmasticrA, Bush.
Amastigia, Busk, 1852, p. 40.
Caberiella, Levinsen, 1909, pp. 134, 135,
Anderssonia, Kluge, 1914, p. 617.
Scrupocellaria (pars) and Menipea (pars), auctt.

Zoarium usually unjointed, the branches in nearly all cases pluriserial.
Frontal surface typically convex, so that the marginal zoccia face outwardly,
the basal surface more or less flat, the zocecia of the median rows as a rule
partially or completely excluded from it. Spines, scutum, frontal and
marginal avicularia present or wanting. Basal heterozocecia typically

330 SIR SIDNEY F. HARMER ON

present, in the form of avicularia, occasionally vibraculoid, or of vibracula.
Rootlets given off by the basal heterozowcia or from the sides of the marginal
zocecia, passing proximally as marginal bundles down the colony and
frequently extending round the axils of the bifurcations. Ovicells typically
small, often with a frontal fenestra.

Distribution, cireumpolar (Southern), reaching as far north as Victoria
in shallow water and Valparaiso in deep water.

The characteristic features of this genus appear to be the pluriserial
branches, which are usually more or less semi-cylindrical, with the zocecia
opening on the curved surface ; the tendency for the basal walls of some of
the inner zocecia to be reduced ; the occurrence of basal heterozocecia (some-
times wanting) ; and the arrangement of the rootlets in marginal bundles.
The last character also occurs in Menipea and Notoplites. The genus is of
special interest as demonstrating, better than any other, the intimate relation
between avicularia and vibracula. Joints definitely associated with a bifur-
cation seldom occur, but there is reason to believe that even in “unjointed ”
species they may be formed secondarily, as fractures occurring at some point
of an internode, the fracture being mended by the formation of chitinous
connecting tubes formed externally to the zocecia and not asa modification of
their own body-walls. It is not unlikely that I have included too many
species in this genus, which may have to be further subdivided.

1. Amasticia NuDA, Busk. (PI. 17. figs. 21, 24, 25; Pl. 19. figs. 50, 51.)
Amastiyia nuda, Busk, 1852°, p. 40, pl. 36. figs. 1-5. Tierra del Fuego.
by » (pars), MacGillivray, 1887, Trans. Proc. R. Soe. Vict. xxiii. p. 200.
Victoria.

The interest of this remarkable species seems to have completely escaped
recognition since the publication of Busk’s original account. MacGillivray
records it, on the authority of J. Bracebridge Wilson, from Victoria, and the
record is confirmed by slides in the British Museum. Kluge (1914, p. 613)
mentions the genus only to dismiss it.

The type-slide (Brit. Mus. 54.11.15.87), from material collected by Charles
Darwin, shows some interesting peculiarities which were not noticed by Busk.
The branches are at first biserial, and the zocecia of the two rows here meet
one another, in a normal manner, in the middle line of the basal surface.
They later become 3-serial, and then 5-serial, by the intercalation of three
additional rows. A basal view of a 5-serial region shows the appearance
indicated in Pl. 1%. fig. 24. The greater part of the basal surface is formed
by the marginal zocecia (1, 5), while the median (3) and submedian (2, 4)
zocecia have only a restricted origin from this wall. The frontal surface is
very convex, the middle region being much thicker than the sides, and the
marginal zocecia face obliquely outwards. As shown in Busk’s figs. 2, 3,
there is a large, undivided scutum, rounded distally. A frontal avicularium
may occur on the stbmedian zocecia, but there is usually a pair on those of

CELLULARINE AND OTHER POLYZOA. 331

the median row, the rostrum directed obliquely proximally except on the
distal side of an ovicell, where the direction is reversed, as in allied species.
A characteristic feature of Busk’s specimen is the occurrence of well-
developed lateral avicularia on the marginal zocecia. The most interesting
point is the presence of basal avicularia, which give off rootlets running on
the inner side of the lateral avicularia, as shown in Busk’s figs. 4, 5. These
have an acute rostrum, directed proximally, and they are rather longer than
wide, but not vibraculoid. The spines are well developed, though thin ;
four occurring on the outer side of the marginal zocecia and two, in addition to
the scutum, on the inner side. Scutum (Pl. 19. fig. 50) with a cavity which
is very narrow at its commencement but dilates towards the free edge. The
other zowecia haye three spines on each side, one of them being a scutum on
one side. In several of the marginal zocecia the distal external spine, which
is jointed at its base, is enormously elongated, being much longer than a
zooecium, and running straight distally, just on the outer side of the margin
of the branch, and parallel with it. he thick walls of these enlarged spines
appear to be entirely chitinous. The ovicells have a frontal fenestra.

In addition to the type-slide the British Museum possesses the following
specimens which I refer to A. nuda: —

97.5.1.246. Port Phillip Heads, Victoria, J. Bracebridge Wilson.

97.5.1.266 and 267. Same locality and donor (labelled Menipea funiculata).

97.5.1.250. Port Phillip, Victoria, Miss E. C. Jelly (labelled A. nuda).

99.7.1.823. Australia, Busk Collection (labelled A. nuda).

87.12.9.68. Kerguelen, ‘Challenger’ Collection, Stat. 149 I., 45-127 fathoms.
Recorded by Busk (1884, p. 19) as Menipea benemunita.

97.5.1.266 and 267 agree with the type, and differ from the other Victorian
specimens, by possessing conspicuous marginal avicularia. Some of their
spines are considerably enlarged, notably the second and third marginal ones,
and the spine on the distal side of the scutum. I have hesitated whether to
consider them distinct from the other Victorian specimens, but I think it is
safer to regard them as one species. Basal avicularia are plentiful in all,
sometimes completely alternate on the two sides of the branch and sometimes,
in the same colony, meeting one another in pairs (PI. 17. fig. 25). The
branches are at most 5-serial, and the number of spines and the frontal
avicularia (fig. 21) agree with the corresponding parts of the type. It is
noteworthy that ovicells occur in all five rows.

The ‘Challenger’ specimen from Kerguelen is of more slender habit than
typical A. benemunita, and is 5-serial. Its scutum is smaller than in that
species, and its spines’ are 4, 2+scutum, on the marginal zovecia, and 3, 3
(one being a scutum) on the others. In all these respects, as in the characters.
of the marginal and frontal avicularia, it agrees with A. nuda, to which
I refer it. If my determinations are correct, this species has a cireumpolar
distribution, from Tierra del Fuego to Kerguelen and Victoria.

332 SIR SIDNEY fF. HARMER ON

On isolating the rows of zocecia, after boiling in Eau de Javelle, the
method by which the zocecia of the median rows become partially separated
from the basai surface becomes apparent. As shown by Pl. 19. fig. 5],
the basal wall is very sinuous, the vertical diameter of the body-cavity
being greatest on the distal side of the length of the zocecium, and smallest
at the proximal end. The prominent part of the wall reaches the basal
surface of the braneh, while the indented proximal part is excluded from it,
the lateral walls of the adjacent zocecia sending lobes across this part, nearly
or quite meeting one another. The lozenges formed by the median zocecia
on the basal wall are thus produced; and the arrangement will be more
easily understood by referring to Pl. 17. fig. 27 (A. rudis), in which the
complete lateral outline of the distal median zocecium is represented. It will
be seen that the greatest prominence of the basal wall corresponds with the
widest part of the zocecium, while the proximal region of the zocecium is
narrower, and is either completely excluded from the basal wall or only meets
it in a linear median region. I believe this to be the method universally
adopted in this group of species, when the zocecia meet the basal wall
partially.

2. AMASTIGIA RUDIS (Busk). (PI. 1%. figs. 26, 27; Pl. 19. figs. 49, 52.)
Caberea rudis, Busk, 1852! p. 877 ; 1852 2, p. 38, pl. 46. figs. 1-3.
5 » MacGillivray, 1887, Prodr. Vict., Dec. xiv. p. 137, pl. 136. figs. b.
Menipea marginata, Hincks, 1884, A.M.N.H. (5) xiv. p. 276, pl. 9. figs. 1-1 d;
1893, Ibid. (6) xii. p. 143.

Branches 2-8-serial, coarser than in the preceding species, the zocecia
larger in all their measurements. Frontal surface strongly convex, the
marginal zoccia facing outwards. Basal surface flat or concave, the
marginal zocecia often projecting strongly asa broad, convex, longitudinal
band on each side, the region of the median rows depressed and concave.
Marginal zocecium forming a disproportionate part of the basal surface, the
others reaching the wall in regularly alternating lozenge-like areas, the
proximal end of each of these zocecia being overlapped by the broad parts of
thosé of contiguous rows, and excluded from the basal wall or meeting it in
a linear region. Opesia longer than in A. nuda, the cryptocyst not quite so
much developed. Spines more uniform in size than in A. nuda, those of the
marginal zocecia 4, 1, the fourth external more nearly vertical than the
others. Spines in the other zocecia 2, 2; the second of one side in the form
of a scutum, which does not fill the opesia, its distal lobe pointed (PI. 19.
fig. 49), its cavity as in A. nuda. Frontal avicularia paired on some of
the median rows, meeting in a median suture, their rostra directed as in
the preceding species; those in relation with an ovicell distant from one
another. Outer frontal avicularium suppressed on the marginal and often
on the submarginal rows, where the inner avicularium is often enlarged.
On the marginal zocecia the single frontal avicularium is commonly

CELLULARINE AND OTHER POLYZOA. 333

gigantic (Pl. 1%. fig. 26), with a strongly hooked rostrum, its articular
condyles strong and spike-like, and the mandible triangular and pointed
(as in the other avicularia), and with an incurved acuminate tip. Marginal
avicularia wanting. Basal heterozocecia in the form of curved vibracula
(fig. 27), placed in alternating pairs, the tip of the rostrum of each vibra-
-culum meeting the middle of its predecessor on the opposite side of the
branch. Rootlets as in other species. Ovicells resembling those of A. nuda.

Described from specimens in the British Museum, including Busk’s type
(54.11.15.82).

The basal heterozocecia of A. rudis (Pl. 1%. fig. 27) are of the kind
described by Levinsen (1909, p. 135) as “curved or angularly bent
vibracula ” in A. benemunita, as noticed under that species. LLevinsen founded
the genus Caberiella on this character, not recognizing the fact that Menipea
benemunita, Busk, is referable to Amastigia. Vibracula of this type occur
in profusion in A. rudts, varying in size but not in form. The rostral groove,
in which the long seta is received, is very long. ‘The occurrence of different
types of basal heterozocecia in Amastigia nuda and A. rudis indicates that
these structures are in a plastic condition in the genus, assuming the form
indifferently of avicularia and vibracula, or being completely absent.

In side view (Pl. 19. fig. 52) the zoweia are very different from those
of A. nuda, being larger in all their dimensions. The terminal wall is
more vertical, and the proximal end of the zocwcium much deeper, the
median zocecia being more completely represented on the basal wall than
in that species, a fact with which the smaller amount of sinuosity of the
basal wall (Pl. 19. fig. 52) is associated. The lateral communication-plates,
-as in other species of Amastigia (as well as in Menipea), are two in number.

The basal view (Pl. 17. fig. 27) of a branch of A. rudis is very charac-
teristic. The marginal zocecia occupy a disproportionate amount of this
wall, but the others, though here reduced, are not nearly as much so as in
A. nuda. Hach zocecium is wide distally, where it meets the basal wall
along its whole width. It becomes narrower proximally, as shown in the
distal zocecium of the median row; but the greater part of this region
is excluded from the basal surface by the union or close approximation
-of the widened parts of the preceding zocecia of the rows on either side
of it. The basal walls of all the zocecia except those of the marginal
rows thus appear as a series of regularly alternating spindle-shaped figures.
‘The basal vibracula, which are not drawn on the younger zoccia, are another
very characteristic feature of the species.

A. rudis differs from A. nuda in the larger number of series of zocecia in
its branches, in the smaller number of spines, in the distal lobe of the
scutum which is usually pointed, in the character of the basal heterozoccia,
in the gigantic frontal avicularia of the marginal rows, and in the larger
measurements of all of its parts.

334 SIR SIDNEY F. HARMER ON

3. AMASTIGIA CRASSIMARGINATA (Busk).

Cabereu crassimarginata, Busk, 1884, p. 28, pl. 11. figs. 1-16. §.E. of Buenos Aires,
600 fathoms,

_ Rootlets running down the margins of the branches. Vibracula covering:
only a small part of the basal surface, and resembling those of A. rudis.
Although this species is biserial the characters of its rootlets, vibracula and
scutum are in agreement with those of other species which I refer to
Amastigia. Waters (1913, Proc. Zool. Soc. p. 480) has suggested that it
should be placed in Canda.

4, AMASTIGIA BENEMUNITA (Bush).

Menipea benemunita, Busk, 1884, p. 19, pl. 4. figs. 4,4 @ (specimens from Stations.
303 (probably 308), 318, 314, 315 only). Straits of Magellan, W. of S. end of
S. America, Falkland Is., 5-175 tathoms,

Scrupocellaria benemunita, Jullien, 1888, Miss. Sci. Cap Horn, vi. p. 69, pl. 8.
figs. 1-8. 8S. of Cape Horn, 99 metres.

Caberiella benemunita, Levinsen, 1909, p. 185, pl. 22. figs. 8a, 8b. ‘ Challenger”
Exp., Stat. 313.

British Museum, ‘ Challenger’ Collection, 87.12.9.69-73. The type-slide is.
87.12.9.70, Stat. 313, E. end of the Straits of Magellan, 55 fathoms. Sir
John Murray thought that a mistake was made in one of the records, and
that Stat. 303 (slide 87.12.9.69) should have been Stat. 308, the depth of
which was 175 fathoms, and the locality near the W. coast of the 8. end
of S. America.

This species has a close resemblance to A. nuda, and it is curious that
Busk did not refer it to his own genus Amastigia. It can be distinguished
from A. nuda by its much larger scutum, with a large, upturned, truncate,
distal lobe, and by its flatter branches, which may be at least 7-serial, in
which all the zocecia appear to meet the basal wall normally, although the:
width of the median zocecia, basally, is less than that of the marginal zocecia.
The spines of the marginal zocecia are 3, 1+scutum. Marginal avicularia
are minute and commonly wanting. Under A. rudis I have alluded to.
Levinsen’s account of the basal heterozocecia, which he found in a ‘Challenger’
specimen from Stat. 313. I have not found heterozoccia of this type in the:
British Museum slide (87.12.9.70) from this Station, and the specimen seems.
to have only a single basal heterozocecium closely resembling those of
A. nuda, In 87.12.9.72, from Stat. 315, basal vibracula as described by
Levinsen occur ; and they resemble those figured by me (PI. 1%. fig. 27) in
A. rudis. The occurrence of two different types of basal heterozocecia in
different specimens of what I think must certainly be referred to the same
species is an interesting fact.

The ‘ Challenger’ slide from Stat. 149, Kerguelen, referred to this species.
by Busk, appears to belong to A. nuda.

CELLULARINE AND OTHER POLYZOA. 335

5. AMASTIGIA FUNICULATA (MacGillivray) .

Menipea funiculata, MacGillivray, 1886, Trans. Proc. R. Soc. Vict. xxii. p. 128, pl. 1.
figs. 8, 8@; 1889, Prodr. Zool. Vict., Dec. xviii. p. 285, pl. 177. figs. 6-6 6..
Victoria.

The British Museum possesses several Victorian specimens (87.12.10.38 ;
8$.11.14.221, 241 ; 97.5.1, 247, 269) of this species. It resembles A. nuda
in general appearance, but the frontal surfaces of the peripheral branches are
flat, and all the zocecia meet the basal wall normally. Some of the spines
are long and thick, particularly the second of the outer group of three in the
marginal zowcia. The single inner spine and the third or proximal outer
spine are short and thick, often crossing one another over the orifice of the
zocecium. ‘The scutum is oval, with a simple cavity, and it fills most of the
opesia. The branches are in the main triserial, sometimes 4-serial, and
marginal avicularia occur, though not commonly ; both these and the frontal
avicularia being small. I have not found basal heterozowcia. The branches
do not form joints at the bifurcations, but although typically unjointed there
seem to be indications of the occasional occurrence of secondarily formed
Joints. The more proximally placed branches may be narrow, with a very
convex frontal surface.

6. AMASTIGIA ABYSSICOLA (Kluge).
Scrupocellaria funiculata, Waters, 1904, ‘Belgica’ Bryozoa, p. 23, pl. 8. figs. 1a, 6-
Antarctic, 2800 metres.
Serupocellaria abyssicola, Kluge, 1914, p. 611. Antarctic, 2450 metres.

Kluge doubts Waters’ determination, and has suggested a new name for
the deep water Antarctic form observed by Waters and himself. Basal
heterozocecia not described.

7. AMASTIGIA CABERIOIDES (Kluge).

Serupocellaria caberioides, Kluge, 1914, p. 612, pl. xxvii. figs. 9,10. Antarctic, 350-
385 metres.

Unjointed, 2—4-serial. Scutum originating distally, not filling the opesia,
oval or long and narrow, the stalk inserted into the distal end of its lamina.
Marginal and frontal avicularia, the latter sometimes gigantic. Basal
avicularia paired, directed obliquely proximally, the tip of the rostrum
slightly vibraculoid. Median zocecia much reduced basally. Kluge rejects
his own suggestion that this species might be placed in Amastegia.

§. AMASTIGIA KIRKPATRICKI (Levinsen, MSS.), n. sp. (PI. 17. fig. 20;
Pl. 19. figs. 46-48.)

Menipea marionensis (pars), Busk, 1884, p. 21, pl. 14. fig. 9 (specimens from Stat.
144 a, Marion Island, 50-75 fathoms, only).

Branches biserial, becoming triserial at a varying distance from the
proximal end. Internodes usually of 3-6 pairs of zocecia, with at least one

336 SIR SIDNEY F. HARMER ON

median zocecium preceding the bifurcation, often with a row of 2-4 median
zoocia. Rootlets arising from the proximal members, mainly attached to
the edges of the branches, some of them passing down the basal surface.
Opesia oval, rather more than half the frontal length, with a moderate
cryptocyst, extending completely round the opesia, and of practically equal
width all round. Spines one external (rarely two) and one internal, at the
distal end, the external spine placed rather more distally than the internal ;
occasionally more numerous (fig. 46), the figure probably representing a
zocecium in an early state of growth of acolony. Scutum wanting. Frontal
avicularia single on most of the zocecia, those of some of the median zocecia
enlarged or even gigantic (fig. 48); much raised, directed obliquely
proximally, and situated on the inner side of the zocecium, on the proximal
side of the opesia, which they do not reach. Two frontal avicularia may
be found on the axillary zocecium. Marginal avicularia small, often
absent. Vibracula large, on the inner side of the lateral zocecia, placed
nearly longitudinally, the rostrum or groove for the seta greatly prolonged
beyond the main part of the vibraculum, and reaching the middle line of the
branch ; a single vibraculum in the axil of the bifurcation. Ovicells large,
longer than wide, the frontal surface rather flat, becoming concave just
distally to the rim of the oritice of the ovicell. The surface is imperforate,
and the ectocecium covers only a narrow part of the frontal surface.
Bifurcation (fig. 20) as described below ; both branches jointed at their base,
the joint involving the proximal end of the opesia of the outer zocecium.

Type-slide (Brit. Mus.), 87.12.9.97, ‘ Challenger’ Coll., Stat. 144 a.

In his ‘Challenger’ Report, Busk described as Jentpea marionensis
specimens from the two following localities :—

Stat. 142, off Cape of Good Hope, 150 fathoms; Stat. 144 a, off Marion
Island, 50-75 fathoms.

On Oct. 4, 1910, the late Mr. G. M. R. Levinsen wrote to Mr. R. Kirk-
patrick, at the British Museum, stating that a fragment of a ‘ Challenger’
specimen of “ Menipea marionensis,” from Stat. 144a, had come into his
hands, through the Dundee Museum, and that it proved to be a Caberea.
On Jan. 26, 1911, he wrote further, to the effect that he had described and
figured the species from Stat. 144a, as Caberea hirkpatricki ; and, having
examined material, from Stat. 142, which had been sent to him by
Mr. Kirkpatrick, he had found that “the great plurality of the frontal
avicularia are internal” in M. marionensis. In returning the material which
had been lent to him, he added a label to the specimens from Stat. 144 a, as
follows :—“ Busk had wrongly named the specimen J. marionensis, the type
of which is from Stat. 142, Cape of Good Hope.”

Levinsen died in 1914, and, although I have made every effort to satisfy
myself on the subject, I have failed to find evidence that his results were
published. It must be assumed that the description referred to in his letter

CELLULARINE AND OTHER POLYZOA. 337

of Jan. 26, 1911, remained in MS. at the time of his death. Having
examined the whole of the material from the two Stations, I can confirm all
Levinsen’s statements of fact quoted above, although I do not agree with
him in placing the new species in Caberea. There is no doubt that Busk
confused two species, and it is unfortunate that marionensis must be applied,
as shown by his description, to a species from the Cape of Good Hope, not at
present known to occur off Marion Island.

I have adopted Levinsen’s MS. name kirkpatricki for the new species ; and
I give below a new diagnosis of Menipea marionensis. The two species can
readily be distinguished under a low magnification. In Menipea mario-
nensis the joints are placed at some distance from the axil of the bifurcation,
the ovicells are small and short, there are no vibracula, and none of the frontal
avicularia are enlarged. In Amastigia kirkpatricki the joints are at the level
of the axil, the ovicells are large and long, conspicuous basal vibracula are:
present, and the frontal avicularium of the axillary zocecium (or of the
proximal median zocecium if more than one is present) is usually much
larger than the others. This last feature is indicated in Busk’s fig. 9
(PI. 14.), which I suppose to have been drawn from a specimen obtained
at Stat. 144a, and therefore to belong to Amastigia hirkpatricki. In
describing a species obtained from more than one locality, it is always
expedient to indicate the locality of the specimens figured ; and this omission,
by Busk and others, gives rise to many difficulties.

With reference to Levinsen’s proposal to place this species in Caberea, I
think it desirable to exclude from this genus those species in which the.
rootlets pass down the margins of the branches instead of down the middle
of the basal surface. I think that C. rudis, Busk, should be placed in
Amastigia, and that C. crassimarginata, Busk, may be referred to the same
genus. It must be admitted that A. kirkpatricki has a considerable claim to.
be included in Serupocellaria, which it resembles in its mode of bifurcation
(Pl. 17. fig. 20). The branches are well jointed at their commencement,
but a median series of zocecia, usually commencing near the proximal end of
the internode, is present, in opposition to what is found in typical Serwpo-.
cellaria. The first median zowcium (EH) originates from a zocecium (“ A’’)
situated on the outer side of a branch with reference to the preceding
bifureation.- It gives to a second (1”), which may be followed by others in
a median row. he distal member of this series (H*in fig. 20) behaves as.
an axillary zocecium. |

A. kirkpatricki thns seems to unite the characters of more than one genus,
which may be explained by the assumption that Amastigia represents, better:
than any other genus, the ancestral character of the Scrupocellariidee, and
that its species show approaches in various directions to those of other genera.
I have suggested above that the pluriserial condition was probably antecedent
to the biserial condition ; and on this view A. kirkpatricki is a species which

338 SIK SIDNEY I. HARMER ON

has almost become biserial, while its basal heterozocecia have acquired the
full yibraeular form, so as closely to resemble those of Serupocellaria.
A. kirkpatrickt from Marion Id., A. gauss from the Antarctic, and A. erassi-
marginata from off Buenos Aires all come from localities within the range of
Amastigia and apparently outside that of Serwpocellaria ; and their distri-
‘bution supports the conclusion I haye come to on other grounds as to their
generic position.

The vibraculum (fig. 47) possesses a rootlet-foramen of the usual kind,
‘situated opposite the proximal end of the muscular mass and given off from
a small chamber separated by a calcareous wall from the rest of the vibraculum.
The figure shows what appears to be another rounded chamber on the distal
side of the rootlet-chamber. In side view this is seen to be continuous with
the main vibracular chamber, and it is really the foot by which the vibraculum
is connected with the proximal end of its zocecium, a communication-pore
occurring in the wall which separates it from the zocecium.

9. AMAsTIGIA GaussI (luge).
Scrupocellaria gausst, Kluge, 1914, p. 609, pl. 27. figs. 38,4. Antarctic.
Closely allied to A. kirkpatricki, and differing from it mainly in possessing
-a scutum, and in being 2—5-serial.

10. Amasticra SoLIDA (Kluge).
Scrupocellaria solida, Kluge, 1914, p. 611, pl. 27. figs. 7, 8. Antarctic, 350-385
metres. :

Unjointed, 3-serial. Stalk of scutum connected with the middle of its
lamina, which is lobed externally. Frontal avicularia occur, but marginal
avicularia are not described. Basal avicularia paired, directed transversely
inwards, the rostrum rather more vibraculoid than in the last species.
Median zocecia reduced on the basal surface.

11. Amasticia ANTARCTICA (Kluge).
Anderssonia antarctica, Kluge, 1914, p. 618, pl. 38. figs. 3. 4. Antarctic, 76-170
metres.

Unjointed, 6—8-serial. Scutum and spines wanting, the opesia with
‘straight lateral margins. The marginal zocecia face outwards, and they form
the whole of the basal surface. Frontal avicularia present. The lateral
avicularia described appear to be morphologically basal avicularia, since they
are placed on the proximal and not on the distal ends of the zowcia. They
are large and are arranged alternately on the basal surface, the rostrum
directed obliquely proximally and being linear distally. A rootlet-foramen
is present, a structure which seems to decide the question that these are basal
heterozoccia.

This species differs from the others described by Kluge in the absence of
spines and scuta ; and in these respects it has a considerable resemblance to

CELLULARINE AND OTHER POLYZOA. 339

the Craspedozoum group of Menipea. Its reference to Amastigia seems to be
indicated by its well developed basal heterozocecia and by the complete
exclusion of all the zoccia except those of the marginal rows from the basal
wall.

12. AMASTIGIA PATERIFORMIS (Busk).
Menipea pateriformis, Busk, 1884, p. 22, pl. 5. figs. 4, 4a, Off Valparaiso, 2160
fathoms.

The type-specimen (Brit. Mus. 87.12.9.101) has not been satisfactorily
described by Busk. It appears to be allied to A. antarctica, which it
resembles in the absence of scutum and spines. It agrees with Amastigia in
possessing basal heterozocecia, which may be regarded as avicularia, although
the mandible is distally linear and rather long. The rostrum is directed
obliquely proximally. The colony is apparently unjointed, the branch thick
and semicylindrical, the marginal zocecia facing obliquely outwards ; a median
row of zoccia, or two submedian rows, occurring in parts of the colony.
Rootlets in two lateral groups. Frontal avicularia stout and prominent, the
mandibles broadly triangular proximally, becoming narrow distally. Ovicells
large, and distinetly long and narrow.

2. Menrera, Lamourouer.
Menipea, Lamoureux, 1812, p. 183.
Craspedozoum, MacGillivray, 1886, Trans. Proc. R. Soc. Vict. xxii. p. 131.
Flabellaris (pars), Waters, 1898, J. Linn. Soc. xxvi. p. 672.
Flabellina, Levinsen, Vid. Medd. Naturh, Foren, Copenhagen, 1902, p. 21.
Cellularia (pars), Busk, et auctt.

Zoarium jointed (rarely unjointed) ; the joints, if present, traversing the
opesia of each of the inner zocecia at a bifurcation (Types 17, 18, Pl. 17.
figs. 17, 18). Branches biserial to pluriserial. Scutum wanting. Frontal
and marginal avicularia present or wanting ; basal heterozocecia found only
in M. vectifera. In several species a frontal avicularium may grow inwards
into the body-cavity, instead of occupying the position normal to these
avicularia. Rootlets associated with a bifurcation originating on the distal
side of the joint. The rootlets are nearly always in lateral bundles, as in
Amastigia and Notoplites.

The genus Menipea, as usually understood, embraces a number of species
which are apparently not nearly allied. I lay special stress, in re-defining
the genus, on the fact that (in all the jointed species) the joint traverses
the opesia of the znner zocecium, in each branch. This may happen in such
a way that a portion of the opesia lies on each side of the joint (Pl. 17.
fig. 17); or the proximal end of the opesia may lie in the region of the
joint (fig. 18). The relation of the joint to the opesia is implied by Busk
(1852?, p. 21) when he states that in I. cirrata one of the lower zocecia in the
internode is more or less aborted.

340 SIR SIDNEY F. HARMER ON

The species here included in Menipea form a group with a considerable:
resemblance to one another in general facées. It will be observed that (No. 2
excepted) they are all Southern forms, and that none of the Northern species.
are included:—a result which has already been anticipated by Norman

(1903, A. M. N. H.(7) xi. p. 578).

1. Menrexza orispa (Pall.). (PI. 17. fig. 17.)
Cellularia crispa, Pallas, 1766, p. 71.
Cellaria cirrata, Ellis and Solander, 1786, p. 29, pl. Ge figs. d, DO. “ Kast Indies.”
Menipea cirrata, Busk, 18522, p. 21, pl. 20. figs. 1, 2. 8S. Africa.
Menipea crispa, Marcus, 1922, p. 11.

Nasily recognizable by the characteristic inflexion of the branches, described
by Pallas and well represented in Hllis and Solander’s fig. d, and by the very
long, cylindrical frontal avicularia (Busk, fig. 1), coated on the proximal
border of the opesia, which they overarch. There are typically six zocecia in
the internode, which commonly has a single, large lateral avicularium.
Three strong, curved spines, jointed at the base, are commonly present on
the outer side of the distal end of the zocecium.

2. MENIPEA FLABELLUM (Ll. and Sol.).

Cellaria flabellum (pars), Ellis and Solander, 1786, p. 28, pl. 4. figs. ¢, C.

Bllis and Solander state that two distinct species are included in their
account of C. jlabellum, one from the Bahama Is. and the other from the
B. Indies. From the description it would appear that the figured specimens.
are from the Bahamas, although they might quite well be taken to represent
the South African form to which this name has usually been applied.
Perhaps a mistake in the locality of the specimens was made in the original
account ; but in view of the uncertainty of the question I think it best to
use the name given by Busk, as indicated in the next species, for the
common 8. African species. /. flabellum, Hill. and Sol. may be regarded as _
at present unrecognizable.

3. MenIPEA oRNATA (Bush).
?Cellularia floccosa, Pallas, 1766, p. 70.
Cellularia ornuta, Busk, 1852*, p. 20, pl. 26. figs. 8, 4. Algoa Bay.
Menipea flabellum, Busk, 1884, p. 21. Simon’s Bay, Cape of Good Hope.
A Marcus, 1922, p. 13, figs. 7 a-e.
labetlanes flubellum, Waters, 1898, J. L.S. xxvi, p. 672.

Tnternodes often increasing regularly in width from the proximal to the
distal end, their lateral margins nearly straight ; the number of constituent
zocecia reaching at least 13. Marginal avicularia and spines wanting.
Frontal avicularia inconspicuous, not prominent, often confined to the zocecia
BE, F, and G; when on F and G oceurring on the part of the zocecium
on the proximal side of the joint, as shown in Busk’s fig. 3,

CELLULARINE AND OTHER POLYZOA, 341

4, MENIPEA PATAGONICA, Busk.
Menipea patagonica, Busk, 1852, p. 22, pl. 23. fig. 1; pl. 25. figs. 1-8; pl. 26.
figs. 1,2. Falkland Is., Patagonia.
Menipea patagonica, Jullien, 1888, Miss. Sci. Cap Horn, vi. p. 71. Tierra del Fuego.
? Serupocellaria patagonica, Kluge, 1914, p. 615, text-fig. 4. Kerguelen.
Internodes commonly consisting of six zoccia. A single spine typically
present on each side, or two on the outer side, distally. Marginal avicularia
well developed, frontal avicularia small.

5. MENIPEA MARIONENSIS, Busk. (PI. 17%. fig. 22; Pl. 19. figs. 43-45.)
Menipea marionensis (pas), Busk, 1884, p. 21, pl. 4. figs. 3, 3a@ (specimens from
Stat. 142, Cape of Good Hope, 150 fathoms, only).

Branches mainly biserial, the axillary zocecium sometimes preceded by a
more proximal median zocecium. Internodes usually of 3-5 pairs of zocecia,
in addition to the median zocecium or zocecia, but sometimes with a larger
number, particularly in those with ovicells. Zoccia facing obliquely
outwards, from a raised keel passing down the middle of the branch. Opesia
nearly oval, but distinctly wider distally, the eryptocyst more developed
proximally than elsewhere. Strong calcareous spines given off basally from
the edge of the eryptocyst, in young zocecia, becoming more numerous with
increased age, and some of them directed frontally. Spines usually two
external and one internal, often one on each side, the external spine the more
distal. Scutum wauting. Frontal avicularia single on most of the zoccia,
tbe rostrum directed obliquely proximally ; not quite reaching the opesia.
Two symmetrically placed frontal avicularia may occur on the axillary
zowcium, where two median zocecia are present. Frontal avicularia of the
usual type may be replaced by internal avicularia (Pl. 17. fig. 22; Pl. 19.
figs. 43, 44), situated in the body-cavity. Marginal avicularia moderate,
constantly present. Vibracula and basal avicularia wanting. Ovicells very
short, wider than long. At the bifurcation (fig. 22) both branches are
jointed, the distal end of the parent-internode being prolonged beyond the
axil like the base of a tuning-fork. The joints are thus at some distance
from the axil and traverse the proximal end of the opesia of each inner
zoecium. Zoccia F and G meeting in a long median suture on the basal
side of H.

The material which was described by Busk as J/. marionensis included two
very distinct species, as has been explained above (p. 336), under Amastigia
kirkpatricki. The ‘Challenger’ specimens (Brit. Mus.) referable to AZ. marion-
ensis are four slides, 87.12.9.94, 95 and 99.7.1. 698, 699, all from Stat. 142 ;
87.12.9.94 being marked as the type.

In examining the spirit material of MJenipea triseriata obtained by the
‘Challenger’ from Simon’s Bay, Cape of Good Hope, I found a frag-
ment of another species, the original of my figs. 22, 43, and 45 of this

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 20

34.2 SIR SIDNEY F. HARMER ON

species. Its determination was confirmed when I remounted in Canada
balsam one of Busk’s dry slides (99.7.1.698) of MZ. marionensis from Stat.
142. Fig. 44 has been drawn from Busk’s slide thus treated.

The calcareous spines developed from the eryptocyst are one of the most
striking features of a Canada balsam preparation of MJentpea marionensts.
They are relatively few in young zocecia, in which they are directed towards
the basal surface (figs. 22, 43,44). In the older zocecia (fig. 45) they are
much more numerous, some of them being branched, and some of them being
directed towards the frontal membrane. These spines appear to be of the
same nature as the similar structures which have been described by Levinsen
(1909, p. 188, pl. il. fig. 7, c-e) in Menipea roborata. As in other members
of its family, cach distal zocecium of Menipea marionensis overlaps its
predecessor to a considerable extent on its basal surface, as indicated in
fig. 22. In the older zocecium represented (fig. 45) p is the proximal Wall
of the distal zocecium, and the opesia has been restricted by the formation of
a horizontal calcareous lamina, the free edge of which is in the form of a
Gothie arch (e) ; while the opesia is further reduced by the enlargement of
the proximal cryptocyst.

The remarkable internal avicularia of this species are described above, in

Sect. III.

6. MenipHa TRISERIATA, Busk. (Pl. 17. fig. 18; Pl. 19. figs. 40-42.)
? Crista elegans, Lamouroux, 1821, p. 6, pl. 65. figs. 4-7. Cape of Good Hope.
Menipea triseriata, Busk, 1852°, p. 22, pl. 28. figs. 2-4. 8, Africa.

Busk, 1884, p. 21. Cape of Good Hope.

% i Waters, 1898, J. L. 8. xxvi. p. 672, pl. 48. fig. 12.

Internodes more or less triserial, composed of numerous zocecia. Hxternal
spines 2, internal 2, on outer zocecia; 1, 1 on median zocecia. Cryptocyst
strongly developed on the proximal side of the opesia, covered with minute
tubercles, its free margin here strongly thickened, and usually produced into
a strong median tooth which may be directed basally. Marginal avicularia
on most of the zocecia. Frontal avicularia often two on the median zocecia,
one of which may be replaced by an internal avicularium ; reversed (the
rostrum directed distally) on the distal side of an ovicell, an arrangement
commonly found in this genus. The outer frontal avicularium is wanting
on the marginal zowcia. Ovicells short, usually broader than long.

The British Museum possesses a number of slides, all from South Africa.

A specially interesting feature of this genus is the possession of internal
avicularia (figs. 41, 42), which have been described above, under Sect. ITI.

7) ”

7. MENIPEA QUADRATA (Bush).
Cellularia quadrata, Busk, 1884, p. 18, pl. 5. figs. 5-5 b. Kerguelen, Heard Island,
Internodes long and narrow, consisting of numerous zocecia. Spines and
frontal avicularia wanting. Marginal avicularia small.

CELLULARINE AND OTHER POLYZOA. 343

8. MENrIPEA FLAGELLIFERA, Busk.

Menipea flagelfera, Busk, 1884, p.21, pl. 4. figs. 1-16. Kerguelen, Marion Id.,
Straits of Magellan, W. of Falkland Is.
Serupocellaria flagelhfera, Kluge, 1914, p. 615, text-fig. 5. Kerguelen.

Jnternodes of moderate length. Usually a single external spine. Mar-
ginal ayicularia large. Frontal heterozocecia vibraculoid.

The frontal heterozocecia of this species are so peculiar as to suggest the
possibility that it should be placed in a new genus. I refer it to Menzpea
because it has the mode of bifureation characteristic of the genus, the joints
traversing the opesia of each inner zocecium. The frontal heterozocecia,
which are constantly present, have the seta of a vibraculum, but they do not
possess the rootlet-foramen which is always found in a basal heterozocecium,
whether it is an avicularium or a vibraculum. In view of the fact that in
Amastigia avicularia and vibracula are interchangeable on the basal surface,
a corresponding plasticity may perhaps be conceded to Mentpea. The struc-
tures in question replace the frontal avicularia of other species ; and, in

spite of the presence of a greatly elongated mandible, I think they may be
regarded as avicularia.

9. Munipea spicata (MacGillivray). (Pl. 17. fig. 19; Pl. 18. figs. 32,
34, 35.)
‘Craspedozoum spicatum, MacGillivray, 1886, p. 182, pl. i. figs. 2,2a@; 1889, Prodr.
Zool. Vict., Dec. xviii. p. 283, pl. 177. figs. 3-36. Victoria.

Unilaminar. Internodes curved frontally, composed of numerous zoecia,
‘biserial at their origin, becoming 4-serial distally. Bifurcation cymose,
owing to the fact that only one of the branches is jointed; the ultimate
branches being unilateral cymes and the larger stems being bilateral (fig. 19)..
While the lateral internodes give off a single branch, those of the larger
‘stems give off two branches, one on each side. The proximal branch comes
off near the commencement of the internode, and forms the commencement
of a lateral branch. The distal branch is placed in the line of the main
stem, of which it forms a segment, and the distal, unjointed termination of
the parent-internode diverges from the stem, on the same side of it as the
proximal branch. The mode of branching of I. spicata does not seem to
have been previously noticed, but a similar suppression of one of the joints
occurs also in the allied genera Notoplites and Tricellaria. The affinity of
M. spicata to the other species which I place in Menzpea is shown by the fact
that the joint traverses the opesia of an inner zocecium.

M. spicata has two well-developed spines on the outer side of the marginal
zocecia and one on the inner side ; and one spine on each side of the median
zocecia. The cryptocyst is strongly developed on the proximal side of the
-opesia, and calcareous spines project into the body-cavity from the lateral
walls, as in MW. roborata. The ordinary frontal avicularia do not occur on all

25*

344 SIR SIDNEY F. HARMER ON

the zocecia, but when present they are large and prominent. They are very
asymmetrical, the rostrum much elevated, and situated at the outer side of
the zocecium, indicating that the avicularium is one of the pair found in
other species, its direction being as required by this hypothesis. Internal
avicularia (figs. 32, 35) occur, always in zowcia which do not succeed an
ovicell, and, as noticed on p. 319, they are concealed by the proximal
eryptocyst, in frontal view. They are considerably smaller than in M. robo-
rata, and they lie close to one side of the zocecium, being directed almost
vertically towards the basal surface. Ovicells occur only on the two median
rows, and two strong calcareous thickenings of the frontal wall converge
distally, uniting with one another to form a strong recurved spike projecting
in the middle line beyond the ovicell (fig. 34). The basal walls of the
median zocecia are considerably constricted near the middle (fig. 35).

This and the next two species were placed in Craspedozoum by MacGillivray
(1886, Trans. Proc. R. Soc. Vict. xxii. p. 131), who instituted this genus for
their reception. They are all provided with the lateral bundles of rootlets
which usually occur in Menipea. The pore-chamber for the rootlet (fig. 35)
is situated in the usual place, at the proximal end of a marginal zocecium,
and it probably represents a vestigial basal heterozocecium. It is incon-
spicuous in M. spicata, in which there are no blister-like pore-chambers on
the terminal walls, at their insertion into the basal wall.

The British Museum possesses several slides of Victorian specimens
(88.11.14.97, 298; 97.5.1.459, 460; 83.10.15.46).

10. Menrera ticuLara (MacGillivray). (PI. 18. figs. 31, 33.)
Craspedozoum ligulatum, MacGillivray, 1886, p. 132, pl. 1. figs. 3-3a; 1889, Prodr.
Zool. Vict., Dec. xviii. p. 283, pl. 177. figs. 1,14, 2. Victoria.
Flabellaris roborata, var. ligulata, Waters, 1898, J. Linn. Soc. xxvi. p. 672.
Menipea ligulata, Levinsen, 1909, p. 140, pl. 2. figs. 8 a-e.

Unilaminar, the frontal surface convex, with the marginal zocecia facing
partly outwards, the basal surface flat or concave. Branches composed of as
many as nine rows of zoccia, unjointed. Proximal eryptocyst less extensive
than in M. spicata, usually prolonged into a thick calcareous spike or column
(tig. 81), descending basally into the body-cavity, on one or both sides. When
paired these spikes are not on the same level, one being usually just visible
in frontal view, at the proximal end of the opesia, the other completely con-
cealed by the eryptocyst and best seen in a basal view of a Canada balsam
preparation. Spines 1, 1, well developed. Frontal avicularium single,
symmetrical, its rostrum broadly triangular and directed proximally, shorter
and wider than in J/. spicata ; not developed on the distal side of an ovicell.
Internal avicularia (fig. 33) rather short, nearly horizontal, the base wide,
the rostrum just appearing in the proximal end of the opesia. Ovicells
longer than wide, the two calcareous bars uniting in a wide Gothic arch near
the distal end, with a short median mucro. Basal walls of zocecia strongly

CELLULARINE AND OTHER POLYZOA. 345

constricted laterally, the arrangement being as in Amastigia rudis (PI. 1%.
fig. 27), the basal wall correspondingly sinuate. Branches bordered by lateral
groups of rootlets, as in the allied species, given off from pore-chambers at
the proximal ends of the marginal zocecia. The terminal walls may or may
not be provided with blister-like pore-chambers at their insertion into the
basal wall.

Victorian specimens, Bracebridge Wilson Coll., in British Museum,
97.5.1.453,454.

Waters regards this as a unilaminar variety of J/. roborata, but I think its
characters entitle it to specific rank.

11. MenipEa roporata (Hincks). (Pl. 18. figs. 28-30.)

Membranipora roborata, Hincks, 1881, A. M. N. H. (5) viii. p. 128, pl. 2. figs. 8, 3a,
Curtis Id., Bass Straits; 1892, Zbid. (6) ix. p. 331.

Flustra membraniporides, Busk, 1884, p. 54, pl. 32. figs. 7a,6. Port Jackson, Bass
Straits.

Craspedozoum roboratum, MacGillivray, 1886, p. 131, pl. 1. fig. 4; 1889, Prodr. Zool.
Vict., Dec. xviii. p. 284, pl. 177. figs. 4,5, 5a. Victoria.

Flabellaris roborata (pars), Waters, 1898, J. L. 8. xxvi. pp. 660, 662, 672

Flabellina (Flabellaris) roborata, Levinsen, Vid. Medd. Naturh. Foren. Copenhagen,
1902, p. 21.

Menipea roborata (pars), Waters, net: A. M.N. H. (5) xx. p. 183.

Menipea roborata, Levinsen, 1909, pp. 3, 9, 131, 132, 138, pl. 2. figs. 7 a-k (Llabellina
on plate).

Bilaminar, unjointed, pluriserial, with at least 17 rows of zocecia, the
branches bordered by marginal bundles of rootlets, widening terminally and
assuming an almost Flustrine appearance. Basal walls of the median zocecia
only slightly constricted, laterally, near the middle. Spines 1,1. Crypto-_
eyst less developed than in MZ. ligulata. Thickening bars of ovicells uniting
in the form of a Gothic arch and more or less acuminate ; or in a rounded
curve, without a mucro. Frontal avicularia paired (except on the lateral
zoccia, where the avicularium is single) close together, the rostrum directed
obliquely proximally ; two constantly present on the distal side of an ovicell,
widely separated, their direction reversed. Internal frontal avicularia large,
directed nearly horizontally, on the basal side of the cryptocyst; their distal
end visible at the proximal end of the opesia. Rootlet pore-chambers of the
marginal zocecia inconspicuous. Blister-like pore-chambers on the proximal
side of the insertion of the terminal walls into the basal wall present or
absent.

The synonymy shows that there has been much difference of opinion with
regard to the generic position of the present species; but I agree with
Waters and others that it must be placed in Menipea. Hincks (1892) and
Waters have both expressed the opinion that I. ligulata and M. spicata are
varieties of this species. That it actually belongs to Menipea is indicated by
MM. spicata, which has the typical unilaminar character of the genus, while

9

346 SIR SIDNEY F. HARMER ON

retaining the characteristic position of the joint at the bifurcation. This
species has a close resemblance to M. spicata in all essential details of
structure ; and its internal avicularia (figs. 29, 30) found also in other
species (see Sect. III.) are a striking point of agreement. Another resem-
blance is the oceurrence in both species of calcareous spines projecting into
the body-cavity, as originally described by Levinsen. On separating the
two laminze of a branch of M. roborata, after boiling in Hau de Javelle, it is
found that while rosette-plates are wanting in all the zocecia of the median
tows, typical lateral rosette-plates occur on all those of the two marginal
rows. WM. ligulata shows a distinct tendency to curve the margins of its
branches towards the basal surface. If this process were continued until
the edges united to form a hollow unilaminar cylinder, with the basal
surface internal and the frontal surface external, and if this cylinder were
then compressed so that the cavity disappeared and the basal walls of opposite
sides came into contact, a bilaminar branch would result. At euch lateral
margin the external zocecia would be united by rosette-plates, while none of
the others would be thus united. This is the arrangement actually found in
M. roborata.

Specimens (slides) in the British Museum: 97.5.1.455, 457, Victoria,
Bracebridge Wilson Collection ; 81.10.21.352-354, Port Jackson ; 50.5.2.2.

12. MmNIPEA VECTIFERA*,n. sp. (PI. 17. fig. 23; Pl. 18. figs. 36-39.)

Unilaminar, the frontal surface flat, the basal surface also flat or slightly
convex; the branches composed of as many as seven series of zocecia, and
bordered by lateral bundles of rootlets. Apparently not normally jointed,
although secondary breaks are formed across the branches, becoming
converted into joints by the development of irregular chitinous tubes.
Habit as in M. spicata, but less calcified. Cryptocyst (fig. 23) finely tuber-
cular, the proximal part extensive, especially in the marginal zowcia, the
free edge narrow on the distal side of an ovicell. A pair of long bars, °
expanding at their free ends in a palmate manner (figs 37, 39), descend
from the thickened edge of the proximal eryptocyst into the body-cavity,
their direction being nearly vertical, but with a slight slope proximally.
Spines small, 2, 2, the inner or distal pair commonly vestigial and recognizable
by their tubular cavities trasversing the oral frame, as seen in a Canada
balsam preparation. Frontal avicularium single, asymmetrical, commencing
at one edge of the proximal cryptocyst, the rostrum directed transversely or
slightly proximally, the mandible rather elongated and linear distally. On
the distal side of an ovicell two frontal avicularia occur, widely separated,
their rostra reversed so as to point distally and outwards. Internal
avicularia (fig. 36) found in several cases in the more proximal zocecium of
the two formed when a row is doubled, close to the concave side of the’

* Veetts, a bar ; referring to the two bars borne by the proximal cryptocyst.

CELLULARINE AND OTHER POLYZOA. 347

zocecium and directed more or less basally ; concealed beneath the proximal
eryptocyst in frontal view. Marginal avicularia (fig. 23), on the marginal
zocecia, small, with a deep cavity excavated in the distal part of the outer
calcareous wall. Basal surface of zocecia moderately constricted laterally,
the terminal wall showing one, two, or even three blister-like pore-chambers
projecting into the proximal zoccium, close to the basal wall. A con-
spicuous pore-chamber giving off a rootlet at the proximal end of each
marginal zoccium, situated just distally to the marginal avicularium. A
single basal heterozocecium (fig. 38) sometimes present at the angle of the
bifurcation, its rostrum narrow and elongated distally. Ovicells without
distinct thickening bars, the free edge of the ectocecium forming a rounded
curve, passing near the sides and distal margin, and not giving rise to a
mucro. Operculum distinct and Membraniporiform, without basal sclerite,
but with well-marked lateral, triangular, occlusor flanges.

Described from three specimens in the British Museum Collection, all
from New Zealand :—Hincks Collection, 99.5.1.630 (Type) and 458 (labelled
Mentbranipora roborata) ; Busk Collection, 99.7.1.703 (labelled Menipea
multiseriata).

This interesting species would be referable to Craspedozowm if that genus
were recognized as distinct. In the presence of internal avicularia it
agrees with the other three species which were placed in that genus by
MacGillivray. But this feature allies the Craspedozowm group with typical
species of Menipea such as M. triseriata and M. marionensis, while C. spicatum
shows a further agreement with Menipea in the character of ifs jointing.
The present species, in retaining a single basal heterozocecium at the bifur-
cation, indicates that these structures form a part of the original inheritance
of the genus, as in other genera of Scrupocellariide, and notably Amastigia
and Notoplites. The presence of marginal avicularia, which I have not
found in other species of Craspedozoum, is a further feature of interest.

13. Menipra muLtisEertara, Bush.
Menwpea multiseriata, Busk, 1852 *, p. 22, pl. 60. figs. 1, 2. New Zealand.

Unilaminar, multiserial, jointed, bordered by marginal groups of rootlets,
the number of series of zocecia at least 8. Basal walls of the zoccia hardly
contracted laterally. Spines 1,1. Ovicells closely resembling those of
M. roborata. Frontal avicularia 2 on the median zoccia, the rostrum
directed obliquely proximally ; two constantly present on the distal side of an
ovicell, the rostrum directed nearly transversely outwards; one on the
marginal zoccia. A small lateral avicularium on the marginal zocecia.

Busk considered this species nearly allied to M/. triseriata, and possibly a
variety of it. The locality was said to be unknown by him. The type-slide
(99.7.1.112.D) was not returned to the Museum on the completion of the
Catalogue, but it came later, with the remainder of the Busk Collection

348 SIR SIDNEY F. HARMER ON

received in 1899. It was then unnamed, but it had been marked by Busk
“New Zealand.” The specimen was determined by Mr Kirkpatrick, in
1904, as the missing type-slide. The evidence in favour of this view seems
to me conclusive, A portion of the specimen exactly resembles Busk’s
fig. 1; and there are certain agreements between the specimen and the
figure, particularly the occurrence of a minute emargination in the outer
outline of the uppermost marginal zocecium, on the left side, which show
that the figure had been drawn with great accuracy. It must therefore be
assumed that Busk had obtained evidence, after the publication of his
description, that the figured specimen came from New Zealand. It may
here be noted that Waters (1887, A. M.N.H. (5) xx. pp. 84, 183), who does
not admit the specific distinctness of M/. roborata and the allied forms, and
has stated that a New Zealand unilaminate form is jointed, may have had
M. multiseriata under observation. It is possible, on the other hand, that he
was referring to M. vectifera.

It is remarkable that the resemblance of M. multiseriate to M. roborata
has not previously been noticed. I have felt some hesitation in separating
them, but I think this procedure may be justified on the following grounds:
M. multiseriata is unilaminar. I do not think it isa specimen of JZ. roborata
in which the laminz had become separated, because the marginal zocecia
show no rosette-plates and because occasional rootlets pass across the
basal surface. The species under consideration has marginal avicularia,
which do not occur in A. roborata; and it is jointed, another definite
difference. In a normal bifurcation of the type-specimen it appears to me
that only one of the branches of the fork is jointed, and that in this one the
joint traverses the opesia of the innermost zocecium. The jointed branch
consists of only 2 or 3 zocecia at its base. In other parts, secondary joints
occur as in certain other species of Menipea. The joint in these cases
traverses the whoie width of a branch, without having any relation to a
bifurcation. It has clearly been formed by absorption of the calcareous
matter, but the separate edges are joined by irregular chitinous tubes, which
have the appearance of being new formations, and not simply the chitinous
lining of the zone of the zocecium which lies in the region of the joint.

I have been unwilling to interfere with the type-specimen to the extent
that would be necessary in order to obtain evidence with regard to the
occurrence of internal avicularia; and I must leave this point undecided.

3. Noropuires *,n. gen. Genotype, JV. rostratus, n. sp.
Cellularia (pars), Menipea (pars), and Scrupocellaria (pars), auctt.
Bifurcation of type 15 (fig. 15), one or both branches jointed, the proximal
segments of F and G in contact and much longer than the corresponding

* voroy, back ; omdirys, an armed man; in allusion to the basal avicularia.

CELLULARINE AND OTHER POLYZOA. 349

parts of J and K. The joints are on the distal side of C and D, and traverse
the proximal ends of GK and FJ without passing through an opesia.
Rootlets given off on the distal side of the joint. Zoccia elongated, the
opesia relatively short. Basal avicularia, approaching a vibraculoid form,
typically present, and then usually confined to the neighbourhood of the
axils ; the rootlets given off by them or from other parts closely applied to
the margins of the branches and strengthening the axils. Scutum, frontal
and marginal avicularia present or absent. Ovicells typically large, and with
a frontal fenestra. Zoarium biserial.

I propose this genus primarily for several deep-water species represented
in the ‘Siboga’ Collection, all of them supported by an elongated stalk
composed of parallel rootlets, which diverge at the base to form an anchoring
tuft—a type of growth frequently found in Cheilostomes from deep water.
I select as the genotype JV. rostratus, of which I give a preliminary diagnosis ;
this species representing the genus in what I consider the full expression of
its characters, some of which are lost in other species. The features which I
regard as specially characteristic are: (1) the mode of bifurcation, the joints
being at a considerable distance from the axil (a convenient way of
recognizing members of the genus), and the position of the joints, which are
completely on the distal side of C and D; (2) the occurrence of basal
heterozocecia, commonly confined to the bifurcations. Certain species do not
possess these structures, but their affinity is indicated by the mode of
branching.

Notoplites appears to be allied to Amastigia, as shown by the occurrence of
basal avicularia ; but it differs from that genus in being biserial and jointed,
in the longer zocecia, and in having the basal avicularia confined, as a rule, to
the bifurcations. It differs from MJenzpea in the fact that the joint does not
traverse the opesia of the inner zocecium at a bifurcation. It will be observed
that the genus, as here understood, occurs from the Arctic to the Antarctic
Oceans, but that a connecting-link is afforded by the ‘Siboga’ species, from
the neighbourhood ot the equator.

1. NoroPLiTes ROSTRATUS, n. sp.

Zoarium reaching a length of 55 mm., attached by a proximal tuft of
rootlets which separate from a main stem, about 20 mm. long, composed of
parallel rootlets. Zocecia long and narrow, their outer outline concave ; the
opesia, which oecupies less than half the frontal length, oval, much reduced
proximally by a broad, crescentic eryptocyst. Distal spines 2 external,
1 median, | internal, with another arising from the basal surface. Scutum
jointed at the base, the lamina lobed, its distal portion the smaller and
generally acutely pointed and narrow. Frontal avicularium, in ordinary
zocecia, variable in size, elevated, at some distance from the opesia, on the
inner side of the cryptocyst, reclining against the opesia of the preceding

350 SIR SIDNEY F. HARMER ON

alternate zocecium, the rostrum linear and directed obliquely inwards and
distally. Frontal avicularium succeeding an oyicell gigantic, directed trans-
versely outwards, the upcurved rostrum resting on the distal border of the
ovicell, the much elongated linear mandible on the distal side. Marginal
avicularia varying in size, usually small, the rostrum directed obliquely
outwards, basally and proximally ; occasionally gigantic ; on the fertile zocecia
gigantic, originating slightly on the basal side, the rostrum directed distally,,
nearly parallel to the edge of the branch, the distal half free, the mandible
on the outer side. Basal avicularia (one pair) occur on the inner zocecia
succeeding the joints, inflated proximally, the long linear rostrum directed
distally ; each giving off a rootlet at its proximal end, in line with itself, the
rootlet running closely attached to the margin of the axil. Ovicells large and
very long, imperforate except for a transversely elongated fenestra, on the
frontal surface, near the orifice.

‘Siboga’ Expedition, Stat. 211, 5° 40’ 7” S., 120° 45/ 5” H., 1158 metres.

Various species which have been referred to other genera appear to belong
to Notoplites; and, in some of these, basal avicularia have already been
described or have been found by me in specimens in the British Museum
Collection. In other species basal avicularia seem to be wanting ; but here,
as in other genera of Scrupocellariide, it may be assumed that an important
generic character may fail to develop, the affinities of the species being
indicated mainly by the mode of bifurcation.

(a) Species in which basal avicularia occur.

2. NoropuirEs BILOBA (Bush).
Cellularia biloba, Busk, 1884, p. 18, pl. 8. figs. 2-26. Azores, 900 fathoms.

Resembling NV. rostratus in habit, in the form of its zocecia, seutum,
marginal avicularia, and ovicells. Frontal avicularia wanting. The ‘ Chal-
lenger ’ slides (87.12.9.63, 64) show a pair of small transverse basal avicularia,
which were not noticed by Busk, at the bifurcation.

3. NoOTOPLITES CRATERIFORMIS (Bush).

Cellularia crateriformis Busk, 1884, p. 16, pl. 3. figs. 1-16. E. of Buenos Aires,
1900-2650 fathoms.

Resembling the preceding species in habit and in the marginal avicularia.
Scutum and frontal avicularia wanting. Of the ‘ Challenger’ slides (87.12.9.
51,52) 51 shows a pair of small basal avicularia, directed nearly transversely,
at a bifureation. These seem to be referred to by Busk, who says:
“Occasionally a radical tube may be seen supporting, instead of an ordinary
zocecium, a small curiously formed avicularium.”

CELLULARINE AND OTHER POLYZOA.

oo
Or
eR

4, NoToPLITES AVICULARIE (Yunagi § Okada).

Scrupocellaria avicularie, Yanagi & Okada, 1918, Ann, Zool. Japon. ix. p. 418, pl. vi.
fig. 4, text-fig. 4. Japan, 78 fathoms.

Scutum large, with a complicated cavity. Frontal and marginal avicu-
laria small. Basal avicularia small, the rostrum directed obliquely proximally
and inwards, not confined to the bifurcations, but occurring alternately, on
the proximal ends of the zowcia. The text-figure shows clearly the origin of
a rootlet from a basal avicularium.

5. NoToPLitEs MARSUPIATUS (Jullien).
Serupocellaria marsupiata, Jullien, 1882, Bull. Soc. Zool. France, vii. p. 506, pl. 13.
figs. 17-20. N.W. of Spain, 2018 metres; 1888, Miss.
Sci. Cap Horn, vi. Zool. p. 69.

4s a Calvet, 1907, Exp. Sci. ‘Travailleur’ et ‘Talisman,’ viii.
p: 377.
$5 Waters, 1888, ‘ Ohalleneer? Rep. pt. Ixxix. p. 9.

Meninee clausa, Busk, 1884, p. 20, pl. iv. figs. 5, 5a. W.of Azores, 1675 fathoms.

Jullien and Waters both pointed out in the same year (1888) that Busk’s
species is a synonym of S. marsupiata. A special peculiarity of the species is
that the convex seutum so completely fills the opesia that it appears at first
sight to be the frontal wall. It is attached to its stalk at the distal. inner
corner, and its free border is marked by characteristic radial slits, in the
‘Challenger’ specimen (87.12.9.83) at least. Frontal and marginal avicu-
laria small. Jullien states that he founda single basal vibraculum in this
species, and Waters points out that Kirkpatrick has found vibracula on the
basal side, on two zocecia.

(b) Species in which basal avicularia are not recorded.
6. NoroPLitEs ELONGATUS (Bush).
Cellularia elongata, Busk, 1884, p. 19, pl. 3. figs. 8-56. Kerguelen, 28 fathoms.
Scutum elongated, the distal lobe the smaller. The ‘ Challenger’ slides
(87.12.9.65, 66) show that this species agrees in its bifurcation with Noto-
plites. Marginal avicularia small, frontal avicularia wanting. I cannot find
any case in which two median zocecia occur, as shown in Busk’s fig. 3 a.

7. NoropLites JEFFREYSII (Vorman).
Menipea jeffreysii, Norman, 1868, Q. J. M.S. (a.s.) viii. p. 213, pl. 5. figs. 4-8 *. Shet-
land; 1898, A. M. N. H. (6) xii. p. 446, pl. 19. fig. 1. Hardanger
and Trondhjem Fjords, Finmark; 1903, Led. (7) xi. p. 579.
; 0 Hincks, 1880, p. 42, pl. 9. figs. 1,2. Shetland.
op . Nordgaard, 1918, Troms? Mus. Aarsheft. xl. (1917), p. 35.
In his 1893 paper Norman pointed out that the scutum is not correctly
described in his original account and in Hincks’ fig. 1, also referring to the

* The figures are wrongly given in the text as 3— 5, as pointed out by Norman (1903, p. 446).

O52 SIR SIDNEY F. HARMER ON

original specimen, but that it fills the entire opesia (in the same way as in
NV. marsupiatus, Jull.). He refers toa figure by Alder, reproduced in his
own original account (fig. 6), supposing that certain avicularia which appear
to be on the basal surface are really frontal avicularia seen through the
zocecia by transparency. It appears possible that these are really basal avicu-
laria, and in a specimen from Norway in the Cambridge Collection I have
observed structures on the basal surface which may perhaps be of this nature.

8. Novropiires smirrit (Vorman).

Cellularia ternata, forma duplex, Smitt, 1868, Ofy. K. Vet.-Akad. Férh. xxiy. (1867),
pp. 288, 312, pl. 16. figs. 25, 26. Spitsbergen.

Menipea snitti, Norman, 1868, Q. J. M.S. (n. s.) viii. p. 214.

3 p Hincks, 1880, p. 43.

Menipea duplex, Levinsen, 1887, Dijmphna-Togtets zool.-bot. Udbytte (Copenhagen),
p. 809, pl. 26. figs. 1, 2. Kara Sea.

Scrupocellaria smittu, Waters, 1900, J. L.S. xxviii. p. 57, pl. 7. figs. 8-11. Franz
Josef Land.

Hincks remarks of this species, which has frontal and lateral avicularia
but no scutum, that it is nearly allied to WM. jefreysii. The mode of branching
characteristic of the genus may be seen in Smitt’s fig. 25; but it is better
shown by Waters (fig. 8), who emphasizes its difference from certain other

species.

9. Noropiires anrarcricus ( Waters).
Serupocellaria antarctica, Waters, 1904, ‘Belgica’ Bryozoa, p. 25, pl. 1. figs. 5 a-e;
pl. viii. figs. 2a, b. Antarctic, 435-480 metres.
Kluge, 1914, p. 606, pl. 28. fig. 1. Antarctic, 350-385
metres.

cP) 7

Frontal and marginal avicularia and scutum present. Waters’ fig. da
shows the mode of bifurcation well.

The three species next following, described by Kluge from Antarctic
waters, may be referred to Notoplites, on the evidence of the bifurcation, as
shown in the figures. SS. watersi appears to bifurcate in the same manner ;
while S. perdita, in which there is no evidence with regard to this point, has
an ovicell resembling that of other species of Notoplites.

10. Novopiires TENUIS (Kluge).
Scrupoeellaria tenuis, Kluge, 1914, p, 608, pl. 27. fig. 2. Antarctic, 46-385 metres.
Frontal avicularia and scutum present. Marginal avicularia wanting.
Bifurcation clearly represented.

11. Noropuires DrYGALSKII (Kluge).
Scrupocellaria drygalski, Kluge, 1914, p. 609, pl. 27. fig. 5. Antarctic, 70-385
metres,
Frontaland marginal avicularia present. The scutum fills the opesia, as in
LV. jefreysu. Bifurcation figured.

CELLULARINE AND OTHER POLYZOA. 353:

12. NoropLites VANHOFFENI (luge).
Scrupocellaria vanhoffent, Kluge, 1914, p. 610, pl. 27. fig. 6. Antarctie, 350-385.
metres.
Frontal and marginal avicularia and seutum present ; differing from the
preceding species in the scutum, which has a complicated or “ cervicorn ”

cavity. Bifurcation figured.

3. NoropLitEs WATERSI (Kluge).
Scrupocellaria waterst, Kluge, 1914, p. 607, pl. 27. fig. 2. Antarctic, 350-385.
metres.
Frontal and marginal avicularia and scutum present. The bifurcation
represented in the figure is typical except for the zocecium B. I suspect
that a mistake was made with regard to this point.

14. ? NoropiirEs PERDITUS (Aluge).
Serupocellaria perdita, Kluge, p. 618, text-tig. 2. Antarctic, 2450 metres,
Bifureation not described. The scutum and ovicell resemble the corre-
sponding parts in NN. rostratus.

incerte sedis (possibly belonging to Notoplites).

15. Menipea normant, Nordgaard, 1900, Norske Nordhavs-Exp., p. 4, pl.i.
figs. 2-8. Off Norway, 408-1134 metres.

16. ? NoropLirEs SIMPLEX (Kluge).
Scrupocellaria simplex, Kluge, 1914, p. 607, pl. 27. fig. 1. Antarctic, 385 metres.

Branches biserial, the opesia elongated, oval, with much raised margin.
Seutum and spines wanting. A single frontal avicularium, and occasionally
a minute marginal avicularium. Ovicells unknown, basal heterozocecia not.
described. Rootlets at first passing down the margins of the branches, given
oft from a chamber at the proximal end of the zocecium. Kluge states that
this species is ordinarily unjointed, but that the branches may be jointed
*€ oberhalb der Gabelung.” The zowecium and the opesia resemble the corre-
sponding parts of Notoplites elongatus (Busk) ; and it seems possible that the
species should be referred to the same genus.

4. TriceLuarta, Fleming.
Tricellaria, Fleming, 1828, p. 540.
Cellarina, Van Beneden (pars), 1848, p. 70 (nec Cellarina, D’Orbigny, 1851, p. 181).
Ternicellaria, D’Orbigny, 1851, pp. 40, 47.
Bugulopsis, Verrill, 1880, p. 190.
Jellularia (pars) and Menipea (pars), auctt.
Zoarium jointed, the branches biserial. Internodes commonly constituted
by three zocecia, at least in the main stems. Zocecia more elongated than in
Emma, the opesia occupying a small proportion of the front and without a

354 SIR SIDNEY F. HARMER ON

large development of cryptocyst proximally. Bifureation of types 9-12
(figs. 9-12), the joints traversing the proximal ends of the zocecia CG and
DF; one or other of the joints occasionally suppressed. F and G usually
separated on the basal side of H, sometimes by its whole width, their
proximal segments shorter than the corresponding parts of Cand D. The
proximal ends of F and G may be suppressed, in which case the internode
consists of a single zowecium at its proximal end, and the joint consists of a
single chitinous tube. Rootlet given off on the proximal side of the joint.
Scutum, marginal and frontal avicularia present or wanting. Basal avicu-
laria and vibracula wanting, but probably represented by the rootlet pore-
chamber.
Among the species which I include in 77icellaria are the following :—

A. Internodes biserial at the proximal end. Joints composed of two
chitinous tubes (figs. 9-11).

1. TRICELLARIA TERNATA (Zl. § Sol.). (Fig. 9.)

Cellaria ternata, Ellis & Solander, 1786, p. 80. Aberdeen.
Menipea ternata, Hincks, 1880, p. 38, pl. 6. figs. 1-4. British coasts.

This common species has often been described. I may draw attention to
the possibility, mentioned on p. 324, that the structure from which a rootlet
may arise in this species may represent a basal heterozocectum—a structure
-otherwise absent in the genus.

La. TRICELLARIA TERNATA (Zl. § Sol.), var. GRACILIS, Smitt.

Cellularia ternata, forma gracilis, Smitt, 1868, Ofyv. K. Vet.-Akad. Forh. xxiv. (1867),
pp- 282, 805, pl. 16. figs. 10-16. Spitsbergen to Britain and Belgium.

Menipea gracilis, Busk, 1878, in Nares, Narr. Voy. Polar Sea ‘ Alert’ and ‘ Discovery,’
ed. by H. W. Feilden, ii. p. 284.

It may be observed that gracilis as the trivial name of this form is ante-
dated by Cellarina gracilis, Van Beneden, 1848 (see Sect. II.).

2. TRICELLARIA OCCIDENTALIS (7rash), 1873 *. (Fig. 10.)

Menipea occidentalis, Trask, 1873, Proc. Calif. Acad. Nat. Sci. i. (1854-1857), 2 ed.
p. 118, pl. 4. fig. 4.

By a Robertson, 1905, Univ. Calif. Publ., Zool. ii. p. 254, pl. 6. figs.
22-25. California.

i compacta, Hineks, 1882, A. M. N. H. (5) x. p.461. Queen Charlotte Is.

3. TRICELLARIA PRIBILOFI (Jtobertson).

Menipea pribilofi, Robertson, 1905, ¢. cit. p. 257, pl. 7. figs. 32, 33; pl. 8. fig. 34.
Alaska, islands of Bering Sea.

* The Proceedings of the California Academy for 1854-1857 were originally printed in a
newspaper, ‘The Pacific,” but were republished, verbatim, in 1878. Dr. Trask’s paper was
read at the meeting, Mar. 30, 1857.

oo ~

CELLULARINE AND OTHER POLYZOA. 355

4, TRICELLARIA SYMPODIA (Yanagi 5° Okada).

Menipea sympodia, Yanagi & Okada, 1918, Ann. Zool. Japon. ix. p. 410, pl. 6. fig. 1,
text-fig. 2. Japan, 250 fathoms.

As indicated by its name, this species is an instance of the sympodial
mode of growth, due to the suppression of one of the joints at the bifurcation.
The remarkably elongated, cylindrical frontal avicularia resemble those of
Menipea crispa and of ? Notoplites norman.

5. TRICELLARIA PEACHII (Bush). (Fig. 11.)

Cellularia peach, Busk, 1851, A.M.N. H. (2) vii. p. 82, pl. 8. figs. 1-4. British
coasts; Busk, 1852°, p. 20, pl. 27. figs. 3-5.

This common species has been frequently described. It appears to repre-
sent the extreme amount of loss of structures which typically occur in the
genus, as shown by the disappearance of the frontal and marginal avicularia
and of the scutum. It was made the genotype of Bugulopsis by Verrill ;
but its mode of bifurcation is that of Z7icellaria, with which I think it
‘should be placed.

6. TRICELLARIA MONOTRYPA (Bush).

Cellularia monotrypa, Busk, 1852", p. 368. Bass Straits.
5 cuspidata, Busk, 18527, p. 19, pl. 27. figs. 1, 2; 1884, p. 17.

In 1884 Busk remarked that it might be proper, conditionally, to revert to
his original name, monotrypa. There seems no question that this should be
done, as Busk was not justified by the Rules of Nomenclature in substituting
a new name without adequate reason, which was not given in his second work
published in 1852.

B. Internodes uniserial at the proximal end. Joints composed of a single
chitinous tube (fig. 12).

I regard this condition as secondary, and as due to the suppression of the
proximal segments of the zocecia F and G. A similar suppression is found
in Emma.

7. TRICELLARIA ACULEATA (J? Orb.). (Fig. 12.)

Bicellaria aculeata, D’Orbigny, 1839-1846, Voy. Amér. Mérid. y. 4, p. 8, pl. 2.
figs. 1-4 (Tricellarta aculeata on the plate). Falkland Is.
Menipea aculeata, Busk, 1884, p. 20, pl. 4. figs. 2,2a@. Falkland Is., Patagonia,
5-175 fathoms*.
5 7 Marcus, 1921, Vid. Medd. Dansk naturh. Foren. Ixxiii. p. 93.
Campbell Id. ‘

* Busk’s record of Stat. 303, 13825 fathoms, should probably have been Stat. 308,
175 fathoms (see under Amastigia benemunita, p. 334). j

356 SIR SIDNEY F. HARMER ON

Menipea fuegensis, Busk, 1852”, p. 21, pl. 19. figs. 1-8. Tierra del Fuego, Falkland!
Is.; 1879, Phil. Trans. vol. 168, p. 194, Kerguelen.
3 9 Jullien, 1888, Miss. Sci. Cap Horn, vi. p. 70, pl. 12. figs. 1, 2;
pl. vii. figs. 8-10. Tierra del Fuego.
Scrupocellaria fuegensis, Waters, 1904, ‘ Belgica’ Bryozoa, p.24. Straits of Magellan.
rs bifurcata, Kluge, 1914, p. 614, text-fig. 8. Kerguelen.

There is some uncertainty with regard to 7. aculeata and its synonyms.
In 1879 Busk expressed the opinion that Menipea fuegensis (Busk, 1852) is
asynonym of M. aculeata ; and | think this opinion is correct. The type-
specimen (54.11.15.262) of AM. fuegensis is from the Falkland Islands, and it
is of rather coarser habit than those described and figured in the ‘ Challenger’
Report ; while its scutum is linear instead of being branched. The scutum
is known to be variable, within the limits of a single species, or even of the
same colony, in other Scrupocellariide ; and the differences in question do.
not seem to be sufficient for the establishment of two species from material
from substantially the same locality. The fertile internodes figured by
Jullien as MZ. fuegensis and by Kluge as Serupocellaria bifurcata are very
long, and Waters has expressed the opinion that Jullien’s species should be
considered distinct. I am inclined to regard them as different forms of a
single species, as indicated in my synonymy. D’Orbigny’s figures are
probably not correct in all details, but some latitude must be allowed to
observations made so long ago. All the specimens I have examined in the
British Museum Collection, whether from the Falkland Islands, Patagonia,
or Kerguelen, agree in the important character of having the internode
composed of a single zocecium at its proximal end, associated with the
existence of a single chitinous tube, instead of two tubes, in each joint.

8. TRICELLARIA LONGISPINOSA (Yanagi § Okada).

Menipea longispinosa, Yanagi & Okada, 1918, Ann. Zool. Jap. ix. p. 408, pl. 6. fig. 3,
text-fig. 1. Japan, 312 fathoms and depth not stated.

5, Huma, Gray.
Emma, Gray, 1843, p. 293.
» Busk, 1852?, p. 27.
Menipea (pars), auctt.

Internodes very short, typically composed of two or three zocecia. Bifur-
cation of types 13, 14 (figs. 13, 14), the two branches given off symmetrically
or asymmetrically. Opesia reduced by an extensive development of crypto-
eyst, sloping considerably ina basal direction. Marginal avicularia on the
proximal side of the opesia, or opposite its proximal end. Scutum usually
present, sometimes wanting. Joints conspicuous, the internodes very narrow
proximally.

Gray gave no generic characters, which were later described by Busk.
Although the genus has been considered unnecessary by MacGillivray (1881,

OELLULARINE AND OTHER POLYZOA. 357

Prodr. Zool. Vict., Dec. vi. p. 32) and others, I think it may conveniently be
retained for the following group of southern species. As pointed out by
Busk, Hmma is nearly related to Tvrieellaria.

A. Internodes biserial at the proximal end. Joints composed of two
chitinous tubes (fig. 14).
1. EMMA CRYSTALLINA, Gray.
Emma crystallina, Gray, 1843, p. 293. New Zealand.
y a3 Busk, 1852°, pp. 28, 33, pl. 40. figs. 1-8. New Zealand, Bass Straits.
Menipea crystallina, MacGillivray, 1881, Prodr. Zool. Vict., Dec. vi. p. 31, pl. 58.
figs, 2 et 2b.
Waters, 1887, A. M.N.H. (5) xx. pp. 88, 265. New South
Wales, etc.
Levinsen, 1909, pp. 152, 183, pl. 2. figs. la, 1d.

” ”

2. HMMA TRICELLATA, Busk.
Emma tricellata, Busk, 1852°, pp. 28, 33, pl. 41. figs. 1,2. Bass Straits, New Zealand.
Menipea tricellata, MacGillivray, 1881, t. czt. p. 34, pl. 58. figs. 5-5 b.

3. Emma BusKi (Wyv. Thoms.) (fig. 14).

Menipea buskw, Wyville Thomson, 1858, Nat. Hist. Rey. vy. Proc. of Societies, p. 144,
pl. 12. fig. 1. Tanta, New Zealand.
9 MacGillivray, 1881, ¢. cit. pp. 35, 32, pl. 58. figs, 6-60.
buskt, Levinsen, 1909, pp. 59, 131-133, pl. 2. fies. 3a-8e.

”

4, HMMA cErvicornis, MacGill.
Emma cervicornis, MacGillivray, 1869, Trans. Proc. R. Soc. Vict. ix. p. 127.
Victoria.
Menipea cervicornis, MacGillivray, 1881, ¢. cit. pp. 34, 32, pl. 58. fies. 4-46. '
var., Waters, 1887, A. M. N. H. (5) xx. p. 88, pl. 4. fie. 1.
New South Wales.
Levinsen, 1909, pp. 59, 132, 133, pl. 2. figs. 4a, 4.

” ”
” 9)

B. Internodes uniserial at the proximal end. Joints composed of a single
chitinous tube (fig. 13).

5. Huma cyatuus (Wyv. Thoms.) (fig. 13).
Menipea cyathus, Wyville Thomson, 1858, ¢, cit. p. 148, pl. 15. figs. 10, 10a. Bass
Straits; Port Fairy (Victoria).
MacGiilliqrar 1881, ¢. cit. pp. 33, Ree pl. 58. figs. 8-3.
Levinsén, 1909, pp. 152, 138, pl. 2. figs. 2a, 26.

” 27

” Pr)

6. SCRUPOCELLARIA, Van Beneden.

Zoarium jointed, biserial, bifurcation of type 8 (fig. 8), the joints crossing
the zowcia CG and DEF; frequently traversing the opesia of the outer
zoccia, © and D, but never those of F and G. Proximal segments of F and
G in contact on the basal side of H, shorter than the corresponding parts of

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 26

358 SIR SIDNEY F. HARMER ON

Cand D. Rootlets not forming marginal bundles, some of them given off
by basal heterozocecia. Hach zocecium is typically provided with frontal and
marginal avicularia and with a basal vibraculum, the last at its proximal end,
the marginal avicularium at its distal end. Cavity of the vibraculum
divided by a calcareous septum into a larger chamber containing the muscles,
and a smaller chamber, from which a rootlet originates in the proximal
members of the colony at least. Seta of the vibraculum without lateral
branches.

This genus is readily recognizable, and as but little confusion exists with
regard to its limits, I do not think it necessary 1o discuss the species in detail.
The relations of the joints to the outer zowcia at the bifurcations give useful
assistance in characterizing species ; a point which will be brought out more
fully in my forthcoming Siboga Report. In some cases, as in fig. 8, the
joint traverses the opesia at its middle. In other cases it merely involves the
proximal end of the opesia ; and in others it passes entirely on the proximal
side of that opening.

Kluge (1914) has described 17 species which he refers to Scerupocellaria,
One of these, S. berthollet?i, Aud. (p. 616) is from Cape Verde, and is rightly
placed. The remainder are from Kerguelen (1 species) and Antarctic
localities (15 species). All of these may more properly, in my opinion, be
distributed among the genera Amastigia, Notoplites, Menipea, and Tricellaria,
and I think it has still to be proved that the genus occurs within the limits
of the Antarctic Circle. Scrupocellaria is a characteristic genus of Northern
latitudes, it is well represented in’ the Siboga Collection from near the
Hquator, and it is well known to occur in localities much further south, as for
instance off the south coast of Australia. Its possible relation to Amastigia
is indicated on p. 337, under A. kirkpatrichi.

Although Serupocellaria exhibits the tendency, shown in other members of
its Family, to suppress important structures like the scutum and the frontal
and marginal avicularia, all these structures are more commonly retained
than in certain other genera. The basal vibracula are very rarely wanting ;
and in most cases they occur regularly on every zowcium. The occurrence
of either one or two vibracula in the axil of the bifurcation is a useful
systematic character, as has been pointed out by Waters (1897, J. L. 8. xxvi,
p. 7). Levinsen (1909, p. 134) has stated that the vibraculum of Scrupo-
cellaria is not divided into two chambers. I do not understand this assertion,

which is at variance with my own results.

CELLULARINE AND OTHER POLYZOA. 359

EXPLANATION OF THE PLATES.

The drawings made with an A objective were reduced to 3, and those with a
C objective to % their original size.

PuatE 16.

Diagrams of bifurcation in Cellularine genera (Text, Sect. [V.). The branches are all seen
from the basal surface, and a uniform notation has been adopted for the zocecia, of which
A is the more proximally situated zocecium which gives rise to two distal successors, The
preparations from which the drawings were made were, in nearly all cases, Canada balsam

mounts.

Fig. 1—Type 1. Didymozoum triseriale (‘Siboga’).

Fig. 2.—Type 2. Stirpariella (undescribed species in the ‘ Siboga’ Collection),

Fie. 3.—Type 3. Bugula johnstone (‘Siboga’); e.p., connecting process.

Fig. 4.—Type 4. Bugula scaphordes (‘Siboga’); ,, 5 90

Fig. 5.—Type 5. Bugula dentata (‘Siboga’) ; ‘ He 3

Fig. 6.—Type 6. Huoplozoum cirratwn (‘Siboga’). The dotted lines indicate the joints.

Fig. 7.—Type 7. Kinetoskias (undescribed, ‘Siboga’ Collection).

Fig. 8—Type 8. Serupocellaria feroa: ( (sional? ). The joint traverses the opesia (dotted
lines) of the outer zooscia.

Fig. 9.—-Type 9. Tricellaria ternata (Norway). A rootlet occurs on the proximal segment

oD,

Fig. 10.—Type 10. Tricellaria occidentalis vay. dilatata (Japan) ; ., rootlet.

Fig: 11.—-Type 11. Zricellaria peachii (Durham coast).

Fig. 12.—Type 12. Tricellaria aculeata (‘ Challenger,’ Stat. “803,” but probably Stat. 308;
see text, p. 355).

Fig. 13.—Type 18. Emma cyathus (Bass Straits).

Fig. 14.—Type 14. Emma buskii (Victoria).

. 15.—Type 15. Notoplhites rostratus, n. sp. (‘Siboga’). A rootlet occurs on the distal

segment of K.

Prater 17.

Figs. 16-20. Diagrams of bifurcation, continued. The specimens from which figs. 21 and
23-27 were drawn had been cleaned with Kau de Javelle, involving the loss of the spines
and chitinous parts.

ig. 16.—Type 16. Cornucopina (from a specimen in the ‘ Siboga’ Collection) ; 7.p., rosette-

plates.

@. 17.—Type 17. Menipea crispa (Algoa Bay, 8. Africa). The joint traverses the opesia

(dotted lines) of the inner zocecia.

.18.—Type 18, Menipea trisertata (Simon’s Bay, 8. Africa, B.M. 87.12.9.99).
g. 19.—Type 19. Mentpea spicata (Victoria). A system of branches, showing a sympodial

form of colony, associated with the aon of one of the joints at each
bifureation (B.M. 97.5.1.460).

, 20.—Type 20. Amastigia kirkpatr zk (Devs MSS.), n. sp. (Marion Id., B.M. 87.12.9.97).
ig, 21.—Amastigia nuda (Victoria, B.M, 97.5.1.246). Frontal view. Obj. A.

26*

360 SIR SIDNEY F. HARMER ON

Fig. 22.—Menipea marionensis (Simon’s Bay, S. Africa, ‘Challenger, B.M. 87.12.9.99;
found with M. triseriata). Basal view, showing 4 internal avicularia (¢.av.) ;
l.av., lateral avicularium. Obj. A.
'.. 23, —Menipea vectifera, n. sp. (New Zealand, B.M. 99.5.1.630). Frontal view. Obj. A.
24.— Amastigia nuda (Tierra del Fuego, B.M. 54.11.15.87, type-specimen). Diagram
of basal surface; /.av., lateral avicularium; 1, 5, marginal zocecia; 3, median
zoceciim; 2, 4, submedian zocecia.
Fig. 25.—Amastigia nuda (Victoria, B.M. 97.5.1.246). Basal view of a 5-serial branch,
showing 4 basal avicularia and 2 ovicells on marginal zocecia. Obj. A.
Fig. 26.—Amastigia rudis (Victoria, B.M. 97.5.1.462). Frontal view, showing gigantic
frontal avicularia on the marginal zocecia. Obj. A.
Fig. 27.—Amastigia rudis. Basal view of the same branch, a seta having been inserted
from another specimen which was not cleaned with Eau de Jayelle. Obj. A.

PLATE 18.

All the drawings are from specimens which had been boiled with Kau de Javelle, after
which treatment the longitudinal rows can be separated by gentle pressure of a needle on
the basal surface. The drawings were made with a C objective and reduced to one-third
their original size,

Fig. 28.—Menipea roborata (Australia, B.M. 50,5.2.2), Lateral view of two zomcia, with
ovicells (ov.) and frontal avicularia (f.av.); 1.p., lateral rosette-plates; sp., base
of spine; @.sp., internal calcareous spines.

Fig. 29.—Menipea roborata (same slide). Lateral view of two zocecia without ovicells;
f.av., frontal avicularium ; 7.av., internal avicularium; ¢.r.p., terminal rosette-
plate.

. 30.—Menipea roborata (same slide), Frontal view; e., the part of the eryptocyst
which overlies the internal avicularium (?.av.), of which the proximal end (p.)
is seen partly through the frontal wall and partly through the lateral wall of the
zocecium ; ¢.7.p., terminal rosette-plate.

Fig. 81.—Menipea ligulata (Victoria, B.M. 97.5.1.454). Lateral view, showing two ovicells,
a frontal avicularium, and a strong calcareous process (¢.p.) projecting vertically
into the body-cavity at the proximal end of the opesia.

Fig. 32.-—Menipea spicata (Victoria, B.M. 83.10.15.46). Lateral view of two zocecia without
ovicells; f.av., frontal avicularium ; ¢.av., internal avicularium.

Fig. 88.—Menipea ligulata (Victoria, B.M. 97.5.1.454). Frontal view of a zocecium with
an ovicell and an internal ayicularium (7.qv.).

Fig. 84.— Menipea spicata (same slide as fig. 82).—Lateral view of two zocecia, each with

an ovicell produced into a conspicuous spike.

-, 35.—Menipea spicata (same slide). Basal view; two of the zowcia with internal
ayicularia (7.av.); 7, origin of marginal rootlet.

. 36.—Menipea vectifera, n. sp. (New Zealand, B.M. 99.5.1.630). Basal view of two
zocecia, one with an internal avicularium (7.av.).

Fig. 87.—Mentpea vectifera, n. sp. (same slide). Basal view of two zocecia, showing the
characteristic calcareous bars (¢.b.) projecting into the body-cayity from the
proximal cryptocyst.

Fig. 88.—Menipea vectifera, n. sp. (same slide). Basal view, showing the large basal
avicularium (d.av.) at the bifurcation ;: Z.av., lateral avicularium,

Fig. 89.—Menipea vectifera, n. sp. (same slide). Lateral view of two zoccia, showing an
ovicell, two frontal avicularia, and the characteristic internal calcareous bars.

Journ. Linn. Soc. Zoon. Vol. XXXV. Pl 17

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Journ. Luyn. Soc. Zoon, Vol. XXXV. P18

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Reid,Lith.Edint

CELLULARINE AND OTHER POLYZOA. 361

PuatE 19.

The originals of Figs. 40-42, 51 and 52 were treated with Hau de Javelle.
Scale of figures as in Pl. 18.

.40.—Menipea triseriata (Simon’s Bay, 8. Africa, ‘Challenger,’ B,M. 87.12.9.99).

Lateral view of two zocecia with ovicells and frontal avicularia.

. 41.—Menipea triseriata (same slide), Basal view, showing an internal avicularium

(<.av.), a lateral avicularium (/.av.), and the origin of a marginal rootlet (7.).

. 42.— Menipea triseriata, Frontal view of the same zocecia; f.av. frontal avicularium ;

2.av., internal avicularium; /.av., lateral avicularium, above which is seen the
origin of a rootlet.

. 43.—Menipea marionensis (Simon’s Bay, ‘ Challenger,’ B.M.87.12.9.99). Frontal view;

cer., cryptocyst; f.av., normal frontal avicularium; @.av., base of an internal
avicularium, showing the origin of its muscles from the frontal surface.

. 44.— Menipea marionensis (Cape of Good Hope, ‘Challenger,’ Stat. 142, B.M. 99.7.1.698).

Basal view, showing two internal avicularia (¢.av.), with their mandibles.

. 45.—Menipea marionensis (same slide as fig. 43), An old zocecium, showing an

excessive development of the internal calcareous spines; cr., cryptocyst;
p., proximal end of the next distal zocecium, connected with the edge of the
opesia (@.) by a calcareous film; fiav., frontal avicularium; /.av., lateral
avicularium.

». 46.—Amastigia kirkpatricht (Ley., MSS.), n. sp. (Marion Id., ‘Challenger,’ B.M.

87.12.9.97). Young zocecium, with 5 oral spines.

47.— Amastigia kirkpatrickt (same slide). Basal view, showing vibracula, the upper-

most axillary.

o, 48.—Amastigia kirkpatricki (same slide). Frontal view, showing a gigantic frontal

avicularium and two ovicells.

o, 49.— Amastigia rudis (Victoria, B.M. 97.5,1.462). Part of a zocecium, with scutum, in

frontal view.

, 50.—Amastigia nuda (Victoria, B.M. 97.5,1.246). A similar preparation, frontal view.
g. 51.—Amastigia nuda (same slide as fig. 50). Lateral view of two zocecia, with

frontal avicularia (paired).

. 52.—Amastigia rudis (same slide as fig. 49). A similar view of two zocecia with

frontal avicularia.

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MR. J. N. HALBERT: NOTES ON ACARI. 363

Notes on Acari, with Descriptions of New Species. By J. N. HALperr,
M.R.I.A. (Communicated by Dr. W. T. Catmay, F.R.S., Sec.L.8.)

(PiatsEs 20-22.)
[Read Ist March, 1923. ]

WitH a few exceptions the Acari recorded in the following paper were
collected in Ireland since the publication of the Clare Island Survey reports
seven years ago. ‘The list of Acari there recorded (23) was practically a
complete one in so far as the Irish terrestrial forms were then known. A few
years later an account was given of the species found living in the zones of
the sea-shore affected by the tides (24). At the present time, collections are
being made with the intention of preparing a more complete list of our
Acarid fauna, but much remains to be done in this respect. Meanwhile it
is desirable to record a certain number of new and interesting forms as a
contribution to our knowledge of the Acari in these islands.

The literature dealing with the Acari is very considerable ; large numbers
of species have been described by Huropean workers in recent years, and, as
very many of these species are unfigured, the difficulties of identification are
greatly increased. In the present paper short descriptions only are given,
and such figures as seem necessary in a group where the details of structure
are so varied and often of so comparative acharacter. Inall, some sixty-eight
species are recorded ; sixteen of these, and also a few varieties, belonging
to the families Gamaside, Oribatide and ‘lrombidiide, are apparently
undescribed. Including these new forms, about forty species are now
recorded for the first time from the Britannic area. The types of two of the
new species, in Lasioseius, were collected by Mr. A. D. Michael on the coast
of Cornwall some years ago.

I am indebted to a few friends for assistance in the collecting of specimens,
more particularly to my colleague, Mr. A. W. Stelfox ; the discovery of some
of the more interesting forms is due to him. I have also to thank
Dr. A. Berlese for kind help in the identification of some of the species :
others are recorded on my own responsibility. The type-specimens of all
the new species are in the collection of the Irish National Museum.

Gamasus (HUGAMASUS) CRASSITARSIS, sp. nov. (PI. 20. figs. 1 a-c.)

Male. A large robust species measuring 1600 in length; general
characters as in the subgenus Hugamasus, to which the species belongs.
Shoulders well marked; hairs long, those on the second dorsal shield
numerous and somewhat adpressed. Sternal region of ventral plate with
faint seale-like markings on its anterior half, concave at the centre of the

364 MR. J. N. HALBERT : NOTES ON ACARI,

front margin. Genital aperture strongly chitinized, posterior extremities
pointed, reaching to middle of second cox. Hpistome with a long central
spine, and the ends of the lateral spines slope slightly inwards. Maxillary
plate transverse, with four long hairs. Chelicerze (fig. 1 a) somewhat like
those of G'. oudemansi, Berl.; fixed chela armed with two stout teeth, the
extremities of which are truncated and minutely dentate, apex of segment
bilobed; free chela with one strong tooth slightly recurved.

First, third, and fourth legs long and slender ; third segment of last pair
with a sharply-pointed ventral spine. Second legs (fig. 16) very stout ;
femoral calcar of moderate size, curved posteriorly, bluntly pointed ; cap of
axillary process flattened and produced towards apex of calcar as in G. oude-
mansi, Berl. ; on the outer side of the femur there is a marked chitinous
prominence (fig. 1c) with a terminal hair; genual process well developed,
cap concave ; process of fifth segment not prominent, its extremity does not
project beyond the ventral outline of the segment. ‘Tarsi strongly swollen
at the base and again at the apex; a stout conical tooth stands near the
middle of the inner surface.

This fine species is evidently allied to Gamasus magnus, Kramer, and to
G. oudemansi, Berl. It is however larger, and differs from both in the
presence of a chitinous projection on the outer side of the second femora ;
the tarsi and chelicerze are also different. It also resembles the species
redescribed as ‘ Hugamasus loricatus, Wankel” by Oudemans (88, p. 114),
especially in the form of the tarsi, but differs from this species in the
armature of the second legs and in other particulars.

Locality. Two males found under a sunken stone at high-tide mark in
company with the local Chelifer dubius (Camb.) at Mount Garrett Wood, near
New Ross, Co. Wexford, by Mr. R. J. Phillips, 12th March, 1922.

GAMASELLUS (PROTOLALAPS) GRANULATUS, sp. nov. (Pl. 20. figs. 2 a, 6.)

A very distinct species belonging to the subgenus Protolwlaps, Berlese
(13, p. 187). It is remarkable for the comparatively large size of the ventral
plates, more especially of those enclosing the peritreme. Verified by
Dr. Berlese. :

Female. Colour yellowish brown, shape pyriform. Cuticle very distinctly
punctate-striate. The two dorsal plates are of equal breadth, leaving a .
rather wide uncovered lateral margin ; the second plate is undulate in front
and almost reaches end margin of body. These plates are minutely punc-
tured, and there are faint traces of reticulate markings. Hair armature short
and stout. Jugular plates absent. Sternum with a bow-shaped front
margin, rather long, reaching middle of third coxze ; end margin truncate.
Metasternal plates with a stout hair on outer margins. Genital plate sub-
quadrate (length 75), anterior half lying between last pair of coxe.
Ventro-anal plate large, twice as broad as long (length 100 p, breadth 200 ~);

WITH DESCRIPTIONS OF NEW SPECIES. 365

front margin flattened, end bluntly pointed and reaching margin of body.
All of these ventral plates are minutely punctured. Peritreme strongly
sinuate, lying close to the legs on the inner margin of a large plate which
projects in a broad tongue-shaped process beyond, but does not enclose, the
last pair of legs. Rod-like endopodial plates lie between the third and
fourth coxe. j

Epistome with a long central spine fringed with minute hairs. Maxillary
plate quadrate ; the small curved lobes are placed on a raised central part
which projects beyond the front margin of the plate. Chelicerze very minute.
Palps normal. Legs robust, of moderate length, clothed with short hairs ;
ambulacra on all pairs. Length 363 yu, breadth 275 pw.

Locality. Apparently a rare species ; found on two occasions, in January
and November, under stones in the Tolka Valley, near Dublin.

GAMASELLUS (PROTOLHLAPS) MUCRONATUS (G. et I. Can.).

1881. Gamasus mucronatus, G. et R. Canestrini, 19, p. 1081.—1882. G. et R. Can., 20,
p: 52.—1885. G. Can., 18, p. 78.—1887. Cyrtolelaps mucronatus, Berlese, 1, Fase. xliv.
n. 5.—1921. Berlese, 15, p. 81.

This species may be recognized by its large size, measuring 1200 w in
length, and the large strongly-chitinized dorsal plates. In addition to a
number of minute hairs, both plates carry two pairs of strong spines feathered
at their extremities ; one pair at the end of the second plate project well
beyond the posterior margin of the body. ‘The peritreme lies close to the
coxee on the inner margin of a large peritrematic plate. The ventro-anal
plate is large, V-shaped, emarginate in front, its extremities lying close to
the genital plate and the posterior margin of the body.

Locality. Females found in rotting potatoes at Drumcondra, near Dublin,
in April. The identification has been verified by Dr. Berlese, who refers
(in lit.) the species to his subgenus Protolelaps. The male has not been
described.

GAMASELLUS (PROTOLHZLAPS) SUBNUDUS ?, Berl. (Pl. 20. fig. 3.)
1918. Berlese, 13, p. 138.

The following is a brief description of a Gamasellus, as well as of its
larva and nymph, which probably belongs to the present species, although
the identification is given with some reserve.

Adult female pear-shaped. Hpistome with a long sinuate spine, minutely
spiculate at the base. Dorsal shields of moderate size and uniform breadth,
distinctly punctate, the punctures lying in transverse rows; margin of
second shield broken up into small folds, and there is a wedge-shaped fissure
in the middle of the front margin ; hairs sparse and weak. Wpidermis
strongly granulate. Sternum of moderate size (length 132), hinder
corners slightly rounded. Endopodial plates very strongly developed and

366 MR. J. N. HALBERT : NOTES ON ACARI,

produced in sharply-pointed processes between the coxe. Peritreme undu-
late, and protected by a narrow plate which reaches to the end of the fourth
coxee. Ventro-anal shield very small, rotund, broader than long (breadth
165), placed on the posterior margin of the body. Legs of moderate
length, second pair slightly more robust than the two hinder pairs. Length
742 w, breadth at shoulders 430 pw, and in posterior third of body 512 p.

Larva. Shape much as in adult; close to the end it slopes rapidly to a
truncated central part of the posterior margin, at each corner of which
is a curved hair; immediately behind these on each side are stouter hairs
with slightly clubbed and spiculate extremities ; the dorsal hairs are weak.
Dorsal shields undeveloped, and the epidermis at end of body is distinctly
areolated. Legs and palps very stout, carrying numerous short hairs; ambu-
lacra consisting of a single pair of rather narrow leaf-like upper lobes, and
two long, lanceolate, lower processes. Length 340, breadth 198 py. (P!. 20.
fig. 3.)

Nymph. Side margins more parallel than in the adult; end margin
truncated, with a small central part projecting below. Dorsal plates large,
granular. Side margins of body with short hairs. Hpistome, palps, and
legs much asin the adult. Length 570 w, breadth 320 p.

Locality. Adult female, larva, and nymph found in decaying turnips,
North Dublin, in April 1921.

MacrocHELES VAGABUNDUS (erl.).

1889. Holostaspis vagabundus, Berlese, 1, Fase. li. n, 8.—1902. Oudemans, 37, pp. 11,
43.—1918. Berlese, 3, p. 172.

Berlese places this species in the subgenus Maerocheles (sensu stricto, 18)
of which the type is Acarus marginatus, Hermann. In the same paper he
records varieties of WM. vagabundus from South America, South Africa, and
Australia, so that the species is of unusually wide range. It may be known
. by the beautiful sculpturing of the dorsum, which, as well as being distinctly
punctured, is ornamented with reticulations and branched chitinous markings
on the epidermis. The strong, curved hairs on the margin of the dorsal plate
bear numerous very short secondary hairs.

Locality. The only specimens (¢ ?, length 930 ~) found in Ireland were
under moist pieces of wood lying on the sea-bank at Malahide, Co. Dublin,
in September.

DENDROLELAPS CORNUTUS (Kramer).

1886. Setus cornutus, Kramer, 31, p. 257.—Dendrolelaps bicornis, Hull, 25, p. 57.

This species is well described and figured by Kramer (81), and is easily
recognized by the presence of two long chitinous horns near the end of the
dorsum in the male. In the female (length about 495 ~) the body is rather
long and narrow. The sternum fills the entire space between the very large
second cox, and is gradually widened to its hinder corners. The genital

WITH DESCRIPTIONS OF NEW SPECIES. 367

and ventro-anal shields are remarkably long and narrow, of almost equal
breadth, and the latter reaches to within a short distance of the end margin
of the body.

Locality. Both sexes occurred under the moist bark of cut pine-logs lying
on the Royal Canal bank, close to Dublin, in October 1918. The logs had
been brought from an inland locality.

Myonyssus pecumMANus, Tirabosche.

A few adult females were found on a Field Mouse (Mus sylvestris) caught in
the Rocky Valley, Co. Wicklow, by Dr. R. F. Scharff. On the same host
were adults of Lwlaps stabulasis, C. L. Koch, and L. agilis, C. L. Koch. It
has been found on the common Brown Rat in Hngland (Hull, 25), and
Waterston records its occurrence on the House Mouse in the Shetland Islands.

ANTENNOPHORUS UHLMANNI, Haller.

A single specimen was found in a nest of the ant, Lasius miztus, Nyl., near
Graiguenamanagh, in Co. Kilkenny, by Mr. A. W. Stelfox. This is the only
Antennophorus hitherto found in Ireland ; all the described species, four
in number, have been recorded from Hngland.

Lauars (IPHIS) ACULEIFER, Can.
1983. Lelaps acuierfer, Canestrini, 17, p. 6.—1885. Hypoaspis aculeifer, Canestrini, 18,
p. 84.1892. Lelaps aculeifer, Berlese, 1, Fasc. Ixviii. n. 10.
Evidently a fairly common species. I have found the female in numbers
under bark of decayed trees at Tallaght in April and Greystones in July,
under pieces of damp wood in the Tolka Valley in January, and in decayed

potatoes at Drumcondra in April. All these localities are in the vicinity of
Dublin.

LA&LAPS LATISTERNUS, sp. nov. (PI. 20. fig. 4.)

Female. Shape broadly ovate, shoulders not prominent. Dorsal plate very
large, surface with scale-like markings ; hairs short, becoming longer towards
end margin, where there are two comparatively long hairs. Sternum very
wide and strongly produced between the coxze at its anterior and posterior
corners, end margins slightly undulate, hairs rather long. The genital plate
is also rather large (length about 165), marked with lines as in figure.
Peritreme comparatively straight, not fused with the pedal plates ; the latter
are well developed and enclose the fourth cox; small obliquely-placed
inguinal plates lie close to them. Ventro-anal plate small (length 220,
breadth 176 ~), subovate, flattened at end and reaching margin of body.

Epistome in the form of a long, sword-shaped, central spine and two strong
lateral teeth. Maxillary plate rather narrow (77 4). Palps normal. Legs
long and robust, first pair of very uniform thickness (length about 506 p) ;
the fourth pair (length 550 ~) have slightly curved femora, and the tarsi are
elongate ; there are traces of a division of the terminal part of the tarsus close

368 MR. J. N. HALBERT: NOTES ON ACARI,

to the middle. Hair armature short, that of first pair stronger than the
others. Ambulacra normal. Length 530 p.
Locality. Found under stones in the Lucan Demesne, Co. Dublin, February.

L&LAPS SIMPLEX, sp. nov. (PI. 20. fig. 5.)

Female. Long oval, shoulders not well marked. Dorsal plate straight-
sided, reaching end of body, with three double rows of weak hairs, two on the
posterior margin much longer. Hpidermis distinctly striated. Sternum of
rather uniform breadth, reaching to middle of third coxee ; front margin
undulate, with slight lateral extensions ; posterior corners pointed ; end margin
with a small notch at the middle; the three pairs of hairs are placed close to
the margin of the shield. Genital plate exceptionally long (length 110 p)
and narrow, being more than twice as long as broad, reaching well beyond
the last pair of coxze, only slightly widened at the extremity, which is some-
what rounded. Ventro-anal plate small (length 90 4, breadth 95 yu), widely
removed from genital plate and placed on the end margin of the body, evenly
rounded in front and truncated behind. A pair of small inguinal plates lie
near the side margins. The inner pedal plates lie between the last pair of
coxze, which are partly enclosed by a chitinous extension of the peritreme.
Chelicere small, though of robust structure. Legs and palps stout and of
moderate length ; inner side of second palp segment with one, and third
segment with two stout spines. Length 450 u, breadth 260 p.

Locality. Vaken under rotten wood at Glendalough, Co. Wicklow, in
September.

LL4LAPS FIMBRIATUS, sp. nov. (PI. 20. fig. 6.)

Female. Long oval. Dorsal plate rather narrow; hairs long, increasing a
little in length at end of plate. Sternum rather short ; front margin bow-
shaped, extending between the coxze; sides deeply emarginate. Metasternal
hairs present, but the plates are indistinct. Genital plate large (length about
150 w along middle line), elongate, constricted at the middle; front margin
produced and elaborately fimbriate, hind margin straight ; anterior part of
plate marked with longitudinal lines. Ventro-anal plate large (length 175 p,
breadth 150 ~), hatchet-shaped, the bluntly-pointed end directed towards the
posterior margin of body; surface with transverse lines; front margin much
wider than genital plate. ‘The outline of the plate of an earlier nymphal form
is indicated in the drawing. Peritreme with a slight regular curve; there is
no protecting plate, anda weakly-chitinized post-stigmatic process encloses the
last pair of coxe. Maxillary plate quadrate, corniculi narrow and placed
close together. Legs long and slender (length of first pair 450 pu, of fourth
pair 550). Ambulacra rather long and narrow, but otherwise of normal
structure. Length 528 w, breadth 320 p.

Locality. Found on sprouting potatoes at North Dublin in March of the
present year.

ite)

WITH DESCRIPTIONS OF NEW SPECIES. 36

CopripHts (ALLIPHIS) HALLERI (G4. et R. Can.).

1881. Gamasus halleri, G. et R. Canestrini, 19, p. 1077.—1882. Lelaps halleri, G. Can-
estrini, 20, p. 57.—1885~99. Iphis halleri, G. Canestrini, 18, p. 93.—1892. Iphis
halleri, Berlese, 1, Fasc. lxvii. n. 6.

Localities. Males and females occurred commonly in decayed roots of
Henbane at Clontarf, near Dublin, in August 1918, and under bark of cut
pine-logs on the Royal Canal bank in October 1918. Canestrini records
it as occurring in decayed potatoes and amongst rotting leaves in
December.

Lastoserus (LAsios.) GRACILIS, sp. nov. (PI. 20. figs. 7 a-c.)

Female. Shape ovate, shoulders not marked. Dorsal plate large, almost
completely covering body, surface with polygonal markings. Hairs long,
formed of a strong rib and blade, curved and serrulate, those on end margin
exceptionally so (length 65). Frontal bristles shorter and strongly pec-
tinate. Sternum of moderate size, corners not produced between the coxee ;
hinder margin straight, with rounded corners about on a level with end of
second coxee. Metasterna small. Genital shield broad posteriorly and rather
strongly narrowed towards the front ; lying behind it are two thin chitinous
bars and two pairs of hairs. Ventro-anal plate of medium size, a little
broader than long (length 145 w, breadth 150 yw), flattened in front and
regularly rounded behind. Anal aperture placed a little in advance af the
centre of the plate. Inguinal plates linear, placed near side margins.
Maxillary plate transverse, evenly rounded at base; corniculi with their
apices trifid, directed inwards, and there is a strong hair at their bases.
Chelicerze small; fixed limb swollen at the base, where there are four sharply-
pointed teeth placed close together ; free chela with an oblique chitinous
blade and traces of two or three teeth just before the apex. Palps normal.
Legs of moderate length, rather slender, furnished with short, curved hairs,
some of which are minutely spiculate; length of first leg about 352 pw,
of fourth leg 363m, not including ambulacra. Length 474 wu, breadth
310 pw.

Dr. Oudemans has partially described (37, p. 17) the protonymph, deuto-
nymph, and adult of an Acarid which he calls Sevwus plumosus, though it
seems doubtful if all of the forms are referable to the same species. L. gracilis
is allied to “ Setulus plumosus,” but the legs are obviously longer and more
slender, and the hairs of the dorsum are much longer and not so strongly
blade-shaped as they are in the Dutch species, which was found on a bat
( Vespertilio dasyeneme) and on a Squirrel (Sciwrus vulgaris).

Locality. Found on sprouting potatoes at the Albert Model Farm, near
Dublin, by Mr. J. G. Rhynehart, early in March.

370 MR, J. N. HALBERT : NOTES ON ACARI,

Lastosetus (Lastos.) omETES, Oudem. (PI. 20. figs. 8 a-c.)

A Lasioseius found under the bark of trees in Ireland seems referable to
this species. I have to thank Dr. Oudemans for kindly lending me his type-
specimen for comparison.

Form as in Z. muricatus, C. L. Koch (Berlese,1, Fasc.xli. n.6). Sternum
with a wide and deep cleft in the posterior margin extending to about the
middle of the plate; the sides of the cleft are usually irregular in outline.
Metapodial plates vestigial. Genital plate of the usual shape, punctured ;
end margin rather strongly chitinized ; immediately behind it are four small
linear plates arranged in a transverse row. Ventro-anal plate very large,
with distinct transverse markings; side margins concave towards apex ; the
end margin is somewhat truncated, and reaches the posterior margin of the
body. The peritreme is normal, and its plate is continuous with the inner
pedal plate, the extremity of which reaches the third coxee. Tritosternum with
rather short processes ; its base lies in the central depression of the bow-
shaped jugular bar. Fixed chela with a row of about ten small teeth, and
there are three teeth on the free chela (fig. 8c). Legs, palps, and capitulum
much as in L. muricatus, C. L. Koch. Length 580 w; breadth 319 pw.

Localities. Females found under bark of decayed logs at Tallaght, and in
the Carton Demesne, Co. Kildare, in May.—(See note on p. 392.)

Lastosrerus (Lasios.) LALAPTOIDES (Berl.).

1887. Epicrius lelaptoides, Berlese, 1, Fasc. xl. n. 10.—1916. Berlese, 10, p. 33.—
1921. Berlese, 15, p. 82.
Locality. Female found under damp wood lying on the ground in the
Tolka Valley, near Finglas, Co. Dublin, in May. Not previously recorded
from Britain.

LasiosE1us (HPIsErus) @LABER, Berl., var. CURTIPES, nov. (PI. 20. fig. 9.)

General structure as in the type-form (Berlese, 1, Fase. xxx. n. 9); differs
in the much shorter first pair of legs; in his description of these, Berlese
says : ‘antici exiliores, corpore fere duplo longiores,” and his accompanying
figure shows the first legs much longer than the body. In the present variety
the first legs are only slightly longer than the body, and the remaining pairs
are somewhat stouter. The ambulacra are of the modified, bristle-like type.
The genital plate is longer and the ventro-anal plate is broader. Triigardh has
described a variety (minor, Triig., 46, p. 432) found in the Sarekgebirge ; from
this the present variety may be known by the more strongly-produced meta-
podia, and the form of the ventral plates is different.

Female. Dorsum with scale-like markings, which are distinct only towards
the margins. Hair armature weak, stronger towards end margin, on which

WITH DESCRIPTIONS OF NEW SPECIES. 371

are a number of very short spines. Sternum rather large (length 187 p,
breadth 165). Jugularia absent. Metasternal plates extremely small.
Genital plate longer than broad, with a thick hinder margin. Metapodial
plate much produced, and ending in a bluntly-pointed extremity reaching
beyond the front margin of the ventro-anal plate. The latter is large and
much broader than long (length 120 wu, breadth 154 w), front margin trun-
cated, posterior margin evenly rounded. Inguinal plates inconspicuous.
Length 429 w, breadth 252 p.

Locality. Two specimens found amongst wet moss colleeted on Lambay
Island, Co. Dublin, in July.

LasiosEtus (HPISEIUS) SPHAGNI, sp. noy. (PI. 20. fig. 10.)

This species bears a superficial resemblance, especially in the form of the
ventral plates, to L. lelaptoides, Berl., but, apart from other differences, it
may be easily known by the elongate tarsi and modified ambulacra, which are
of the Hpiseius type, with long terminal bristles, while in L. lelaptoides
these parts are of the ordinary L. muricatus type.

Female. Colour light brown, shape broadly ovate. Dorsal plate very
large, surface not reticulate, but with imdistinct areolations on its anterior
part. Hair armature weak, a row on side margins stronger, and there are
three pairs of deflexed hairs on the end margins. Jugular plates absent.
Sternum normal (length 110). Metasternalia distinct. Genital plate
broad, front margin indistinctly serrulate ; lying behind this plate are four
small linear plates. Ventro-anal plate large, broader than long (length
about 200 uw, breadth 264 w), front margin flattened. Sides produced in a
rounded prominence in front, emarginate towards apex, which is somewhat
truneated and minutely punctured. Canal of peritreme broad, with the
usual post-stigmatic continuation, and protected by a narrow outer plate.
Metapodia strongly chitinized, ending in bluntly-pointed extremities, much as
in L.2talicus, Berl. Hpistome tricuspid. Maxillary plate quadrate; corniculi
small and slightly sinuate, and three pairs of long hairs stand near their bases.
Palps normal. Legs long and robust; length of first and fourth pairs about
equal (562 w); ambulacra with three terminal bristles and a pair of small
membranous lobes.

Locality. wo specimens found in sphagnum collected in the Black Valley,
Howth, Co. Dublin, in October.

LasiosEius (Heiserus) iranicus, Berl., L. (H.) MICHAELI, sp. nov.,
L. &.) tenvuipxs, Halbt., and L. (H.) masor, sp. nov.

From a study of the species belonging to the “taliews” group of the sub-

genus Hpiseius, Hull, which includes those species of Lasioseius in which the

tarsi are elongate and the ambulacra armed with bristles instead of the usual

372 MR. J. N. HALBERT : NOTES ON ACARI,

lobes (Hull, 25 ; Halbert, 23, 24), I believe there are at least four closely-
allied species in these countries. The male and female of three of these are
now known to me, and although the general structure is much alike, the
males possess excellent characters in the form of the mandibular calcar.
These three species are L. (/.) italicus, Berl., L. (E.) michael, sp. nov.,
and L. (H.) major, sp. nov. The male of L. (£.) tenuipes, Halbt., has not
been discovered, and until this is found I-must rely on the characters of the
single specimen described (23, p. 78). The females in the case of two of the
species, i.e. L. (£.) ttalicus and L. (H.) michael, are difficult to separate ;
indeed, the one description would suit both species almost equally well.
Unfortunately the males, as in many Acarid genera, are decidedly rare; as
an instance, I may mention that out of a gathering from one locality of about
one hundred examples of “‘italicus” aggregate three males were found, and
this only by close searching of moss brought home from a suitable locality.
The species frequent wet moss, liverworts, &., and may be found amongst
submerged mosses growing on stones in mountain-streams, in company with
mites of the genera Calonyx, Panisus, and Aturus.

The female of L. (#.) italicus, Berl., is described in 4, p. 234, and is
excellently figured by Dr. Berlese in a later paper (“ Redia,” vi. pl. 19. fig. 35),
while the male is briefly described in 10, p. 34. In view of the great
similarity of the females of these species, it will probably be sufficient to give
a short description of L. (1/.) italicus, and then briefly tabulate the chief
characters of the four species. Figures of the male mandibular calcars of
three species and of the female of L. (/.) major, sp. nov., are given in the
present paper.

L. (B.) italicus, Berl. Female. Shape pyriform. Dorsal shield large,
reaching end of body: sides often slightly rounded, surface reticulate. Hair
armature weak on centre of dorsum, much stronger and somewhat adpressed
at the margins, where they are arranged in a double row. There is a raised
tuberele near the posterior margin. Epistome of the tricuspid Episeius type.
Sternum rather large and wide; both the front and hinder margins are
slightly concave, and the latter reaches to middle of the third cox. Genital
plate hatchet-shaped ; hinder margin straight, standing well beyond the
fourth coxee. Ventro-anal plate large, much broader than long (length 210 pu,
breadth 300 ,), flattened in front; side margins concave near centre, surface
with irregular lozenge-shaped markings. Between the last two plates are
three pairs of very minute plates arranged in two rows. Peritreme broad
and strongly sinuate. ‘he united plates of the peritreme and metapodia
extend beyond the last pair of coxze in an obtusely-pointed triangular form.
Legs very long, with attenuated tarsi, and the ambulacra have bristle-like
lateral lobes and a lanceolate upper lobe. Length about 640 mw, breadth
420 p.

WITH DESCRIPTIONS OF NEW SPECIES 373

I. A raised tubercle present near end of dorsum in both sexes.*

A. The extremities of the metapodial plates slope downwards in an oblique line
towards the body margins.

a. Male: Length 495 , breadth 330. Calcar of mandible (Pl. 20. fig. 11) long,
consisting of a thickened rib and narrow blade, flagelliform, curved strongly
downwards under the maxillary plate, extremity sinuate. Female: Length
640 y, breadth 420 1; body usually broader and more rounded than in the
following species. Length of first leg 716 p, of fourth leg 768». (Localities:
Streams at Glencullen, Glencree, Kilmashogue, in the Dublin Mountains;
Reservoir at Greystones, July. Lambay Island, July, &c.)

Lastoseius (Episeius) ttalicus, Berl.

b. Male: Length 495 p, breadth 330. Calear of mandible (fig. 12) long (length
about 210 1, not including base of mandible), almost straight except at the
apex, where it is deflexed. Length of first leg 4701, length of fourth leg
614. Female: Length of type 640, breadth 390; length of first leg
692,.; leneth of fourth lee 768. (Locality: Mill Bay, Land’s End,
England ; taken by My. A. D. Michael in November 1892.)

; Lasioseius (Episeius) nuchaeli, sp. nov.

B. Extremities of metapodial plates truncated, lying in a straight line across the
venter. Male unknown. Female described in 23, p. 78. Length about
740; breadth 486. Length of first leg 742, of fourth leg 820 p,
including ambulacra. (Locality: Mountain-stream near Glencree, May.)
Lasiosetus (Episeius) tenwpes, Halbt.
II. No tubercle at end of dorsum.

Male: Length 520, breadth 374. Calear of mandible (fig. 130) short (length
about 70 4, not including base of mandible), straight, or slightly curved; apex
deflexed. Female (fig. 13), length 7404, breadth 486. Body broadly pyri-
for, posterior margin somewhat flattened. Ventro-anal plate large (length
220 p, breadth 396 2), distinctly reticulate. Length of first leg 820, of fourth
lee 890 ». (Localities: Streams at Kilmashogue (¢ & 2, December) and Glen-
cullen (2 2, October), Mill Bay, Land’s End, England (g & 2, November,
coll. A. D, Michael).) Lasiosevus (Episetus) major, sp. nov.

LAStosmius (PLATYSEIUS) SUBGLABER (Oudem.). (Pl. 20. fig. 14.)
1906. Hypoaspis subglabra, Oudemaus, 36, p. 88.

A very distinct species, evidently referable to the subgenus Platyseius,
Berlese (10, p.42). Characteristic features are the broad pyriform shape and
the very long hairs fringing the side margins of the dorsal shield. These
hairs are arranged in an irregular double row and are incurved ; on the end
margin are two shorter, straight hairs. Length 560-600 w, breadth 430 p.
Length of first leg 666 w, of fourth leg 717 p,

* The male of Z. (£.) tenuipes is unknown. In a recent paper on Swiss Acari,
Dr. Schweizer described and figured what he considers is the male of Z. tenwpes, Halbt.
(41, p. 43). I was struck by the similarity of the male calcar there figured with that of
LT. (E.) major now described. Dr. Schweizer very kindly sent me some mounted specimens
(Q @ ) for examination (locality, Quellen am Kellersee, 4. iv. 1819) and they are undoubtedly
referable to this species. The male calear figured (Pl. 21, fig. 18) agrees well with that of
my type of Z. (£.) major.

LINN. JOURN. —ZOOLOGY, VOL. XXXY. 27

374 MR. J. N. HALBERT: NOTES ON ACARI,

The male of this species has not been described. The general structure
resembles the males of Mpiseius. Dorsum as in the female. Free chela
(fig. 14) armed with a long process resembling that of L. (£.) italicus, Berl.,
but shorter and less curved, each chela with one strong tooth, in front of the
tooth on the fixed chela is a row of very small teeth. Length of male 484 y,
breadth 352 yp.

On account of the apparent difference in the length of the dorsal hairs in
the Irish specimens when compared with the published figures (36, Taf. 6.
fig. 45), I sent drawings to Dr. Oudemans, who informs me that they un-
doubtedly represent L. (P.) subglaber. The dorsum of this species is
frequently coated with fragments of débris which are entangled between the
long inwardly-curved marginal hairs, reminding one of what occurs in the
Oribatid genus Dameus.

Localities. In sphagnum pools at Lough Atorick, Co. Clare, June, and
also at Lough Dan, August. Both sexes occurred amongst wet moss
collected on the bank of a stream at Glencullen in the Dublin Mountains,
in October, and at Drogheda in August.

Lastosetus (Lerosgius) minutus (Halbt.).

1915. Sevwlus minutus, Halbert, 23, p. 76.—1918. Zerconopsis minutus, Hull, 25, p. 66.
—1920. Lastoseius minutus, Berlese, 14, p. 171.

Hull has placed the present species in a new genus Zerconopsis, of which
the type is Kramer’s ‘ Gamasus remiger.’ Berlese, however, had already
referred this species to his subgenus Zercoseius (10, p. 33) with S. spathu-
liger, Leon., as the type. On re-examining the Irish specimens of L. minutus,
it seems to me they fall readily into Dr. Berlese’s subgenus Leioseius, briefly
diagnosed by him as follows :—‘“ Ex gen. Lasioseius. Pedes breves et robusti.
Truncus elongatus, lateralibus subparallelus. Typus L. L. minuseulus, Berl.”
An estuarine species (L. salinus, Halbt.) of this subgenus occurs on the
Dublin coast (24, p. 125).

Localities. Amongst sphagnum collected on Croaghmore Mountain, Clare
Island, Co. Mayo, and under rotten wood lying on the ground in the Carton
Demesne, Co. Kildare, May. The male is unknown.

Serius toeatus, C. L. Koch.

Until recently (1916) acarologists referred species of very different facies
to Koch’s genus Seius. Many new genera and subgenera have now been
established by Dr. Berlese, so that a great advance has been made. As
S. togatus is the first species of the genus described by Koch it remains as
the type. It is also a very isolated form with but few congeners; three have
been described from Europe (see Berlese, 11, p. 150).

Locality. A single specimen found by Mr. Norman Stephens under the
moist bark of a pine-stump at the entrance to The Devil’s Glen, Co. Wicklow,

WITH DESCRIPTIONS OF NEW SPECIES. 375

in May. The species has been recorded from Scandinavia (Trigardh),
Germany (Koch), Switzerland (Schweizer), and the North of Hngland (Hull).

ZERCON TRAGARDHI, sp. nov. (PI. 21. figs. 15a, 6.)

Female. Colour pale brown. Shape broad ovate. Side margins
strongly serrated, and the end margin is crenulate much asin Z. trigonus, Berl.
First dorsal shield sculptured with scale-like markings, second shield with
areolations and four crescentic pore-like markings asin Z. triangularis, Koch ;
both shields have a double row of irregularly-shaped pits. The hair arma-
ture is characteristic ; on the side margins there are ten pairs, not including
the frontal spines, of strongly plumose hairs (fig. 156). The dorsal surface
also carries a number of spines which are less strongly plumose than those of
the margins; short spines spring from the marginal serrations. Sternum and
genital plate of normal shape and rather weakly chitinized. Ventro-anal
plate large, elliptical, though somewhat flattened on the front margin, with
a few plain hairs. Legs robust and armed with plain spines, those on the
middle segments of the first pair are somewhat stronger and are carried on
distinct tubercles. Length 340 uw, breadth 250 p.

This species is allied to 4. ornatus, Berl. (3); the body is relatively
narrower, and the sides less strongly rounded, the margins are more dis-
tinetly serrated and carry fewer plumose spines. The last-mentioned
character also separates it from Z. radiatus, Berl. (7, p. 9), in which the
spines are more numerous.

Locality. Female found at roots of decayed Henbane plants in August.

ZERCON PERFORATULUS, Berl.

1904. Berlese, 3, p. 269.—1914. Berlese, 9, p. 186.

Originally described as a variety of Z. triangularis, C. L. Koch, and later
raised to specific rank. Hasily distinguished by the marking of the dorsal
shields. In 4. triangularis these bear scale-like markings, while in the
present species the markings are largely replaced, more especially on the
second shield, by distinct punctures, and the hair armature also is weaker.

Localities. Glendalough, Co. Wicklow, found under chips of wood lying
on the ground, April; Knappagh Wood, Co. Mayo, in moss, August, in
company with 4. triangularis in both localities. i

CERATOZERCON BICORNIS (Can. et Fanzago). (Pl. 21. fig. 16.)
1877. Sevws bicornis, Canestrini et Fanzago, 21, p. 103.—1881. Gamasus bicornis,
Kramer, 29, p. 14.—1882. Lelaps bicornis, G. et R. Canestrini, 20, p. 78.—1885-
Sejus bicornis, Canestrini, 18, p. 91.—1887. Zercon bicornis, Berlese, 1, Fasc. xli.n. 8.

—1910. Berlese, 7, p. 346.
This species has been described and figured as having only one plumose
spine on the two chitinous horns of the posterior margin. Instead of this, in
27*

376 MR. J. N. HALBERT : NOTES ON ACARI,

the Irish specimen (length 342 w, breadth 220 «), there are two plain spines
(fig. 16), the outer one being a little longer than the other. It is possible,
however, that the specimen may be immature. Canestrini remarks: “le quali
portano all’ apice, nelle forme giovani, ciascuna due setole, nelle piu avonzate
di eta ciascuna una setola robusta diretta in dietro e in dentro” (20).

Locality. One female found under a stone on the Malahide sandhills in
August.

PoLYAsPINUS CYLINDRICUS, Berl. (PI. 21. fig. 17.)
1916. Berlese, 11, p. 134.—1917. Berlese, 16, p. 10.

This interesting genus was founded to receive the present species, and
belongs to the tribe Polyaspidini of Berlese’s recent classification of the family
Uropodidee (16). ‘he species may be recognized by the elongate sub-
parallel shape; the central area of the dorsum is protected by a long, narrow
shield, smooth at the centre, roughened at the sides ; at the end of this are
three small shields, of which the middle one is the least, arranged in a trans-
verse row. The marginal shields are represented by a row of small hair-
bearing plates, and on the actual body margin are numerous similar plates.
The venter is protected by strongly-chitinized and fused plates; their margins
are indicated by thickened ridges. Peritreme placed on the side margins of
the body. Hairs simple and blade-shaped. Legs robust, claws of the first
pair much reduced. Length about 670», breadth 3004. The male is
unknown. Identifieation verified by Dr. Berlese.

Locality. Several females found under logs of wood half buried in the
mossy banks of the stream between Glendalough and Laragh, in April.

TRACHYTES PYRIFORMIS (Kramer).

1876. Trachynotus pyriformis, Kramer, 28, p. 80.—1877. Canestrini et Fanzago, 21,
p. 68.—1882, Kramer, 30, p. 420.—1892. Trachynotus egrota var. pyriformis,
Berlese, 1 (Mesostig. Supp. p. 94).—1894. Michael, 33, p. 313.—1915. Berlese, 9,
p. 184.

The above are the principal records undoubtedly referring to Kramer’s
species, which was described in 1876. There has been confusion between
this species and the form described as Z. wgrota by Koch, ,and if the latter
were clearly recognizable it would be the type of the genus Trachytes.
There is doubt, however, concerning the identity of Koch’s 7. wgrota; a
species supposed to be the same was described and figured by Dr. Berlese (1,
Fase. xxxviii. n. 10), but he has since suppressed this, stating that it may
have been identical with either of his two recently described species,
T. lambda or T. tubifer (9, p. 185). It would then follow that 7. pyrifermis,
Kramer, is the type of the genus 7’rachytes. Fortunately this species was
carefully described and figured by Kramer (28). Mr. A. D. Michael has
recorded both 7. egrota, Koch, and 7. pyrifornus, Kramer, from English
loealities, but he did not describe the first-mentioned form.

WITH DESCRIPTIONS OF NEW SPECIES. 377

Locality. Several females were found under branches and pieces of wood
lying on the river bank at Glendalough in April.

TracHytss pi, Berl., var. PAUPERIOR, Berl.
1915. Berlese, 9, p. 135.

Differs from the typical form (7) in its paler colour, smaller size, and in
the form of the epigynum which is more elongate and punctured. The
length of the Irish specimen is 484, breadth 286. Hvidently closely
allied to 7. minma, Triig. (44, p. 448), but in the present form the anterior
part of the body is much more elongate and the shoulders are less marked.

Locality. A single female found under decayed wood at Poulanass,
Glendalough, in April.

Uroszius acuminatus, C. L. Koch.

Locality. A few specimens were found in company with numbers of Uropoda
obscura, U. lL. Koch, in decaying potatoes, Rocky Valley, Co. Wicklow,
September 1921. The species has been recorded from Italy, Switzerland,
Germany, and Great Britain. Hull queries the “ Cilliba vegetans” of (25)
as the present species, but the nymph recorded as such can have no rela-
tionship with Urosetus. ‘Vhe adult form of the “ Cilliba vegetans”’ of authors
has not been made known.

Diyycaus (PRopinycuus) caRinatus, Berl.
1908. Berlese, 3, p. 247.

This species is briefly described in the above reference, and it is not
figured; Dr. Berlese, who has kindly verified the identification, refers the
species (in lit.) to his subgenus Prodinychus. The size is rather small for’
this genus (length of Irish specimens 622 u, breadth 333 ~) ; shape of main
body a regular oval, anterior part rather acutely pointed. In the middle
line of the body there are two carinz, and between these and the slightly
raised lateral margins.the dorsum is concave.

Locality. 1 have found this distinct species in two localities in Co. Dublin,
once in decayed bulbs in January, and under bark of old moss-grown tree
trunks at Lucan in April.

PHAULOCYLLIBA VIRGATA (fZull).
1918. Cillibano virgata, Hull, 25, p. 44.

The original description is as follows:—‘‘ Sub-cireular, smooth, claret brown.
Dorsal shield divided by a very shallow median furrow forked behind the
middle and curved backwards on each side to the margin, leaving a transverse
part behind which bears a pair of clavo-pectinate setae. Two similar setz
stand in a line with these on the lateral margin. Hpigyne oval, rounded at
both ends, apiculate in front. Ventral slneld reticulate.”

378 MR. J. N. HALBERT: NOTES ON ACARI,

This species should certainly be referred to the genus Phaulocylliba, Berlese.
Apart from other differences it may be easily known from Cylliba by the
absence of the marginal plates, a feature not referred to in the original
description. The epigastric region of the venter is rather indistinctly defined
and includes the anal foramen, so that the original figure (25, pl. 1. fig. 4)
is inaccurate in this respect. The Irish specimens measure 970 u in length.

Locality. A few specimens found at roots of decayed Henbane plants at
Clontarf, Co. Dublin, in August.

TRACHYUROPODA (DinycuRA) CORDIERI, Berl.
1916, Berlese, 11, p. 145.

A species referred by Dr. Berlese to the subgenus Dinyewra, of which it
is apparently the type. A characteristic feature is the double row of small
piligerous plates lying between the extremities of the marginal shields at the
end of the body, much as in the genus Discopoma except that there is but one
row in this genus. These small plates vary greatly in number (16 to 22) in
T. cordieri due to a few of them fusing with one another, or with the
extremity of the marginal shields. The ventral line is thickly chitinized
and divides the epigastric region into two parts, which lie at different planes.
The female resembles the male in general structure; the epigyne is large
and of the usual arch-like form, its ventral surface is strongly punctured and
is produced anteriorly in a long chitinous process which ends in two or three
points. Length of female 563 4, breadth 357. Identification verified by
Dr. Berlese.

The nympha heteromorpha measures nearly 550 p by 385 w ; on the ventral
side the area surrounding the acetabula is distinctly reticulate. The ventro-
anal plate is of moderate size (length 70 u, breadth 165) ; it is supported
by a thick transverse bar which also forms the metapodial line.

Locality. The male, female, and nymph were not uncommon under damp
wood lying on the Malahide sandhills, Co. Dublin, in September.

TRACHYUROPODA TROGULOIDES, Can. et Fanzago, var. CcELTICA, Halbt.
(Pl. 21. fig. 18.)
1907. Tiachyuropoda celtica, Halbert, 22, p. 67.

Described as a new species in the above reference, but more accurately
placed as a variety of the present species, a decision in which Dr. Berlese
agrees (?n lit.). This variety differs from the typical form as described and
figured by Italian acarologists(1, 5, 21) in its smaller size (length of female
820 u, breadth 486; length of male 742, breadth 436). The form
is more parallel-sided, and the raised central part of the dorsum is of more
uniform breadth throughout, while in the typical form its terminal part is
much wider than the rest. The sculpturing of the ventral surface is also
somewhat different (figured in 22).

WITH DESCRIPTIONS OF NEW SPECIES. 379

Localities. Both sexes occurred in nests of the ant Lasius niger under
stones at Tallaght, Co. Dublin, and it was also found in company witb Lasius
flavus on Lambay Island off the coast of Dublin. Mr. A. D. Michael found
it with ants at Land’s Hnd, Cornwall (33), and Mr. H. St. J. K. Donisthorpe
records it from other English localities associated with Laséws niger and
T. cespitum (Entomologist’s Record, 1909 and 1920),

CILLIBANO DINYCHOIDES, Hull.
1918, Hull, 25, p. 45.

A male Cillibano found in company with Phaulocylliba virgata (Hull) is
probably to be referred to the present species. The Irish specimen agrees
with the description except in the measurements (length about 717 ~). The
length of the English specimen is given as 655 pm.

The short description 1s as follows: —“ Sub-circular, smooth, shining reddish
brown, uniformly convex. Leg grooves exceedingly faint. Male genital
aperture between coxz iv. rather large (width 85). Femur ii. with a stout
conical acute spine underneath. A clavate seta on each side of the anus,
projecting beyond the posterior margin.”

As is usual in this genus the marginal plates are entire, and in the present
species they are separated from the dorsal plate by a very thin line running
parallel to the margin in the posterior two-thirds of the body. The meta-
podial line curves inwards from the body margins to a point a little behind
the insertions of the last pair of legs, and it is widely interrupted in the
middle line of the body. The female is unknown.

Locality. One male found at roots of decayed Henbane at Clontarf,
Co. Dublin, August.

LABIDOSLOMMA LUTEUM, Kramer.

The species recorded in the Clare Island Survey Report (23) as L. cornuta,
Can. et Fanzago, is in reality L. lutewm, Kramer. The former is a much larger
species and differs also in certain details of structure ; it has not been found
in Ireland.

L. luteum is probably not uncommon in suitable localities in this country ;
in the Achill and Westport districts of Mayo it occurs under bark and
amongst mosses growing on trees. In the Dublin and Wicklow district I
have found it in pine woods.

PHYLLOTEGEUS PALMICINCTUM (Michael).

1888. Leiosoma palmicinctum, Michael, 32, p. 280.--1898. Liacarus palmicinctum,
Michael, 34, p. 42.—1913. Berlese, 8, p. 92.

Locality. Apparently a rare species in Ireland, two specimens were found
under stones at Doo Lough, near Muckross, Co. Kerry, in May. It is
recorded by Michael as occurring on the lichen Peltigera at Land’s End,
Cornwall; so far as I can ascertain this Acarid has been found only in

Britain.

380 : MR. J. N. HALBERT: NOTES ON ACARI,

CARABODES AFFINIS, Berl.
1913. Berlese, 8, p. 72.

Closely allied to C. marginatus, Michael, but may be known by the clavate
abdominal hairs. Dr. Berlese assures me the Irish specimens are referable
to this species, which he records as occurring under bark at Florence. The
pseudostigmatic organ has a curved upturned stem and a somewhat flattened
club. The lamelle are broad, flat and granulate, and there appears to be a
slightly raised ridge in the middle line of the cephalothorax. Interlamellar
hairs stout and plumose at their extremities. The hairs of the dorsum as
well as the marginal hairs are clavate, and their extremities are distinctly
plumose.

Locality. Common amongst moss and lichens on the Portmarnock
sandhills, Co. Dublin, in January, The Carabodes recorded as C. marginatus,
Michael in (39) is the present species.

ORIBATULA (HEMILEIUS) PLANTIVAGA, Berl.
1892. Oribatula tibialis, Berlese, 1, Fasc. lxiy. n. 1.—1895, Oribatula plantivaga, Berlese,
1, Fase. Ixxvii. n, 5 (reference in footnote).—1916. Berlese, 12, p. 322 (Redescribed).
Localities. Found near Mulranny, Co. Mayo, in September, and at Lucan
near Dublin in company with O. oblonga and other species in February.
Hull records it as occurring on rock lichens, Physcia, on the coast at Whitley
Bay.

Damenosoma MACULOSA, Warburton and Pearce. (PI. 21. fig. 20.)
1905. Warburton and Pearce, 49, p. 567.

A Dameosoma found under bark in the Tolka Valley near Dublin (January )
is apparently referable to this species. As it seemed not quite typical I sent
drawings to Mr. Warburton, who has kindly verified the identification. The
species is remarkable for the long, pointed cephalothorax, also the long
lamelle and the spotted dorsum, though, as the describers remark, these spots
are in the epiostracum and are very easily rubbed off. This is soin the Trish
specimen, in which they remain only near the margins. The pseudostigmatic
organs are bent backwards, and carry on the anterior surface at least four
distinct branches (PI. 21. fig. 196); anterior margin of abdomen truncated.
Length 418 p.

DaAMEOSOMA MINUS, Paoli, var. LAMELLATA, noy. (PI. 21. figs. 19a, b.)
1908. Paoli, 39, p. 48.

The form briefly described here must, I think, be identified as a variety of
the present species. It differs from the type in the larger size and in the
presence of well-defined lamellee on the cephalothorax ; in )). nunus these are
rudimentary.

Cephalothorax rather long and with sharply pointed extremity ; lamellee
present in the form of two strongly curved ridges reaching from beyond the

WITH DESCRIPTIONS OF NEW SPECIES. 381

middle to the pseudostigmata. The latter are circular and strongly chitinized.
Sensory organs with a stout slightly curved stem and a large club-shaped
extremity which is minutely spiculate. Abdomen elongate, pointed at both
extremities, hairs very weak. Hpimera well defined. Legs normal, mono-
dactyle. Length 240 yw, breadth 110 yw.

Locality. Found under dead wood at Poulanass, Glendalough, Co. Wicklow,
in March. The typical form is widely distributed, oceurring in Italy and
near Washington, U.S.A. (40).

CYMBHREMMUS MONILIPES (Michael).
1888. Notaspis monihpes, Michael, 32, p. 681.—1896. Berlese, 1, Cryptostig. ii. p. 37.
Locality. Glendalough, Co. Wicklow, in decayed tree-trunks, March.

HERMANNIELLA GRANULATA (JVicolet).
1855. Hermannia granulata, Nicolet, 35, p. 469.

In his well-known work on French Oribatids, Nicolet described and figured
two species of Hermannia, i.e., H. granulata and fH. arrecta (which he believed
were distinct), differing from each other in the form of the cephalothorax.
As far as | am aware the distinctness or otherwise of these species has not
been settled. It seems likely, however, that Nicolet’s specimens, which were
found in the same localities, represent but a single species, and that the
supposed differences in the form of the cephalothorax may be more apparent
than real. There is a tendency among acarologists to record H. granulata
in preference to //. arrecta, it is also the first mentioned of the two species
in Nicolet’s monograph, so I think it is better to refer the Irish specimens
here until the question has been decided.

Localities, Adults and nymphs were found in some numbers in a decayed
birch log at Glendalough in April, and it has also occurred in moss collected
at Howth, Co. Dublin, in June.

TARSONEMUS LATICEPS, sp. nov. (PI. 21. figs. 21 a-c.)

A species allied to Z. culmicolus, which is recorded by Reuter (41) as
causing injury to meadow grasses in Finland. | Differs in the form of the
body, the broader capitulum, and in other details.

Female. Form rather short and broad (length 195 u-210 w, breadth 110 »);
in ovigerous females the body is more elongate, becoming long oval (length
215 w), the enclosed egg measuring about 115. Side margins sub-parallel,
front margin truncated and half covering the head, end margin rounded.
Division between the cephalothorax and abdomen and the one at the middle
of the latter distinct, but the segments at end of the abdomen are ill-defined.
Hair armature very short except for the usual long pair close to the sides of
the cephalothorax, and a shorter pair at the angles of the front margin. The
sensory organs (PI. 21. fig. 21 a) arising from small cireular stigmata, are large,
they may be either globular, or leaf-like with pointed extremities, and both

382 MR. J. N. HALBERT: NOTES ON ACARI,

forms may be present in the same individual. The capitulum is very large
and wide in relation to the length (breadth about 40), truncated in front
and obtusely round at the end margin. Epimera of the first pair of legs
strongly defined. ‘The first two pairs of legs are short and stout; the long
terminal hair of the end pair is strongly recurved.

Male (Pl. 21. fig. 216). Form of the main body much as in female (Jength
180 w-215 pu, breadth 85 ~-100 4). The capitulum is less transverse (breadth
35), and the hair armature is stronger, there are two pairs of long hairs on
the cephalothorax. Legs short and stout; fourth pair (PI. 1. tig. 21 ¢) shaped
much as in 7’. floricolus, being without blades; second segment straight,
inner margin slightly sinuate at the extremity ; two hairs are present.
Third segment armed with a long spine-like hair, and at the base is another
shorter hair. Asin 7. brevipes the front margins of the third and fourth
epimera are ill-defined.

The male larva is as long as the adult (length 215 w), due to the large size
of the post-abdominal part. Hpidermis distinctly striated. Legs stout and
of very uniform size. The terminal part of the abdomen is strongly
constricted and is narrower (breadth about 50.) than the main body, and
there are two stout outwardly curved hairs at the apex.

The eggs of this species are large (length 125 u-135 pw, breadth 65 w), they
are of a regular long oval form and the surface is marked with numerous
light-refracting punctures.

Locality. Found in numbers in partly decayed Narcissus bulbs in County
Dublin, January. ‘The place of origin of the bulbs is uncertain.

At least three other species of Tarsonemus have been found in Ireland. One
of these, noticed injuring oats at Killyarden, Co. Donegal, in August iast, is
possibly referable to 7. spirifex, Marchal, the female agreeing well with
Korft’s figure of this species which is reproduced by Sorauer (Handbuch der
Pflanzen-Krankheiten, iii. p. 102). The male of 7. spirifex has the fourth
pair of legs peculiarly modified, and until this sex is found this identification
must remain somewhat doubtful.

The form of this female is a regular long oval (length 235 w, breadth 110 y).
The capitulum is rather broad (breadth 35 yw), but less so than in 7’. laticeps.
The sensory organs are very distinctly leaf-shaped with sharply pointed
extremities. The first two pairs of legs are short and stout.

This species was communicated by Dr. G. H. Pethybridge, who tells me it
attacks oats in much the same wayas 7’. culmicolus, Reuter, attacks meadow
grasses in Finland. Reuter says the mites are found on the haulm, presum-
ably within the leaf-sheath within the first node; they suck out the juices so
that the inflorescence wilts and dies.

Females of 7. floricolus were found on gooseberry plants at Lisburn, Co.
Antrim, in July.

Colonel Samman tells me he finds Acarapis woodi, Hirst, commonly in the
trachez of Irish honey-bees.

WITH DESCRIPTIONS OF NEW SPECIES. 383

BIMICHAELIA CRASSIPALPIS, sp. nov. (PI. 21. figs. 22 a, b.)

Allied to B. setagera, Berlese (4, p. 13), but larger, the palps are stouter,
the shoulders less prominent, and the structure of the skin, which is
elaborately sculptured, is different.

Colour white. Form sub-pentagonal (PI. 21. fig. 22 a), shoulders prominent
but less so than in B. setagera. Hpidermis striated and marked with a distinct
hexagonal pattern, the hexagons forming rosette groups round tlie body hairs,
the latter are sparse and are branched at the base (fig. 22). Cephalothorax
not well marked off from the abdomen ; a narrow area enclosed by chitinous
rods lies in the middle line, and two long sensory hairs are placed at its
proximal extremity; behind these hairs are two semicircular stigmata. I can
tind no trace, however, of the small, clubbed, sensory organs which arise from
these in other species of the genus (in a second specimen from another
locality they are also absent) ; it is likely, however, that such organs may be
sometimes present in the species. Abdomen thrown into folds and constricted
at the middle, apex somewhat truncate. Palps stout. Legs short and very
robust, distinctly areolated, hairs similar to those of the body. Length 320 p,
breadth 240 p.

Localities. Found in sphagnum moss collected by Mr. A. W. Stelfox on the
Garron Plateau, Co. Antrim, in July. It has also occurred in damp moss
from Glencullen, Co. Dublin, in April of the present year.

ALIcUS RosTRATUS, Z’rdq.

An Alicus found in two highland localities in Ireland agrees excellently
with the present species, which is recorded from the Sarekgebirge (Swedish
Lappland), except that the rostrum seems shorter than is described and
figured (46). Drawings were sent to Dr. Trigardh, and he believes there is
a difference in this respect. However, they agree in so many characters that
it does not seem desirable to describe the Irish specimens under a new name.

The colour during life is a reddish purple. Length 460 pu, breadth 270 p.
Cephalothorax with a distinct rostrum, and well marked off from the abdo-
men, central area defined by two subcutaneous chitinous ridges, the hinder
part of which is areolated at the middle, and encloses the usual two pairs of
long sensory hairs. Hyes small but distinct, placed on the front of lateral
swellings. Abdomen with moderately marked shoulders and a sparse
covering of short, spiculate hairs; during life the dorsum lies in distinct
folds. Legs longer and more slender than is usual in this genus.

Localities. Found in moss collected on the Comeragh Mountains, Co.
Waterford, in July; and in sphagnum from Glendhu, Co. Dublin, in
October.

Scirus InERMIS, Trdg. (Pl. 21. figs. 23 a, 6.)
This species was described from specimens found on the surface of a small

reservoir (Wassersammlung) at Gizeh, Cairo, in December 1900 (45).
There can scarcely be any doubt that the specimens here recorded are the

384 MR. J. N. HALBERT: NOTES ON ACARI,

same species. The palpi, which are characteristic in the species of this
genus, are identical with those of S. inermis, Trig. Dr. Trigardh, to whom
drawings were sent, agrees with me in this identification. The following is
a brief description of the Irish form:—Colour red (length 490 ~). Body of
the usual subrhomboidal form, shoulders prominent, the body margins
gradually narrowing to the posterior margin where there are three small
lobes. EHpidermis minutely lined. Maxillary plate longer than broad ;
mandible long (length about 286 yw), extremities reaching to near the end of
the second palp segment. Palps (PI. 21. fig. 23 6) long and slender, apparently
four-segmented, second and third segments armed at their ventral extremity
with a strong spatulate hair. Legs long and of very uniform thickness so
that the tarsi are truncated at the ends, where they are armed with two flat
lobes and small claws. Length of the first pair 510 pw, of the fourth 530 pm.

Locality. Found amongst dripping wet moss and liverworts (Conocephalus
conicus) at the overflow of a small reservoir at Greystones, Co. Wicklow,
July 1920, with such species as Notaspis lacustris, Oribata lucasi,
Episeius italicus, and others.

NEOPHYLLOBIUS SAXATILIS, sp. nov. PI. 21. figs. 24 a, b.)

Colour as in NV. elegans, Berl. Form oval, cephalothorax not well marked
off from abdomen. Hyes double lensed. There is a double row of six strong
hairs, placed rather close together, down the middle of the body, and a row
of eight similar hairs along each outer margin ; all of these hairs are spicu-
late, rather strongly curved and arise from small tubercles. Hpimera large,
inner margins indistinct, with from two to three hairs which are much
weaker than those of the dorsum. Genital area tapering to a point and
enclosing a small anal plate with four marginal hairs. Rostrum triangular,
ending in a single lobe. Palps very short and stout, second segment longer
than the two end segments together, with two dorsal spines, one spiculate
and stronger than the other; third segment with one dorsal spine ; fourth
segment ending in two curved spines, and carrying an appendage which is
contracted at the apex and has three spines.

Legs much as in JV. elegans, Berlese ; there is a long spiculate hair on the
third seoment of each leg, that of the last pair bent at the end, length about
264 w (fig. 24). The legs measure about 396 pw, 330 w, 363 w, and 420 w in
length.

A species allied to WV. elegans, Berlese (1, Fasc. xxxiv. n. 5), but differing
in the size and form of the body. The median dorsal series of spines are
more numerous and are placed closer together, and the long hairs on the third
segment of the last pair of legs are shorter and of different form. Verified
by Dr. Berlese.

Locality. Not wncommon on lichen-covered rocks at Howth summit,
Co. Dublin, September.

WITH DESCRIPLIONS OF NEW SPECIES. 385

°
BRYOBIA HUMERALIS, sp. nov. (Pl. 21. figs. 25 a—c.)

A small species, length about 330 yu, breadth 270 ~. Colour red. Body
strongly and suddenly narrowed at both extremities, shoulders very promi-
nent. Froutal tubercles of cephalothorax comparatively short, carrying
strongly-curved spatulate hairs (Pl. 21. fig. 250). End margin of abdomen
with five pairs of similar hairs, the three inner pairs heing very conspicuous ;
there is also a double row of three or four hairs near the middle line of the
body and a pair at the shoulders. Palps very short, appendage of fourth
segment large and stout, terminal claw strongly curved. Legs shorter and
stouter than in B. pretiosa and of more uniform breadth, the second seg-
ment of the first pair being noticeably shorter ; the two end segments are
of nearly equal length (Pl. 21. fig. 25 ¢).

Apart from the difference in the legs, this species may be known from
B. pretiosa by the smaller size, the shorter frontal tubercles, the more
strongly narrowed body, the more prominent shoulders, and the longer body
hairs.

Localities. Found at Clontarf, near Dublin, in March 1907. The corre-
spondent who sent me the specimens had his attention drawn to them by
the patches of red colour on a garden wall caused by the presence of large
numbers of this mite. North shore of Lough Neagh, in wet moss, August
1922. Mr. Hvans has found the species amongst moss collected near
Hdinburgh.

RHAPHIGNATHUS PATRIUS, Berl., var. TRUNCATUS, noy. (P]. 22. figs. 26a-c.)

A robust form evidently allied to . patrius, Berl., and in ail probability
a variety of this species. Differs notably in the narrower body ; of the type
Dr. Berlese remarks: “fere zeque longium ac latum humeratum”; the end
margin is very distinctly truncated, and the areolations of the epidermis are
fewer and larger.

Length 420 w, breadth 320. Colour blood-red, with a central dark area in
front and four dark blotches on each side of the body, much as in Berlese’s
drawing of B. clavatus (1, Fasc. xxii. n. 2). Cephalothorax bluntly pointed
in front, the sides slightly sinuate behind this, and they merge evenly with
the shoulders of the abdomen; the margins of the latter narrow gradually to
the truncated end margin, at each angle of which there is a hair somewhat
longer than the body hairs. There are nine pairs of hairs in all, including
the frontal hairs, and the latter are the only ones which have a trace of
secondary hairs. Hpidermis very coarsely punctured, and there are paired
circular pits close to the shoulders; the end of the dorsum is slightly
depressed. On the ventral side the anal plate and the areas surrounding the
epimera are strongly areolated, and on the hinder margins of the former are
two spine-like hairs. Palps (PI. 22. fig. 26 6) very stout; second segment with
a long, bent hair, which may, apparently, be either simple or trifid at the
extremity ; third segment with a similar dorsal hair; length of palp about

386 MR. J. N. HALBERT: NOTES ON ACARI,

95. Legs very stout (length of first pair 264 yw, of fourth pair 254m), with
strong dorsal hairs, which are spiculate at the base (PI. 22. fig. 26¢).

Locality. A single specimen found amongst wet moss from the banks of a
mountain-stream at Glencullen, Co. Dublin. Collected by A. W. Stelfox in
October 1921.

IT have also found amongst moss in a mountain-stream at Kilmashogue,
Co. Dublin, a Rhaphignathus the identity of which is uncertain. It is very
closely allied to a form described by Trigardh as R. patrius var. brevipalpis
(46, p. 470). The Irish specimen differs in the smaller size (length 368 p,
breadth 275 «); it is less rotund, the puncture less regular, and the palps are
relatively larger.

RHAPHIGNATHUS PLUMIFER, sp. nov. (PI. 22. figs. 27 a, b.)

A very distinct species, which may be easily known from its congeners by
the small size, the strongly plumose hairs, and the beautiful sculpturing of
the epidermis. Verified by Dr. Berlese.

Male. Length 210, breadth 120m. Colour bright red. In shape
resembling A. siculus, being ovate, though the end of the body is produced
in a point. All of the body hairs are strongly plumose. The epidermis is
marked with a very distinct polygonal pattern as well as being punctured
(fig. 27). Ceplialothorax large in relation to the abdomen, carrying three
pairs of hairs and a strongly-curved pair of frontal hairs. Eyes small yet
distinct, and placed on the side margins. The main abdomen is much higher
than the apical part, and is truncated at the end, where there are two rather
long (40) hairs, curved inwards at the extremities. In addition to these
there are six pairs of hairs on the dorsum. LHpimera well defined ; anal
plate large, rounded on the front margin. Maxillary plate transverse;
rostrum sharply pointed. The palps are very stout, second segment with
two strongly-plumose hairs. Legs short and robust, armed with both simple
and plumose hairs like those of the body.

Locality. A single specimen found under a stone amongst heather on

Howth Head, Co. Dublin, in September.

RHAPHIGNATHUS LONGIPILIS, sp. nov. (Pl. 22. fig. 38.)

Colour blood-red with blackish markings. Form short and broad. Front
margin obtusely pointed, end margin rather truncated. Eyes small, placed
near middle of side margins. Upper surface of cephalothorax and abdomen
with a distinct, raised polygonal network. Abdomen well defined, shoulders
rather prominent. Hair armature, including the frontal bristles, very long
(about 125), curved towards their extremities, and there is no trace of
secondary hairs. Hpimera small. Maxillary plate transverse. Palps very
short and robust, distinctly stouter than the legs. The latter are of moderate
length, and rather slender for this genus. Length 290 yw, breadth 380 p.

WITH DESCRIPTIONS OF NEW SPECIES. 387

Localities. Found amongst damp moss at Glencullen in April, and in
moss collected from pools in the River Dodder, at Old Bawn, in May, by
Mr. A. W. Stelfox. Both localities are in Co. Dublin.

STIGMAUS ANTHRODES, Gerl., var. RETICULATUS, nov. (Pl. 22. figs. 28 a, b.)

A large, robust species. The Irish specimens are to be referred to a variety
in which all of the body plates are sculptured with a very distinct, polygonal
reticulation. Colour bright red. Cephalothorax protected by a large central
shield and two small plates at the posterior corners. On the dorsum of the
abdomen are ten shields, comprising three on the anterior part, then four
smaller ones in a transverse row; behind these are two similar plates, and a
large unpaired shield lies at the end of the abdomen. ‘The palps, legs,
epimera, and ventral plates are all marked with a polygonal network.
Leneth 550 pw, breadth 310 w.

Localities. Amongst hay brought into a cave at Doneraile, Co. Cork,
in July. Under refuse on the sea-bank at Malahide, in May. Amongst
garden refuse at Rathgar, Dublin, in April. Mr. A. W. Stelfox found it
abundantly in the last-mentioned locality.

CALIGONUS SCAPULARIS (Koch), Berlese.

A shining red species of very convex form. The eyes are very large and
the body hairs are long and curved.

Localities. In sphagnum, Lough Dan, in August, and in moss from pools

hy the River Dodder, near Tallaght, in May.

CHEYLETUS VENUSTISSIMUS, Koch.

Locality. Found in numbers on a Hay Moth (Caradrina) at Sandymount_
in November by Mr. J. G. Rhynehart. Cheyletus eruditus, Schr., has been
found on pine-shoots in August.

HYDRACARINA.

Drammenia crassipalpis, Sig Thor. The genus Vrammenia was established
by Thor in 1913 (44) with J. elongata as the type-species. In the same
paper Thor partially described a second species, D. crassipalpis, but apparently
the single specimen, found at Drammen in Norway, was lost before a
description had been made; such characters as are mentioned chiefly refer
to points in which this species differs from 1). elongata. As far as one can
judge from these notes, the form briefly described below is in all probability
the same as D. crassipalpis, Thor. Colour yellow with brown cecal markings.
Length 474, breadth 374. Cuticle strongly chitinized, except for a
marginal band separating the dorsal and ventral plates. Front margin
truncated, sides gradually increasing in width to beyond the middle of the
body, posterior margin slightly flattened. Dorsal plate punctured, touching

388 MR. J. N. HALBERT: NOTES ON ACARI,

the front margin of the body, separated from edge of ventral plate at the
side and end margins by a rather broad band of soft, striated cuticle, which
contains about five pairs of pores. On the dorsal plate is a double row of
similar pores, near each of which is a fine hair. Hyes placed at each corner
of the front margin on the dorsal outline of the body. Frontal bristles very
short (Pl. 22. fig. 29a). Hpimeral region much as in D. elongata, except
the outer border of the fourth pair, instead of being absent, is indistinctly
defined (Pl. 22. fig. 296). The genital plate is shorter (length 132 p,
breadth 100). Length of maxillary plate to tip of rostrum 120. Palps
very stout, the second segment broader than the legs (breadth in dorsal view
49 w): on its inner ventral margin is a low, rounded prominence and a strong
hair as in the type-species.. Inner distal angle of the fourth segment pro-
duced in the form of a strong triangular tooth, and beside this is a smaller
tooth, both bearing fine hairs ; near the dorsal line of the palps are a few
short spines (P1. 22. fig.29¢). Legs of moderate length, stout, armed with
short spines and without swimming-hairs; the lengths are about 240 u,
374 pe, 330 pw, and 410 pw.

Locality. A single specimen found amongst sphagnum moss collected on Ott
Mountain, in the Mourne Mountains, in May by Mrs. R. F. Scharff.

Liania bipapillata, Sig Thor, has been found in the River Dodder, near
Dublin, in November ; in the River Dorgle, at Powerscourt, and in the stream
flowing between the lakes at Glendaiough, Co. Wicklow, in April. The colour
of living specimensis pale yellow with brownish markings. Megapus gibberi-
palpis, Piersig (40), occurs in streams on the Dublin Mountains at Kilkee and
Kilmashogue, and in the River Liffey. Also in the stream flowing from Lough
Highter, on Carrantuohill Mountain, in Co. Kerry. Arrhenurus insulanus,
Koenike: a single immature specimen of this species occurred in a small Chara
pool at Gollierstown, Co. Dublin, in October. The male has not been deseribed.
The female is red in colour, and is remarkable on account of the large wing-
shaped genital plates and the long and narrow epimera especially of the third
and fourth pairs. The species is well figured by Dr. Koenike (26) ; the type
was found on Norderney. 1 believe this species has also been found by
Mr. ©. D. Soar in England.

Apart from a few records of the common //ydryphantes ruber, de Geer,
nothing has been published on the species of Hydryphantes occurring in
Ireland. I have recently been making a preliminary examination of
specimens of this interesting genus from various Irish localities, and find there
are at least six species found in this country, counting 1. prolongatus, Thon,
of specific rank. A certain amount of variation occurs in the form of the
eye-plates of Hydryphantes ; and as the structure of the plates is of great use
in the identification of the species, a series of figures are given in the present
paper, in the hope that they may be of interest for comparison with those of
the same species in other countries.

WITH DESORIPTIONS OF NEW SPECIES, 389

The type-species of the genus, H. ruber, de Geer, is common and wide-
spread in Ireland (PI. 22. fig. 30), as is also H. prolongatus, Thon (PI. 22.
fig. 31), which is sometimes recorded as a variety of the former ; in my
experience it is rather the exception to find both of these occurring in the
one locality. The first-mentioned has been found in ponds and lakes in
Donegal, Antrim (L. Neagh), Monaghan, Galway, Wexford and Dublin, and
the latter in Donegal, Dublin, Kildare, Galway and Kerry. HH. prolongatus
is perhaps of more frequent occurrence. The eye-plate and palps of a curious
form, which must, I think, be regarded as an aberration ot Hf. prolongatus,
are figured (Pl. 22. figs. 32a, b). The general structure is as in the species,
but the eye-plate is smaller and narrower especially across the front margin
(length along middle line 3863, breadth 373 4), and the median eye is
placed much nearer the middle of the plate. Unfortunately, only one
specimen was found in a small lake on Lough Salt Mountain, in Donegal.
FA. bayeri, Pisarovic (PI. 22. fig. 33), is apparently rare; I have founda
single specimen in a drain by the River Shannon, at Portumna, in June.
H. bayeri nonundulata, Viets (Pl. 22. fig. 34) was found in a pool at the
entrance to Glenshelane Valley, at Cappoquin, Co. Waterford, inJune. It has
also occurred under stones on the marshy edge of Bount Brown Lough, near
Westport, in Co. Mayo. The water-level of the lough had fallen considerably
at the time (July 1911). In the structure of the eye-plate this form bears a
strong resemblance to H. planus, Thon, but the plate is more abruptly
narrowed behind the anterior corners, the breadth across the hinder part is
relatively greater, and the posterior emargination is less deep than in 7. planus.
The median eye is placed far back as in the type. Viets records this form
from Eastern Prussia (47). H. crassipalpis, Koenike, was found in ponds near
Enniscorthy, Co. Wexford, in May. This species may be easily recognized by
the form of the eye-plate and the short, thick palps ; described by Dr. Koenike
from Borkum and the neighbourhood of Bremen, where it is rare (27). At
Killarney I have found a Hydryphantes which seems referable to a form of
this species. ‘The eye-plate (PI. 22. fig. 35) is smaller (length along middle
line 298 w, breadth 418 ~) and the anterior corners are most acute; the sides
are more deeply indented, and the posterior emargination is deeper. The
palps (fig. 35 6) are shorter (length about 300) than in the typical form ;
the second segment (length 110) has four short dorsal spines on the upper
surface and three longer, feathered hairs placed at the upper corner ot the
inner surface of the segment. This form, which may be called lacustris,
forma noy., was found in Loosecaunagh Lough, between Killarney and
Kenmare, in the month of April. H/. placationis, Thon (Pl. 22. fig. 36), not
common; found in ponds near Galway in June, and in ponds in the Pheenix
Park, Dublin, in April. . dispar (Schaub) is apparently rare in Ireland;
found in company with the preceding species in ponds in the Phoenix Park
in April. ‘The eye-plate is figured (Pl. 22. fig. 37).

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 28

390 MR. J. N. HALBERT: NOTES ON ACARI,

BIBLIOGRAPHY.

BERLESE, A.

1, Acari, Myriapoda et Seorpiones hucusque in Italia reperta. Patavii, Florentiz,

1882-1892.
. Acari nuovi, Manipulus i. Redia, i. 1908.
Acari nuovi, Manipulus ii. Redia, i. 1908.
. Acari nuovi, Manipulus iii, Redia, ii. 1904.
Acari mirmecofili. Redia, i. 1903.
Elenco di generi e specie nuovi. Redia, v. 1908.
Lista di nuovi specie. Redia, vi. 1910.
Acari nuovi, Manipulus vii.vili. Redia, ix. 1913.
. Acari nuovi, Manipulus ix. Redia, x. 1914.
. Centuria prima di Acari nuovi. Redia, xii. 1916.
11. Centuvia secunda di Acari nuovi. Redia, xii. 1916.
12. Centuria terza di Acari nuovi. Redia, xii. 1916.
13. Centuria quarta di Acari nuovi. Redia, xiii. 1918.
14. Centuria quinta di Acari nuovi. Redia, xiv. 1920.
15. Indice sinonimico (A. M. et 8.). Redia, xiv. 1921.
16. Intorno agli Uropodidee. Redia, xiii, 1917.

COHMIARARwWD

CaNESTRINI, G.
17, Acari nuovi o poco noti. Atti R. Ist. Veneto, 1888.
18. Prospetto dell’ acarofauna italiana. Atti Soc. Veneto-Trentina. Padova, 1885-1899.
CANESTRINI, G. et R.
19. Nuovi specie del genere Gamasus. Atti R. Ist. Veneto, vii. (v.), 1881.
20. I Gamasi italiani. Monografia. Atti Soc. Veneto-Trentina, viii. 1882.
CanesrRintI, G., et F. Fanzaco.
21. Intorno agli Acari italiani. Atti R. Ist. Veneto, iv. (v.), 1887.
Havsert, J. N.
22. Acarina of Lambay. Irish Naturalist, xvi. 1907.
23. Clare Island Survey. Acarinida Il. Proc. Roy. Irish Acad. xxxi. 1915.
24. The Acarina of the Sea-shore. Proc. Roy. Ivish Acad. xxxy. 1920.
Hutt, J. i. ,
25. Terrestrial Acari, Tyne Province. Trans. Nat. Hist. Soc. Northumberland, Xc.,
vy. (n. s.), 1918.
Kowntxke, F.
26. Sechs neue norddeutsche Wassermilben. Abh. Nat. Ver. Bremen, xx. 1911.
27. Neue und neubenannte Wassermilben. Abh. Nat. Ver. Bremen, xxii. 1914.
Kramer, P.
28. Zur Naturgeschichte einiger Gattungen aus der Familie der Gamasiden. Arch. f.
Naturg., Jahre. xlii. 1876.
29. Ueber Milben. Zeitschr. f. d. ges. Naturwiss. (vi.) liv. 1881.
30. Ueber Gamasiden. Arch. f. Naturg., Jahre. xlvili. 1882.
31, Ueber Milben. Arch. f. Naturg., Jahre. lii. 1886.
Micuast, A. D.
32. British Oribatide. Ray Soc., London, 1883-1887.
33. Notes on the Uropodide. Journ. Roy. Micr, Soc. xxxii. 1894.
34. Oribatidee. Das Tierreich, 3 Lief. Acarina. Berlin, 1898.
Nicoxet, M. H.
35. Histoire naturelle des Acariens qui se trouvent aux Environs de Paris, Archives du
Muséum d’Hist. Nat. vii. 1855.

WITH DESCRIPTIONS OF NEW SPECIES. 391

Ovuprmans, A. C.
36. Acariden von Borkum und Wangeroog. ‘Notes on Acari,’ ix. serie. Abh. Nat.
Ver. Bremen, xviii. 1905.
37. New List of Dutch Acari. Second Part. Tijdsch. v. Kntom. xly. 1902.
38. Acarologisches aus Maulwurfisnestern. Archiv f. Naturg., Jahrg. lxxix. 1913-14.
Paout, G.
39. Monografia del genere Dameosoma Berl. e generi affini. Redia, v. 1908.
Prersie, R.
40. Deutschlands Hydrachniden. Zoologia, xxii. 1897-1900.
Reuter, H.
41. Ueber die Weissabrigkeit der Wiesengriser in Finland. Acta Soc. Fauna Fennica,
xix. 1900.
ScHWEIZER, J.
42. Beitrag zur Kenntnis der terrestrischen Milbenfauna der Schweiz. Ver. Naturforsch.
Gesellsch. in Basel, xxxin. 1922.
Tuon, K.
43. Monographie der béhmischen Hydryphantes Arten. Bull. internat. Acad. Se.
Bohéme, 1899.
THOR, Sic.
44. Drammenia, eine neue Bachmilbengattung aus Norwegen, nebst Bemerkungen iiber
die systematische Stellung von Drammenia und Bandakia. Zool. Anz. xliii. 1913.
TRAGARDH, I.
45. Acariden aus Aegypten und den Sudan. Results of the Swedish Zool. Exp. to the
White Nile, 1901.
46. Acariden aus dem Sarekgebirge. Naturwiss. Untersuch. d. Sarekgebirges in
Schwed. Lappland, iv. Zoologie, 1910.
Viets, K.
47. Hydracarinologische Beitrige, ix.,.x. Abh. Nat. Ver. Bremen, xxix. 1918.
48. Hydracarinen aus norddeutschen und schwedischen Quellen. Archiv fiir Hydro-
biologie, xii, 1920.
Warsurton, C., and N. D. F. Pearce.
49. New and rare British Mites of the family Oribatidee. Proc. Zool. Soc. London. 1905.

EXPLANATION OF THE PLATES.

PratE 20.
Fig. 1. Gamasus (Eugamasus) crassitarsis, sp.nov. Male. a. Chelicere. 6. Second leg.

c. Process on outer side of femur of second lee, dorso-ventral view.

2. Gamasellus (Protolelaps) granulatus, sp. noy. Female. a. Under side. 6. Part
of dorsal plate and soft epidermis.

3. Gamasellus (Protolelups) subnudus?, Berl. Larva.

4. Lelaps latisternus, sp. nov. Female. Under side.

5. Lelaps simplex, sp. nov. Female. Under side.

6. Lelaps fimbriatus, sp. nov. Female. Genital and ventro-anal plates.

7. Lasioseius (Lasios.) gracilis,sp.nov. Female. a. Upper side. 6. Hair of posterior
margin. ¢. Ambulacrum.

8. Lasiosetus (Lasios.) ometes, Oudem. Female. a. Under side. 6. Shoulder bristle.
c. Chelicerze.

9. Lastoseius ( Episeius) glaber, Berl., var. curtipes, nov. Female. Under side.

10. Lasioseius ( Episeius) sphagni, sp. noy. Female.

392

MR. J. N. HALBERT: NOTES ON ACARI.

Fig. 11. Lastoseius (Episeius) italicus, Berl. Male cheliceree.

Fig.

Fig.

12.
18.
14.

Lasioseius (Episeius) michaeli, sp. nov. Male chelicere.
Lasioseius (Episeius) major,sp.nov. Female: a. Under side. Male: 6. Chelicere.
Lasioseius (Platyseius) subglaber (Oudem.). Male chelicere.

(The male chelicerz figured in 11, 12, and 14 are drawn to the same magnification.)

15.

16,
Wi
. LTrachyuropoda troguloides, Can. et Fanzago, var. celtica, Halbt. Upper side.

. Dameosoma minus, Paoli, var. lamellata, nov. a. Upper side. 6. Pseudostigmatic

PLATE 21.

Zercon tragardhi, sp. nov. a. Upper side. 6. Plumose hair.
Ceratozercon bicornis (Can. et Fanzago). One of the terminal tubercles.
Polyaspinus cylindricus, Berl. Upper side.

organ.

. Dameosoma maculosa, Warburton and Pearce. Cephalothorax.
21. Tarsonemus laticeps, sp. nov. Female: a. Anterior part of cephalothorax. Male:

b. Under side; ¢. Fourth leg.

. Bimichaelia crassipalpis, sp. nov. a. Upper side. 6. Hexagonal marking of

epidermis.

. Seirus inermis, Trigardh. a. Upper side. 6. Palp.
. Neophyllobius saxatilis, sp.nov. a. Upper side. 6, End segments cf the fourth leg.
. Bryobia humeralis, sp. nov. a. Upper side. 6. Body hair. ce. First leg.

Puate 22.

. Rhaphignathus patrius, Berl., var. truncatus, nov. a. Upper side. 6. Right palp.

c. Segment of leg.

. Rhaphignathus plumifer, sp. noy. a. Upper side. 6. Epidermis.

. Stagmeus anthrodes var. reticulatus, nov. a. Upper side. 6. One of the dorsal shields.
. Drammenia crasstpalpis, Sig Thor. a. Upper side. 6. Epimeral region. c. Palp.
. Hydryphantes ruber, de Geer. Hye-plate (Riyer Bann).

. Hydryphantes prolongatus, Thon. FEye-plate (Galway).

. Hydryphantes prolongatus, Yhon (Ab.). a. Eye-plate. 6. Palp (Donegal).

33.
. Hydryphantes bayeri var. nonundulata, Viets. Kye-plate.

. Hydryphantes crassipalpis, Koen., forma lacustris, nov. a. Eye-plate.. 6. Palp.
. Hydryphantes placationis, Thon, Kye-plate.

. Hydryphantes dispar, yon Schaub, Eye-plate.

. Rhaphignathus longipilis, sp. nov. Palp.

Hydryphantes bayert, Vhon. Wye-plate (River Shannon).

(All eye-plates of Hydryphantes ave drawn to the same magnification,
with the exception of number 35.)

Note I.—Lasrosrits (Lasios.) OMETES, Oudem.

In a paper just received, Vitzthum describes and figures this species from Austrian
specimens found in the borings of “ bark-beetles.” (Arch. f. Naturg. 89 Jahrg. 1923.)
Note II.— Drammenia crassipalpis, Sig Thor. Additional localities for this species are—
Stream above Lough Nahanagan at 1500 feet, in submerged moss in June. Glencree,
amongst wet moss and sphagnum at 500 feet in September; both of these localities
are in the County Wicklow. Also amongst sphagnum on leatherbed Mountain in
County Dublin at an elevation of 1600 feet, in September 1922.

Ath July, 1923. J.N. H.

HALBERT. JOURN. LINN. Soc., ZOOL. VOL. XXXV. PL. 20.

SPECIES OF ACARI.

JOURN. LINN. SOc., ZOOL. VOL. XXXV. PL. 21.

HALBERYs.

SPECIES OF ACARI.

JourRN. LINN. SOc., ZOOL. VOL. XXXV. PL. 22.

HALBERT

SPECIES OF ACARI.

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Vou. XXXV. ZOOLOGY. No. 236.
CONTENTS.
Page

I. A Form of Dimorphism and Asexual Reproduction in Ptychodera
capensis (Hemichordata). By J. D. F. Grucurist, M.A., D.Sc.,
F.L.S., Professor of Zoology in the University of Cape Town.
(WILD ANGE SSSRST OOS) Gren sdonapeehee sare ancenar re ccaasane tac sasacn st eace 393

II. On the Morphology of the Head Capsule and Mouth-parts of
Chlorops teniopus Meig. (Diptera). By J. G. H. Frew, M.Sc.,
Ministry of Agriculture Research Scholar. (From the Ento-
mological Department, Rothamsted Fxperimental Station,
Harpenden.) (Communicated by Dr. A. D. Ins, FJ..8.)
(Waitine cee Roxct= townes.) aera cea mereenerseen sist, toenails cscs auteioee 399

III. The Crustacean Plankton of the English Lake District. By
Rosert Gurney, M.A., F.L.S. (Plate 23 and 3 Text-
IEGAUTRESS oo te exis dbbeconecoatoudoacs. onadteroetn a DRRt en eEeeeT EenE eer ec Ali

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DIMORPHISM IN PTYCHODERA CAPENSIS. 393

A Form of Dimorphism and Asexual Reproduction in Ptychodera capensis
(Hemichordata). By J. D. F. Gitcurist, M.A., D.Sc., F.LS.,

Professor of Zoology in the University of Cape Town.
(With 7 Text-figures.)
[Read 7th June, 1923.)

Tur Hemichordata, considered as including the three groups represented
by Cephalodiscus, Balanoglossus, and Phoronis, are of such importance as
possible connecting links between the great groups of Vertebrates and Inver-
tebrates that any new facts bearing on their relationship to each other or
to other phyla of the animal kingdom are of particular importance. The
following observations on a species of Balanoglossid, which exhibits a form
of dimorphism, associated with sexual and asexual reproduction, may there-
fore be of special interest.

There appear to be at least four different species of ‘“ Balanoglossus” in
S. Africa: three, which I have already described (Trans. Phil. Soc. S. Afr.
vol. xvii. pt. 2, p. 151, 1908, and Ann. 8. African Museum, vol. vi. pt. 2,
p- 207, 1908), Ptychodera capensis, P. proliferans, P. natalensis, and an
undescribed species, of which only a single imperfect specimen has as yet
been found. The first and second were found in a tidal sandy pool in False
Bay, the third in the muddy mangrove-bordered flats of Durban Bay, the
fourth in a crevice of a rock at low-water mark in False Bay. The first two
are obviously different from the third and from the fourth, which does not
belong to the genus Ptychodera. The first two also seem to be quite distinet
from each other, but their constant association in the same pool, their
anatomical resemblance in all but the posterior regions of the body, and the
fact that there is a normal process of asexual multiplication in one of them,
gave rise to the suspicion that they might represent sexual and asexual
forms of the same species.

Both species belong to the genus Ptychodera, the Tauroglossus of Spengel,
and there appears to be no specifie difference in size, shape, or structure of
the proboscis and collar region. The size, shape, colour, and position of the
genital wings immediately behind the collar are also similar. The differences
in the posterior part of the body are, however, well marked, for (1) in
P. proliferans (fig. 2) there are no hepatic ceca, either external or
internal ; (2) the caudal region is not clearly marked off from the branchio-
genital region, and the bright yellow gonads seem to be continued into this
region as two narrow streaks on either side of the intestine, which is never
filled with sand as in P. capensis ; (3) asexual reproduction takes piace by

LINN, JOURN.—ZOOLOGY, VOL. XXXV. 29 :

SORE,
neal

sy

2,
Com

394 PROF, J. D. F. GILCHRIST ON DIMORPHISM AND

successive fragmentations or proliferations at the extremity of the tail ; (4) a
further difference is that, though the two species were found in the same
locality and under the same conditions, they do not usually oecur at the same
season, P. capensis being found chiefly, often exclusively, in the winter
months, P. proliferans in the summer months, a fact which may, however,
indicate a possible relation between the two species. To determine this point
extensive collections were made at various seasons, but no indication of any

Fie, 1.

“deseeysereytestites:

Ptychodera capensis.

Fie. 2.

Piychodera proliferans.

transition from one form to the other was observed. The external differences
between the two are shown diagrammatically by figs. 1 and 2, in which the

‘bright yellow gonads are indicated by stippled shading and the hepatic czeca

by black. Figs. 3 and 4 represent regenerating fragments, two and four
wecks respectively after breaking off from the parent.

As both species are hardy animals, thriving fairly well in captivity when
supplied with abundant pure sea-water, a number were kept in small glass
tanks partly filled with sand, and were examined from time to time. The

ASEXUAL REPRODUCLION IN PTYCHODERA CAPENSIS. 395

process of asexual reproduction was continuous in the summer months, but
not rapid. In the case of half-a-dozen P. proliferans, kept in a separate
vessel, about four to six small fragments were given off every week. Fora
time these remained in close proximity to the parent, but many of them were
subsequently found free on the surface of the sand, and were readily moved
about by any commotion of the water—probably a means of dispersal.
Some of these fragments were isolated and examined from time to time.
They moved about freely, a proboscis and traces of a collar appeared, and
for three weeks seemed to be developing into a form like their parent
P. proliferans, but, about the end of this period, some of them were seen to
be developing hepatic ceca and genital wings, while still retaining the bright
yellow colour and narrow intestine of the parent. Finally, after about a
month, they had the general characteristics of P. capensis, with a few
well-marked czeea and whitish caudal region, now containing a few grains of
sand. The bright yellow colour had entirely disappeared even from the
branchio-genital region, and they could not be distinguished from small
P. capensis at a stage in which the gonads are not yet developed.

Fre. 3. Fre, 4.
Proliferated part 14 days after Proliferated part 25 days after
separation. separation.

Tn order to ascertain further details of the transformation of the one form
into the other, various stages were examined by sectioning. Sections of the
tail region of P. proliferans show that the lateral septa are well developed
throughout its length, and are in intimate connection with the gonads, which
contain eosinophil globules, and, in some cases, one or two ova. The same
condition is¢ of course, found in the segments which break off from its
posterior extremity, and this is maintained until the stage when the proboscis
with notochord, &e., have appeared.

Figs. 5 and 6 show the conditions in a segment in which the proboscis is
clearly: shown, but not the collar region, and the gonads occur, not as a
continuous streak, but at intervals, so that in some sections (fig. 5) the
gonads still occupy a large part of the body cavity, but a few sections
further on (fig. 6) they are almost entirely absent, though the lateral
septa are still present. Sections of the same region at a later stage (about
4 weeks) (fig. 7) show an entire absence of gonads and no trace of lateral
septa. Fragments of food-material—diatoms, protozoa, and flocculent
organic matter—may now be seen in the intestine.

Evidently, therefore, the two species are identical, P. proliferans being
capable of giving rise asexually to P. capensis. The question then arises as

PAG)

396 PROF. J. D. F. GILCHRIST ON DIMORPHISM AND

to whether this is a case of alternation of generations, P?. capensis being the
sexual form. To ascertain this the genital pleuree of P. proliferans were
examined by teasing, sectioning, and compression, the last-named method
being the most suitable for examining a large number of individuals, as the
ova are fairly large (12 mm.) and conspicuous. The results of the
examination were not consistent, some haying apparently no ova, some only
a few, and some a fair number.

Transverse section of caudal region of the stage shown in fig. 3.
gon. =gonads ; 7.=intestine; sep.=septum.

Another section of caudal region Transverse section of
further back than that shown caudal region at the
in fig. 5. stage shown in fig. 4.

Towards the end of the summer (May) the six P. proliferans kept under
observation had become considerably reduced in size, and a change was
observed. The posterior region of the body became of a paler colour, the
yellow streaks being reduced to a few scattered patches, and the intestine

ASEXUAL REPRODUCTION IN PLYCHODERA CAPENSIS. 397

contained a few grains of sand. At a later date hepatic ceca appeared
about the middle of the body, and subsequently all were transformed into
typical P. capensis, with fairly well-developed gonads, in which, however,
there were no ova. Thus both the parent and its proliferated parts ultimately
assumed the form of P. capensis.

Ptychodera prolijerans would seem, however, to be capable of reproducing
sexually, as the gonads contain well-developed ova. The facts, however, that
(1) ova may occur in the proliferated part, and be used up like the granules
in the growth of the body, and (2) that the ova ultimately disappear in the
reduced adult, seem to indicate that there is no sexual reproduction in this
form. ‘There can be little doubt that P. capensis, into which both buds and
parent are transformed, reproduces sexually, so that the whole process looks
like a modified form of alternation of generations. There is evidence, how-
ever, which seems to indicate that P. proliferans arises from P. capensis by a
division of the latter in front of or at the hepatic region, and subsequent
prolongation and proliferation of the genital region, followed finally by
regeneration of the lost hepatic and caudal region.

The nutritive eosinophil giobules or granules evidently play an important
part in the life-history of the animal, and their relation to the sexual elements
may therefore be further enquired into.

‘They have been noted in many other HEnteropneusta, but their origin and
function remain in doubt. Spengel found that, in P. minuta, they were con-
tained in cells in which no nuclei were seen. In P. flava Willey found no
normal nuclei, and ‘believed that the nuclei of the cells undergo a process
of degeneration analogous to fatty degeneration. They were found to be
present when the germ-cells were fully developed, but both authors found .
that they disappeared at the period of complete sexual maturity in the
species they exainined. As to their functions, Spengel could come to no
definite conclusion ; Willey suggested that they are partly for the nutrition
of the growing germ-cells, but principally for providing an albuminous
covering to protect the germ-cells during maturation.

In P. capensis they are absent only in the young stages. They are present
from the time the germ-cells begin to develop and increase in numbers with
them, and are present when the animals are fully developed. They do not
disappear at complete sexual maturity, though the sexual elements then pre-
ponderate, and they subsequently (towards the end of the winter, when
exceptionally large specimens were found) appear to increase in number,
their bulk greatly exceeding that of the remaining germ-cells.

In P. proliterans stage they constitute the main mass of the gonads.
These are greatly attenuated posteriorly and occur, as already stated, as two
narrow streaks of a bright yellow colour in the elongated posterior region of
the body, which is rounded and without pleural ridges in the living animal.
The relative abundance of ova and globules was ascertained by the methods

398 DIMORPHISM IN PTYCHODERA CAPENSIS.

already stated, and it was found that the well-developed gonads consisted
mostly of globules alone, the eggs being only present, and in diminished
numbers, in some cases.

The functions of globules in this case seem obvious, namely, the sustenance
and growth of the non-feeding P. proiiferans, and of the buds, till they
reached the feeding stage.

The origin of globules and ova may best be studied in sections of P. ca-
pensis at a stage when the gonads are beginning to appear as small yellow
streaks in the genital wings, and their early stages may be seen at the
extremities of their lobate ramifications. ‘The gonads are surrounded with a
layer of cells with elongate nuclei. . The developing ova oceur just within
this layer. They are of irregular amoeboid shape, and the germinal vesicle
is already well marked. Alongside of the ova, and apparently arising from
the same peripheral epithelium, are other cells, the outlines of which are not
well marked, containing lenticular nuclei like those of the outer layer, and
numerous small globules. In the older parts of the gonads the globules
constitute a solid mass, sometimes with many scattered and disintegrated
nuclei, and, in still later stages, only a mass of globules, some small and
some comparatively large, the latter being most probably formed by con-
fluence of the smaller. No globules were seen in the body of the ovum,
which, when mature, is surrounded by a zone of clear tissue with radiating
lines. It appears to be a reasonable inference that the granules are produced
by modified germ-cells.

Though no such process of asexual reproduction has apparently been
observed in other members of the Hnteropneusta, the tendency of some
species to fragmentation and regeneration is well known, and, if their life-
histories could be followed out, it might be found that reproduction by fission
is not an unusual process of multiplication, and is associated with the great
abundance of.eosinophil globules, which otherwise seem so difficult to
account for.

The presence or absence of hepatic ceca, as well as the extent of their
development, has not been found of any great systematic value in the Entero-
pneusta, and this case may indicate that other species may at different times
assume different forms, in which these structures may be absent, or in various

stages of development as in P. proliferans.

Some ten years ago it was noted asa remarkable fact by Korschelt and
Heider that, while asexual reproduction was such a characteristic feature of
Cephalodiscus, it was entirely absent in the closely-allied Balanoglossus and
in Phoronis. Since that date it has been shown by Harmer that it is a
normal process in a species of Phoronis, so that this distinction between the
Pterobranchia, Enteropneusta, and Phoronidea disappears.

HEAD CAPSULE AND MOUTH-PARTS OF CHLOROPS TANIOPUS. 399

On the Morphology of the Head Capsule and Mouth-parts of Chlorops
temopus Meig. (Diptera). By J. G. H. Frew, M.Sc., Ministry of

' Agriculture Research Scholar. (From the Entomological Department,
Rothamsted Experimental Station, Harpenden.) (Communicated by

Dr. A. D. Ins, F.L.S.)
(With 7 ext figures.)
[Read 3rd May, 1923.]

TE following paper deals with a part of the anatomical work done in
connection with an investigation into the life-history and bionomics of the
Gout-fly of Barley (Chlorops teniopus) which is being carried on at Rot-
hamsted under the direction of Dr. A. D.Imms. Considerable attention has
recently been directed to the Dipterous head capsule owing to Peterson’s
valuable monograph on the subject (3), and as the results of the present
investigation seem to necessitate important modifications of some of the
homologies put forward by Peterson, it has been thought advisable to deal
with the morphology of the head capsule of Chlorops in a separate paper.
Peterson’s nomenclature is followed in all cases unless there is very strong
evidence that it is incorrect.

Toe TENTORIUM AND THE HpIcRANIAL SUTURE.

The location of the epicranial suture is of fundamental importance in
determining the homology of the regions of the head capsule, and the’
position of the invaginations of the arms of the tentorium is, in cases of
doubt, the safest guide to the position of the epicranial suture. The epicranial
suture is typically \-shaped. The stem represents the line of junction of the
paired selerites of the head, while between the arms lie the unpaired sclerites,
the frons, clypeus, and labrum. Except ina few forms (Peterson, p. 15) the
stem of the epicranial suture is wanting in Diptera. The tentorium consists
of an arrangement of chitinized rods and plate-like structures. These arise
from three pairs of invaginations on the head—the openings of the posterior,
anterior, and dorsal arms of the tentorium. The invaginations of the posterior
arms are situated one on each side of the occipital foramen and are joined by
a transverse chitinous bar, forming the body of the tentorium and dividing
the occipital foramen into dorsal and ventral halves. ‘“ The invaginations of
the anterior arms are usually associated with the lateral margins of the
clypeus, with one of the points of articulation of the mandibles, and frequently
with the ventral ends of the arms of the epicranial suture. The invaginations
of the dorsal arms are associated with tlie points of attachment of the antenne

400 MR. J. G.H. FREW : MORPHOLOGY OF THE HEAD CAPSULE

and near the dorsal portions of the arms of the epicranial suture ” (Peterson,
p. 26). According to Comstock and Kochi (1, p. 41), ‘‘ Each dorsal arm of
the tentorium arises from the side of the body of the tentorium between the
»nterior and posterior arms, and extends either to the front or to the margin
of the antennal sclerites.”’

If one examines the tentorium of Chlorops it is seen to consist of three
distinct chitinous rods on each side (figs. 1 and 2). One rod on each side
(p.a.) arises at the side of the occipital foramen and passes forwards along
the ventral surface of the head capsule ending at the point marked X in
figure 1, at the ventral corner of the ridge which forms the lateral
boundary of the oral depression in which the proboscis lies when retracted.
These two rods are obviously the posterior arms of the tentorium ; the question

Fire. 1.

Fig. 2.

1:0 mm

Caudal aspect of Head Capsule.

Omm
Cephalic aspect of Head Capsule.

of the position of the body of the tentorium is dealt with in connection with
the morphology of the caudal aspect of the head. From the point X a second
chitinous bar (a.a.) runs along each lateral margin of the oral depression
ending at the point marked Y in the figure. his bar is a thickening of the
mesial wall of a shallow slit-like depression (s.), which extends along each
lateral margin of the oral depression. From the point Y a third bar (d.a.)
on each side extends dorsally along the anterior suface of the head; the
dorsal end of each of these bars is strongly curved and almost encloses
the antennal base of its side in the manner shown in the figure. All three
arms of the tentorium are fused along their entire length with the head
capsule. The dorsal margin of the oral depression is marked by a strong
bar (p.), which unites the two halves of the tentorium.

AND MOUTH-PARTS OF CHLOROPS TANIOPUS. 401

It seems clear that the bars extending from the points Y to the antennal
bases are the dorsal arms of the tentorium, and their antennal ends are
probabiy to be regarded as their points of invagination. The bars on each
side uniting the points X and Y are here regarded as the anterior arms of
the tentorium, their dorsal ends (Y) being their true points of invagination.
As already mentioned, their invaginations are actually slit-like and extend
along their whole length. Peterson has described several Brachycera and
Cyclorrhapha Aschiza in which similar, though apparently not such pro-
nounced, slit-like invaginations run along the anterior arms of the epicranial
suture, and the occurrence of slit-like invaginations in Chlorops cannot,
therefore, be regarded as in any way exceptional. It thus seems clear that
the three arms of the tentorium can be recognized in Chlorops as in less

Fre. 3.

Diagram illustrating the nature of the tentoriai thickenings of Chlorops.
to} to) to}

specialized insects, though their relations to each other are undonbtedly very
different from the relations in more typical tentoria. Peterson, speaking of
generalized insects, says (p. 26): “The small dorsal arms unite with the larger
anterior arms, and these, in turn, join with the posterior arms.” Fig. 3A
represents this diagrammatically ; the points O and P represent respectively
the points of junction of the dorsal arms with the anterior arms, and of the
latter with the posterior arms. Fig. 3B represents diagrammatically the
condition found in Chlorops, and the following provisional explanation is
offered. In Chlorops, as in most species possessing a ptilinum, the rods of
the tentorium have become fused with tlhe head capsule. In the case of the
anterior arm this may be supposed to have resulted in the entire obliteration
of the portion of the rod extending from a.v. (its point of invagination) to O,

402 MR. J. G. H. FREW : MORPHOLOGY OF THE HEAD CAPSULE

so that these two points become coincident, the anterior arm now being
represented only by the portion between the points O and P. Although this
explanation seems quite plausible, an examination of Peterson’s extensive
series of figures has failed to reveal any morphological evidence in support
of it in the shape of intermediate forms, and in view of the extensive series
of forms studied by Peterson it seems unlikely that such evidence is available.

Peterson (p. 16), although he says that the anterior arms of the epicranial
suture are not present in any Dipteron possessing a ptilinum, considers,
nevertheless, that “there is every reason to believe that the tentorial thick-
enings mark the course of the suture.” If one accepts this view the region
of the head lying between the dorsal arms of the tentorial thickenings in
Chlorops must be the frons; but owing to the curved dorsal ends of the
thickenings the antennz arise, quite definitely, on this included region.
Such a position for the antennz does not occur, so far as I know, in any
insect, and one must, therefore, conclude that the region lying between the
dorsal arms of the tentorial thickening is not the frons (or fronto-clypeus).

I consider that the tentorial thickenings between the points X and Y on
each side mark the lower portions of the arms of the epicranial suture, the
shallow invaginations in which the thickenings lie being probably the actual
sutures which have remained permanently open. As mentioned above, the
dorsal edge of the oral depression is formed by a transverse chitinous bar
uniting the two halves of the tentorium, and I consider that this represents
the dorsal and median region of the arms of the epicranial suture. ‘he
following passage from Peterson (p. 28) gives, I think, considerable support
to this view. Speaking of the invaginations of the anterior arms of the
tentorium in Tabanus he says :—‘ The invaginations on each lateral half of
the head are joined together by the arms of the epicranial suture and
resemble the hypothetical type.” The points Y have already been indicated
as probably representing the points of invagination of the anterior arms of
the tentorium.

I consider, therefore, that the dorsal and lateral margins of the oral
depression coincide with the anterior arms of the epicranial suture. Accord-
ing to this view the antennze arise on the vertex, a position which they
occupy in many Orthorrhapha, and one must look for the frons and elypeus
among the structures lying in the oral depression. On the dorsal surface of
the rostrum of the proboscis there is a plate which is the superficial region
of a chitinous structure usually termed the fulcrum. Lowne (p. 134)
regarded this as the clypeus, and Dr. Imms tells me that he is expressing the
same view in a forthcoming publication. With this view I agree, with the
small reservation that, as according to Peterson (p. 17) the frons and clypeus
are separated by a doubtful suture in only a few Orthorrhapha* and are
entirely fused in the majority, it is possible that the sclerite in question

* He also regards the frons and clypeus as fused in the Cyclorrhapha, but here the region
which he calls fronto-clypeus is, in my opinion, part of the vertex.

AND MOUTH-PARTS OF CHLOROPS TANIOPUS. 403

_represents the fronto-clypeus. If it only represents the clypeus, the frons is
represented by the intervening membrane uniting it to the dorsal edge of the
oral depression.

Only in Aycetophila among the Dipterous types studied by Peterson is the
stem of the epicranial suture complete. Some forms (Rhabdophaga, Mycetobia,
Tabanus) ‘show depressions or thickenings along the meson. These marks
may have no significance” (p.15). In Chlorops there is a very distinct
internal ridge running along the meson from the median ocellus to the
median point of the dorsal edge of the frontal suture.

TormM# AND FRONTO-CLYPEUS.

In the base of the rostrum there is a somewhat complex chitinous structure
usually termed the fulcrum (fig. 4), part of which is internal and is formed
by the chitinized basipharynx, and part of which is superficial and is, as
stated above, the clypeus or fronto-clypeus. The internal portion will be

SK ee ee

= os a= =

0:5 mm

Ventro-lateral view of basipharynx, torme, and clypeus.

considered later ; it consists of dorsal and ventral plates enclosing between
them the pharyngeal cavity. A vertical chitinous plate (¢) on each side unites
one side of the superficial plate (clypeus) to the corresponding side of the
basipharynx. Peterson regards these, together with the superficial plate
which I consider to be the clypeus (or fronto-clypeus), as the Torme.

“<The tormee in generalized insects are chitinized pieces which belong to
the lateral portions of the epipharynx in the region of the clypeo-labral
‘suture and connect with the clypeus or labrum at the lateral ends of the
suture.... The torme of generalized Diptera also connect with the inner
surface of the ventral portion of the fronto-clypeus” (Peterson, p. 19). The
basipharynx is composed of united epipharynx and hypopharynx. Thus, by
accepting the view that the superficial plate is the clypeus or fronto-clypeus,

404. MR. J. G. H. FREW: MORPHOLOGY OF THE HEAD CAPSULE

it will be seen that the lateral vertical plates have absolutely typical relation-
ships as tormee. According to Peterson’s view that the lateral plates and the
superficial plate represent torme alone, the tormee of the Cyclorrhapha are
strikingly different in their relationships from the tormeze of generalized
insects or of the Orthorrhapha; a difference which he hardly attempts to
explain. ,

As in the remainder of this paper I shall have no cause to disagree with
Peterson’s views, it may be useful to summarize here the points of difference.
In Chlorops I consider that the tentorium clearly consists of the three typical
arms; this may, however, be a special case, as it certainly appears to be
judging from Peterson’s figures, and not general to all Cyclorrhapha. The
following points, however, are almost certainly applicable to all Cyclor-
rhapha :—

1. The position of the arms of the epicranial suture is marked by the
dorsal and lateral borders of the oral depression.

2. All regions of the head lying dorsal and lateral to the oral depression
are derived from the paired sclerites of the head, and the frons and
clypeus must lie within the oral depression.

3. The antenne arise on the vertex.

. The superficial plate of the fulcrum is the elypeus or fronto-clypeus.

5. The torme are the chitinized plates joining the sides of the clypeus
to the sides of the chitinized basipharynx.

is

It cannot be claimed that absolutely rigid proof of any one of the above
statements has been given. The evidence in support of them seems to me,
however, to be very strong, and their acceptance appears to afford a very
much clearer explanation of the nature of the facial aspect of the Cyclor-
rhaphous head capsule than does Peterson’s homology, and, moreover,
brings the Cyclorrhaphous head capsule more into line with that of the
Orthorrhapha or of generalized insects.

Yhe Ptilinum shows no marked peculiarities. It lies within the head
capsule close against its anterior wall. The frontal suture ( fs.) lies just
dorsal to the antennal bases.

The Labrum (figs. 1, 5, & 6) is a rather heavily chitinized triangular
sclerite lying along the dorsal surface of the mediproboscis, its base being
continuous with the dorsal membrane of the distal end of the basiproboscis,
so that it is separated from the clypeus by a considerable extent of membrane
a condition which is usual, but not universal, in Diptera (Peterson, pp. 20-
21). The vevtral surface of the labrum is longitudinally grooved for the

reception of the epipharynx, which fits tightly within the groove and cannot
be removed without damaging the labrum. The epipharynx does not extend
quite to the distal end of the labrum, and just beyond its termination the
labral groove bears two small conical papillee, one on each side, which have

AND MOUTH-PARTS OF CHLOROPS TANIOPUS. 405

the appearance of being sense-organs. The dorsal surface of the Jabrum is
strongly convex from side to side ; its base articulates with the distal ends of
the stipites (fig. 6). The labrum, together with the epipharynx, is easily
lifted from the surface of the proboscis and rotated dorsally, its base hinging
on the stipites.

The Vertex “is interpreted as including all the cephalic and dorsal
aspects of the epicranium except the front.... The region of the vertex
ventrad and mesad of each compound eye is a gena” (Peterson, pp. 21-22).
It is only necessary to remark that what Peterson calls the front is, in my
opinion, part of the vertex. The term “front” as applied to Cyclorrhapha
has no true morphological significance, but the region defined by the term is
so well understood, and the term is so constantly used in taxonomical writings
on Diptera, that it would, I think, be a mistake to propose any new name for
this region.

The TOsphnnne Byes and Ocelli.—The only noteworthy feature concerning
the eyes is the presence round their margins of a well-marked, ‘Thome
incomplete, ocular sclerite (figs. 1 & 2,0.s.). This ring sclerite is incomplete
anteriorly ; it bears two small peg-like projections, an antero-ventral (a.p.)
and a postero-dorsal (p.p.). The three ocelli are situated on the vertex some
distance dorsal to the frontal suture, and lie on a raised and dark-coloured
triangular area (fig. 2, 0.t.), which is continued forwards in front of the
median ocellus as a dark-coloured internal chitinous ridge running along the
median line of the vertex and ending at the dorsal margin of the frontal
suture.

Occreur AND PosTtGENa.

Owing to the absence of sutures on the caudal aspect of the head (fig. 2
it is not possible to define with certainty the limits of the occiput and ses
genx, and accordingly Peterson’s view is followed that all regions of the
caudal aspect lying dorsal to a transverse line drawn through the middle of the
occipital foramen belong to the occiput, while the areas ventral to this line
and lateral to the mesial membranous area are the posteenze. The dorsal half
of the foramen is strengthened by a thickening of the occiput (p.o.) which
provides articulation for the neck sclerites. I follow Peterson in calling this
the Paroeciput. It is not marked off from the occiput by a secondary
suture, as is frequently the case in Diptera. On each side a chitinous peg
(b.t.) projects mesially across the foramen, the two almost meeting in the
median line and dividing the foramen into dorsal and ventral halves. These
certainly appear to me to be processes of the ventral ends of the parocciput,
but Peterson has named apparently identical structures (e.g. in Chloropisca
glabra, fig. 132) as the body of the tentorium, and as he has examined a
large series of forms and has therefore had the opportunity of tracing the
yariation of the body of the tentorium in Diptera, I have decided to follow

406 MR. J. G. H. FREW : MORPHOLOGY OF THE HEAD CAPSULE

his nomenclature in this matter. On each side the parocciput bears an
articular surface of somewhat complicated nature (art.) with which the
anterior end of the chief neck sclerite on each side articulates.

The ventral half of the rim of the occipital foramen is thickened (pa.p.),
and the posterior arms of the tentorium appear to arise from this thickened
rim. This is the Parapost@enal thickening (Peterson, p. 24).

Two long chitinized ridges (7.) occur on the inner surface of the occiput,
one on each side, arising from the dorso-lateral region of the parocciput and
extending dorso-laterally towards the eyes. A median thickening, which is
present in many Cyclorrhapha in addition to the above,is not found in
Chlorops.

Fre, 5.

la

~ ;

d.ph th

1 ——+—-___}
O5 min

Lateral view of the proboscis.

-— so

The Antenne consist of three basal joints and an arista arising from the
third joint and itself three-jointed. The second joint of the antenna has a
cone-shaped apex which fits into the base of the third joint. The basal joint
is a ring-shaped sclerite and is united to the second joint by a membranous
region of considerable extent. There is a relatively large pit on the under
surface of the thitd joint which is probably a sense-organ, but which has not
yet been closely examined. The arista is minutely pilose.

Mandibles are absent.

The Labium forms the greater part of the proboscis. The following is a
brief résumé of Peterson’s views on the homology of the Dipterous pro-
boscis :—The membranous basiproboscis is made up of submentum, mentum,

AND MOUTH-PARTS OF CHLOROPS TANIOPUS. 407

and’ the cardines and stipites of the maxiile, but the limits of the sclerites
cannot be distinguished. The mediproboscis and the distiproboscis con-
stitute the ligula. The labellie are the paraglossse, and between them are two
»membranous lobes which are the glossze ~.

Tn the membranous basiproboscis of Chlorops only the stipites (figs. 1, 5,
6, st.) can be distinguished as separate sclerites. In the majority of
Cyclorrhapha a distinct galea is found attached to the ventral end of each
stipes, part of it appearing on the surface of the proboscis (the stipites lying

Fie. 6.

0-2 mm

Ventral yiew of labrum, epipharynx, and associated parts.

entirely within the proboscis). “In the Calypteratze and some of the
Acalypteratee the galea articulates with the proximal end of the labrum and
is more or less firmly connected with the same. The ectal exposure of the
galea is very small in these forms” (Peterson, p. 41). In Chlorops I have
been unable to differentiate the gales, and they have probably fused with the

* Other views on the morphology of the proboscis are given by Wesche, W., ‘The
Labial and Maxillary Palps in Diptera” (Trans. Linn. Soc., Zool. vol. ix. 1903-7), and by
Crampton, G. ©., “The Sclerites of the Head, and the Mouth-parts of certain Immature and

-Adult Insects” (Ann. Ent. Soc. Amer, vol. xiv., 1921). The former author considers that
the palpi of Muscidze are labial and not maxillary. Crampton considers that the labella of
Diptera are labial palps.

408 MR. J. G. H. FREW : MORPHOLOGY OF THE HEAD CAPSULE

ventral end of the stipites, which, as already mentioned, articulate with the
base of the labrum. Although very near the surface I could not definitely
determine that any part of these ventral ends was actually superficial. The
stipites are flexible, and probably springy rods. When the proboscis is
extended they are practically straight, but they are considerably curved
when the proboscis is retracted. They probably assist in the extension of
the proboscis by pressing on the base of the labrum ; this point, however,
requires further investigation. According to Peterson the areas mesad
of the stipites are to be considered as developed from the submentum and
mentum, while the areas lateral to them are formed from the maxille.

The Theca (fig. 5, th.) is a broad curved sclerite lying on the caudal] aspect
of the mediproboscis. It is not very heavily chitinized, nor are its limits
very sharply defined. On its dorsal (anterior) aspect the mediproboscis is
channelled by a longitudinal groove whose walls are rather more heavily
chitinized than the general membrane of the mediproboscis ; the hypopharynx
lies in this groove.

Fie. 7.

Diagram of the relationship of the epipharynx and
the hypopharynx. (Ventral view.)

The Labellw (Paraglossee) are shown in fig. 5. A detailed account of their
structure is unnecessary as they are essentially similar to those of Musca.
Hach labella forms half of the stomal disc, the two being united mesially by
a flexible membrane, and each bears three pseudotrachex. The two outer
pseudotraches of each side unite to form a short common duet which
opens into a short but wide groove at the proximal end of the stomal
disc, this groove being continuous proximally with the groove of the hypo-
pharynx. The mesial pseudotrachea of each side opens separately into the
short common groove. The glossee cannot be distinguished. According to
Peterson this is also the case in Chloropisca, where “it is impossible to dif-
ferentiate the glossee from the chitinized groove of the mediproboscis and the
proximal ends of the pseudotrachez ” (p. 49).

In the majority of Diptera each paraglossa is supported by a triradiate
chitinized rod—the Furea—lying on its lateral and caudal aspects, and in
the majority of Brachycera and Cyclorrhapha there is a small sclerite—
Sigma—situated between each furea and the ventral margin of the theca.
Neither of these sclerites is present in Chlorops, but this is not very surprising,
as Peterson mentions a number of Diptera, Chloropisca among them, in

os

AND MOUTH-PARTS OF CHLOROPS THNIOPUS. 409

which he has been unable to distinguish them. ‘“ Kappa” is also absent, but
Peterson has only found this sclerite in Tabanus, Tipula, and Bittacomorpha,
where it lies ““embedded in the membrane laterad of the ventral ends of the

theca.”
The Mawillary Palps (m.p.) are one-jointed and present no peculiarities.

HPrPHARYNX AND HyPoOPHARYNX.

The epipharynx (ep., figs. 1, 6, 7) lies in the groove on the under side of
the labrum and is in the form of a gutter, semicircular in cross-section and
opening ventrally. The hypopharynx (hp.) is a similar but rather more deli-
eate structure lying in the groove on the dorsal surface of the mediproboscis,
its opening facing dorsally so that, when epipharynx and hypopharynx are in
contact, 1 closed hollow cylinder of chitinis formed. At about the point of
junction of mediproboscis and distiproboscis the epipharynx and hypopharynx
unite to form a short closed cylindrical tube, the distipharynx (d.ph.), whose
proximal end articulates with the distal end of the basipharynx (b.ph.), the
cavities of the two being continuous. Although it is certain that the short
distipharynx consists of united epi- and hypopharynx, it has the appearance
of being formed by the former alone, whichis obviously a direct continuation
of it, whereas the hypopharynx is separated from the distipharynx by a
narrow membranous area acting as a hinge, and is easily broken off at this
point.

Just distal to its junction with the distipharynx the epipharynx has a
rather deep groove (e.g., fig. 6) on its surface. The hypopharynx apparently
lies quite freely in the groove on the mediproboscis, from which it is easily
detached except at its distal end, where the two are united. As the labrum
is slightly, and the epipharynx considerably, shorter than the hypopharynx, -
there is a short distal region of the latter which cannot be converted into a
closed groove by the apposition of the labrum or epipharynx. The distal end
of the hypopharyngeal groove is, however, deep, and a study of serial
sections has afforded some evidence of the possibility of the lips of the groove
being brought together to form a closed channel, such a closure being
probably effected by an increase of turgidity inthe mediproboscis due to the
pumping in of body-fluid when the proboscis is extended for feeding.

BastPHARYNX. (Higs. 4 & 5, b.ph.)

The form of the basipharynx is sufficiently well shown in the various
figures. It is an organ of suction. Muscles arise from the dorsal surface of
the basiproboscis and are inserted into the dorsal wall of the basipharynx,
which they raise by their contraction, thus increasing the capacity of the
pharynx. Posteriorly the basipharynx ends in two blunt cornua (cu.).
Peterson interprets the basipharynx as consisting of united epipharynx and
hypopharynx.

LINN, JOURN.—ZOOLOGY, VOL. XXXV. 30

410

HAD CAPSULE

REFERENCES.

AND MOUTH-PARTS OF CHLOROPS

(1) Comsroor, J. H., & Kovnz, C. (1902).

(2) Lownn, B. T.

(3) Permrson, A. (1916).

Insects.”
(1890-95).

Development of the Blow-Fly.”

TENIOPUS.

“The Skeleton of the Head of

The American Naturalist, vol. xxxvi.
‘Anatomy, Physiology, Morphology, and

London.

“ The Head Capsule and Mouth-parts of Diptera.”

Illinois Biological Monographs, vol. iii. No. 2.

Index lettering to Teat-figures.

ad., d.d., and p.a, anterior, dorsal, and pos-

terior arms of the tentorium.

at, dt, and pz. their points of invagi-

ap.

art,
b.ph.
b.t.

cl.

cu.
d.ph.
EE.
e.9.
ep.
fs.
hp.
la,
1b.
Mm.

mp.

nation.
antero-ventral process of ocular scle-
rite.
articular surface for neck sclerite.
basipharynx.
body of the tentorium.
clypeus.
cornu of basipharynx.
distipharynx.
compound eye.
epipharyngeal groove.
epipharynx.
frontal suture.
hypopharynx.
labellee.
labrum.
membrane separating
from distipharyny.
maxillary palp.

hy popharynx -

|

N.S.

O.

neck sclerite.
point of junction of dorsal and an-
terior arms of the tentorium.

of. occipital foramen.

0.8.
0.8.

IE,

ocular sclerite.

ocellar triangle.

point of junction of anterior and pos-
terior arms of the tentorium,

. thickening of dorsal margin of oral

depression.

. parapostgenal thickening.
. parocciput.
. postero-dorsal process of ocular scle-

rite.

, ridge on occiput.
. slit-like invagination containing an-

terior arm of tentorium.

. Sense (?)-organ on labrum,
st. stipes.
. torma.
. theca.

CRUSTACEAN PLANKTON OF THE ENGLISH LAKE DISTRICT. Al1

The Crustacean Plankton of the Hnglish Lake District.
By Rosert Gurney, M.A., F.L.S.

(PLaty 23 and 3 Text-ficures.)

[Read 3rd May, 1923.]

Tax published information with regard to the Crustacea of the Lake District
is surprisingly scanty. Mr. Scourfield has included all available records in
his Synopsis of British Hntomostraca (1903-4), but naturally dealt only with
the district as a whole, without comparison of the fauna of the different
lakes. Miss Pratt’s paper on the Hntomostraca of Lake Bassenthwaite is
the only attempt that has been made to give a comprehensive account of the
Crustacea of any single lake. The phytoplankton, on the other hand, is
fairly well-known asa result of the work of Messrs. West and of Dr. Pearsall.
The following account of the Crustacea of the district lays no claim to
being exhaustive or even to deal with many of the points of special interest,
but at Jeast it may serve as a groundwork for comparison with other lake
areas and to draw attention to questions on which further work is required.
The material at my disposal consists chiefly of a series of plankton samples
taken by Dr. W. H. Pearsall during 1921 and 1922 for the purpose of a
study of the phytoplankton, and I wish to express my thanks to him for the
loan of these collections and for the valuable information which he has
readily given to me during the course of the work. Ihave also myself:
visited the district and have made collections in a few of the lakes. During
the autumn of 1922 I spent a week in the western part of the district
specially investigating the Crustacea of Hnnerdale, Wastwater, and Coniston.
While it is probable that the collections at my disposal give a fairly com-
plete picture of the composition of the crustacean plankton of the various
lakes, it is unfortunately true that the methods which are suitable for the
collection and preservation of phytoplankton are by no means so for the
Crustacea ; and for this reason, though Dr. Pearsall’s collections extend over
a large part of two years, it is not possible to deduce from them any reliable
conclusions as to the seasonal distribution and variation of the Hntomostraca.
Undoubtedly much further work remains to be done with regard to these
two questions, and also in connection with the vertical distribution of the
species. It seems also to be probable that the distribution of the Crustacea
may not be uniform in some of the larger lakes, and that such lakes should
not, therefore, be dealt with as homogeneous units. For instance, the com-
position of the plankton in the deep part of Ennerdale is not the same as
30*

412 MR. R, GURNEY ON THE GRUSTACEAN PLANKTON

that of the shallow western part, and the same may be the case with Ulls-
water, Windermere, and, perhaps, Coniston. Holopedium gibberum was not
found in any of Dr. Pearsall’s collections from Windermere, but it was taken
there many years ago by Beck, and again in 1912 by Mr. P. A. Buxton.
It may be that it is a permanent inhabitant of the lake, but it is restricted to
certain parts of it.

Fic. 1.

Bassenthwaite L.
63

a

ss a

|, “BORROWDALE*\VOLCANIC

oe?

Ganiston Ly

4

SILURIAN FLAGS &
a B y

: 5 Skidd
Silurian series SIateel A

Volcanic Series

Map of the English Lake District, with section showing the relation of
scenery and gradients to the underlying rocks.

(By permission of Dr. W. H. Pearsall and the Council of the Royal Society.)

I have thought it well to add notes on some of the species composing the
plankton, and on the fauna of some of the high-lying lakelets or tarns, in
order to give a more complete picture of the crustacean fauna than would he
conveyed by a statement of the composition of the plankton alone.

OF THE ENGLISH LAKE DISTRICT. 416

With the physical conditions of the district and its lakes it is not my
business to deal. The foundation of our knowledge of these lakes was laid
in the survey of Dr. H. R. Mill (1895), and Dr. Pearsall has, in his papers
on the aquatic flora and phytoplankton, added greatly to our knowledge of
the post-Glacial history and present conditions of these lakes. The accom-
panying map clearly shows relations of the lakes, which occupy valleys
radiating from a common centre.

Tan PLANK Ton.—Comparison of the various Lakes.

One of the chief results of Dr. Pearsall’s work is that “the stage of
evolution of the lake-basin must be regarded as being the fundamental factor
affecting the distribution of the phytoplankton, since it is upon this factor
that the character of the waters depends” (1921, p. 279), and he has divided
the lakes into groups on the basis of the amount of silting of their beds.
These groups are :—

J. Primitive Lakes.
Wastwater, Hnnerdale, Buttermere, and Crummock.
II. Intermediate Lakes.
Hawes Water, Derwentwater, Bassenthwaite.
III. Evolved Lakes.
Coniston, Windermere, Ullswater.
IV. Most evolved Lakes.
Hsthwaite (and Grasmere).

The phytoplankton corresponds fairly well with this grouping, the lakes
of the first group containing a Desmid plankton, while in the remainder
there is a large proportion of Diatoms. But Desmids are also frequent in
Group IL., while Myxophycez form an increasing component of the plankton
of Groups ILL. aid IV., and are at times dominant in Esthwaite. As
Dr. Pearsall intends to deal in detail with the plankton as a whole in relation
to its environment, I shall do no more than attempt to show how far the
distribution of Entomostraca conforms to this grouping.

Any grouping of these lakes on the basis of their Crustacea is by no means
easy, since, with the exception of Buttermere and Crummock, which lie in
the same valley and undoubtedly at one time formed a single lake, each lake
has its own individual character—so much so that it would probably always
be possible to identify the lake from which any given representative sample
of plankton was taken.

In Table 1, I have arranged the lakes in the groups adopted by Dr.
Pearsall, Group I. being the most primitive and least silted, and from this
table the composition of the Crustacean plankton can be seen at a glance.
Every plankton species taken is recorded in the table, but those which
occurred only singly in one collection or were evidently not regular con-
stituents of the plankton are distinguished by a cross enclosed in a circle.

414 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

Tasie 1, showing composition of Plankton. The Lakes are arranged in
Groups as in Pearsall, 1921, p. 276. Crosses enciosed in circles indicate
that the species is occasionally found, but is not a regular member of
the plankton. These species are not included in the total number's.

| . | ( |
| Group I. | Group LI. Group UI. ||GroupIV.
| [eee we | een
| oe | o | &
| Sele Slelelalalel/s ie |
| Ee aS a g || » i) | = g ® « S =
epee |S Peis | BS) Bose eB ls
Seales lelelaleatlaelel2ileles
FISIS/ol/FlS ayo lel oie is
ee a | 8) roan} 12,
Stda erystallina ........ gol GS Mibe ah oe ee | ae | ar + | |
Diaphanosoma brachyurum) @B any } ip | + +) + |
| Elon GARAGE boos) ce | 44) os | co lll oo | so ll oo fl eo | I & ar |
|| |
| Daphnia hyalina s. str. fp] + |
a UGOVISARS Shoood a eal ay epeil Ketseces vera ata foe 0-9 +
A GHZ 5.0.606.5.4 0 Bl Teton tora Na ol hose tds sect: Sea niesre sce |
Re - | pulchella l | |
| Ceriodaphnia qoadiengetal 2.) 28 es fies ih ee ee [ss ay) sp ats
Bosmina longirostris...... at a9 |} ox ap || ar |
|» » obtusirostris ....) +] +] .. | + AG | fe | ao ae |) oe
Polyphemus pediculus .....@|..\--|@ |) @ +) 4] .. SF
| Bythotrephes longimanus ..| + | + | qe ae sei se iy ae | Se] SP lrco ll Se
Leptodora Iindtt ........ = | + ists + | + + }+
Limnocalanus macrwus ..) .. | + | l | |
Diaptomus laticeps ......| .. |. Ine ae | + |
4 gracilis ...... +/t}/4)+) +)4+)4 7 +) 4+) 4447+
Cyclops leuckarti ........ Bea eam hewel Liat NIE +/+ ..) 4+ --
» — AbYSSOTUM «ove ees So og Pee | oe IP se | +)+)+ +
| Number of species Se he | 6 BN Oe 9/6 || 10) 8

It should be noted further that collections have oly been taken from the
shore of Buttermere, and that I have only a single collection from Grasmere;
taken in September 1912. Species which are known to belong to the littoral
fauna, such as Alonopsis elongata, are omitted even when they have some-
times been taken in the plankton. Chydorus sphericus, which is often
limnetic in central European lakes, is not found in the plankton of this
district.

It will be seen at once that the Crustacea do not conform to Dr. Pearsall’s
grouping, and indeed it is diffienlt to trace definite relationships at all.

OF THE ENGLISH LAKE DISTRICT. 415

Group I. of the lakes of the deep and rocky type agree in the limited number
of species and the absence of species characteristic of shallow and warm
waters, and in their predominant zooplankton, while Hnnerdale and Wastwater
together differ from Buttermere and Crummock in the entire absence of
Daphnia. Buttermere and Crummock are characterized by the dominance
in their plankton of the typical form of Daphnia hyalina. But, although
Ennerdale and Wastwater are so alike in physical characters, they differ
markedly in their plankton, not only in the species found, but in their
relative abundance. Hnnerdale possesses two striking species, /olopedium
gibberum and Limnocalanus macrurus, which are absent from Wastwater,
but, on the other hand, lacks both Leptodora kindti and Cyclops abyssorum,
hoth of which are common in Wastwater. Further, Bosmina obtusirostris 1s
abundant in Wastwater, while it is an insignificant component of the plankton
of Ennerdale. It cannot, in fact, be said that the Crustacea indicate any
greater relationship between these two lakes than between either of them
and certain lakes in other groups such as Hawes Water, Ullswater, or
Coniston. In these two lakes Dr. Pearsall’s grouping is put sharply to the
test, for, so far as physical conditions go, two lakes could hardly be more
alike than these, and their phytoplankton also is very similar.

The same difficulty is met with in the other groups, though it is true that
Esthwaite bas characters which distinguish it very clearly. It has a very
varied plankton, which contains the “ warm-water” or lowland Crustacea
Diaphanosoma brachyurum, Ceriodaphnia, and Cyclops leuckarti; but its chief
claim to distinction is the presence (in company with B. obtusirostris) of
Bosmina longirostris, which is a distinctly southern form. Grasmere, except
for containing Holopedium, is very similar to Esthwaite.

In Group II. it is clear that Hawes Water cannot be classed with Derwent-
water and Bassenthwaite with regard to their Crustacea. The presence of
the northern species iaptomus laticeps places it in a class apart, its elevation
of 694 feet giving it almost a position among the mountain tarns. It differs
also from the other two, possibly for the same reason, in the different race of
Daphnia inhabiting it, and also in the absence of Sida, Leptodora, and
Cyclops leuckarti.

Derwentwater and Bassenthwaite also differ considerably, though forming
part of one drainage system, for the latter lacks any species of Daphnia or of
Bosmina, being unique among the lakes in this respect. It is true that both
have been recorded from it, and that a single specimen of Bosmina obtusi-
rostris occurred in one sample examined by me; but it is evident that, at the
present time, neither can properly be included in its fauna. The great
abundance of Diaphanosoma in the plankton of both lakes is a striking
feature in common, and the plankton of Bassenthwaite may perhaps be
regarded as that of Derwentwater, in which certain elements have been
suppressed.

416 : MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

Dr. Pearsall suggests that the impoverishment of the plankton of Bussen-
thwaite may be due to the silt trom lead mines washed into the lake during
the past fifteen or twenty years. It is quite possible that to this cause is due
the difference between the present plankton and the list given by Miss Pratt
in 1898.

Group III. is also by no means a natural group from the point of view of
the Crustacea, although the fact that it is only in the three lakes—Coniston,
Windermere, and Uliswater—that Daphnia of the galeata-form occurs, may
indicate a similarity of conditions. The presence of Holopedium, either
regularly or occasionally, in Ullswater and Windermere may also be regarded
as evidence of similarity, though the status of the species in the district is
rather doubtful. It seems to me that, if any grouping is to be attempted on
the basis of the crustacean plankton, Ullswater and Hawes Water should be
included in the series of primitive lakes, while Windermere, with its )ia-
phanosoma, Ceriodaphnia, and Cyclops leuckarti, belongs more nearly to the
Hsthwaite-Grasmere series. Coniston, with its dominant crustacean plankton,
comes near the primitive type, while Derwentwater and Bassenthwaite lead
on towards the more evolved type of Windermere and Nsthwaite.

lt is impossible, without greatly extended kuowledge of the food require-
ments of the Mntomostraca, to determine what are the factors controlling
their distributions in an area such as this. The evidence is conflicting.
Birge (1898, p. 353) states that aphnia feed readily on filamentous Diatoms
(e.g. Melosira) and also on Aphamzomenon and Anabena, while Diaptomus
prefers Anabena and Aphanizomenon to Diatoms. On the other hand, more
recent investigations lead to the supposition that but little correspondence
will be found between the distribution of the Entomostraeca and the net
phytoplankton, since the Entomostraca appear mainly to feed upon the
minute nannoplankton, which is not taken in the plankton net. Woltereck
(1908) stated that the net plankton is far too large to be taken in by Daphnia,
their food consisting partly of the finest detritus, but mainly of the nanno-
plankton. he latter he found to be more abundant in the Lunzer Obersee,
where Daphnia was more flourishing, than in the Untersee, and attributed
this to the greater organic content of the water. ‘This result was confirmed
by experiment, since it was found possible to maintain limvetic Daphnias in
health on a diet of a pure culture of Chlorella.

Naumann * concluded that it was not the Algze themselves but the detritus
produced from them, and the smaller Flagellates, which provided the food
for the Crustacea, and that the dust-fine organic detritus or peritripton, which
may be introduced into the jake from without, may be of the utmost
importance in the lime-free Paleozoic regions. In such lakes the zoo-
plankton is greatly in excess of the phytoplankton. ‘The lakes of the

* T have not been able to consult Naumann’s paper, but rely on a review of it in Arch.
f, Hydrob. xii. 1920, p. 835.

OF THE ENGLISH LAKE DISTRICT. 417

lowlands of north and middle Europe are, on the other hand, rich in electro-
lytes, the phytoplankton is abundant, and the zooplankton feeds on the
detritus derived from it.

Considerations such as these offer a possible clue to the understanding of
the composition of the zooplankton, but leave untouched the problem of the
distribution of the predaceous Cladocera such as Leptodora.

The factor which appears chiefly to influence the distribution of the
Crustacea in this district is depth of water. A fair idea of the relative
depth of the lakes is given by the following table, showing the percentages
of the area covered by depths of 50 or 100 feet.

TasLe 2.—Depths of the Lakes. Figures from H. R. Mill, 1895.

Maximum Percentage of area covered by
depth cr A ane
| in feet. | 0-50 feet. | 0-100 feet. over 100 feet. |
eer: |

| Wastwater .... 958 | 23 39 61
Windermere.... 219 42 | 63°6 | 364
Ullswater ...... 205 352 | 64 | 36
Coniston ...... 184 371 | 60°8 | 39:2
Ennerdale...... | 148 52:6 Z | 678 | 3272
Crummock...... 144 | 27 47 53
Hawes Water .. 103, 64 | 98:3 | ey
Buttermere .... 94 | 378 | 100 | —
Derwentwater .. 12 93-9 | 100 —
Bassenthwaite .. 70 94-1 | 100 —

i

In some cases this does not give a true impression, since, in the case of
Hnnerdale, Windermere, and Ullswater, there are deep and shallow basins.
Mill divides the lakes into a deep and a shallow class, the latter including
only Derwentwater and Bassenthwaite.

Now, the species of plankton Crustacea may be roughly arranged as Cold-
water, Hurytherm, and Warm-water species, and it is found that the
proportions in which these classes occur in the lakes vary in fairly close
accordance with the depth. It should be noted that, in the following table,
the lakes are arranged according to the percentage of depth over 100 feet,
but Hnnerdale is placed next to Wastwater, in view of the configuration of
its eastern trough. Windermere and Hawes Water are more or less inter-
mediate between the deep and shallow lakes in the character of the plankton,

418 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

but the former should be specially studied with a view to seeing if, as is
probably the case, the plankton of the deep and the shallow basin differs in
composition.

TaBLE 3.—Composition of the Plankton.

| Cold-water Species. | Hurytherm Species, | Warm-water Species, |
Wastwater ....| 2 3 =
Ennerdale ...... | 3 2 —
Yrummock .... 3 | 2 | =
@onistoniyae 2 4 | =

Windermere ... .| 2 4 3 |
Ullswater ...... 3 3 ay
Hawes Water .. 3 3 1
Derwentwater .. 1 5 3
Bassenthwaite .. — 4 3
Hsthwaite...... 1 4 5

The Northern Element in the Plankton.

The general character of the plankton is very distinct from the Central
European and Baltic types and resembles that of Scotland. Characteristic
of it is the absence of Daphnias of the cucullata-group and of Bosmina
longirostris from all the lakes except Esthwaite and Grasmere, and of
Ceriodaphnia from all but Windermere and Hsthwaite.

The general abundance of Bosmina obtusirostris in itself marks the
plankton as of the northern type, but the following species are also
charaeteristic of northern or arctic lakes :—

flolopedium gibberum.
Daphnia hyalina s. str.
Bythotrephes longimanus
Polyphemus pediculus.
Diaptomus laticeps.
Limnocalanus macrurus.
Cyclops abyssorum.

Diaphanosoma brachyurum may be looked upon as belonging to a southern,
warm-water, genus, but it is itself so widely distributed throughout Hurope,
in cold as well as in warm waters, that little importance can be attached to

OF THE ENGLISH LAKE DISTRICT. 419

it in this connection. It is found only in the summer months and has an
extremely short life-cycle, a fact which has probably largely assisted it to
accommodate itself to a great variety of waters.

Leptodora kindti, although in deep lakes it may have a preference for the
lower or colder strata, is by no means restricted to deep lakes, nor is it
characteristic of northern regions. It may rather be regarded as a southern
element in the fauna.

Although the crustacean plankton of the Lake District has a distinct
northern facies, the absence from it of Diaptomus laciniatus and D. wierzejski,
which are not uncommon in Scotland, is noticeable. The latter isa eurytherm
species of remarkably wide range, but characteristic of pools rather than of
lakes, but the former is a definitely arctic-alpine species. Diaptomus laticeps
is a member of the group of northern Diaptomus which is widely distributed
in Scotland, but only occurs in this district in Hawes Water and Goats
Water. It seems probable that it may be a recent immigrant.

From the point of view of the Crustacea the Lake District plankton
certainly belongs to the Scottish Highland type, with the addition of a few
southern forms.

It must be admitted that any Deon as to whether the plankton of any
lake or district has a northern or southern facies is of doubtful value in the
present state of our knowledge of the means of distribution and required
conditions of existence of the different species. Not only is the whole
question of the relation of the present fauna to the effects of the Glacial
period far from being clear, but we have little means of knowing to what
extent the fauna was able to survive that period in its original habitat. In
the case of the Lake District it seems fairly certain that the present lake-.
basins were filled with ice, and it is probable that the whole district was
covered with a mantle of ice, so that the present aquatic fauna is entirely of
post-Glacial origin. It has been re-colonised largely by northern species,
because these species find here, under a lowland climate, the conditions
necessary for their existence, which are depth of water, form of lake-basin,
and chemical composition of the rocks of the drainage area. The similarity
of the plankton of the Scottish and Cumberland lakes, and of both with
that of some Scandinavian and Irish lakes, may be due in large part to their
being excavated in ancient lime-free rocks rather than to climatic influences.
The similarity between the arctic and alpine fauna of Entomostraca may be
accounted for by unlimited powers of dispersal rather than by the influence
of the Glacial period. It is true that the evidence from other groups is in
favour of the current views as to the relation of the alpine to the arctic
fauna, but we require to know far more about means of dispersal before the
matter can be regarded as decided. I will give two instances within my
own knowledge which illustrate the danger of relying on limited powers of
dispersal and upon the nature of the habitat in speculations regarding

420) MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

post-Glacial distribution. In 1907 Apus cancriformis, which had not been seen
in Britain for over 40 years, was found in Scotland by Mr. Balfour-Browne,
but had again disappeared the following year. Limnicythere mirabilis is
claimed by Zschokke as a member of the deep-water fauna of the Swiss lakes,
which is of northern origin and finds only in deep waters the conditions
necessary for existence in its southern station. But in 1905 I found this
species in a shallow rain-pool near Biskra in Algeria, and another species,
L. incisa, in a stream which irrigates the Oasis of Oumach.

The Plankton of Ennerdale Water.

The discovery of Limnocalanus macrurus in Ennerdale indicated at least a
possibility that other “relict” Crustacea might be found there, and made it
most desirable that the lake should be carefully explored. With this
purpose in view I visited the lake at the end of September 1922, and made
special efforts to collect the Crustacea of the deep water. The lake is shallow
at its western end, but the eastern part is a deep narrow trough with a
maximum depth of 148 feet. The shores are rocky, and with a very steep
slope.

The weather during my stay was unfavourable, but I was able to search a
large part of the deep trough with a coarse tow-net and light dredge, and
also to take plankton samples on three successive nights. Neither A/ysis
relicta nor any Amphipods were taken. This negative result should be tested
by examination of the stomachs of Char (Salmo alpinus), but I have not
been able to obtain specimens.

Table 4 gives the composition of the plankton between September 27th
and 29th. It is of course impossible to obtain accurate information as to
the vertical distribution of plankton without the use of a closing net and a
succession of short vertical hauls, but I donot think that such accuracy is, in
general, necessary, and a fair approximation to the truth may be got with
less effort and simpler means. My own collections were made with an
ordinary small plankton net, which was let down, together with a weight, to
the required depth and then towed a distance of about 100 yards. Naturally
the net fished both going down and coming up, but the results of the different
hauls are so distinct that they do in my opinion give information as to the
vertical distribution. I have not attempted to compare different hauls by
absolute numbers, but have counted the individuals of each species in the
whole or in part of the collection, and expressed their frequency as percentages
of the whole number of individuals of all species counted. This method gives
an accurate statement of the comparative frequency, though not of the absolute
numbers of any species, and it admits of a comparison being made between
samples from the same or different lakes however they may have been taken,
Similar figures are given for Wastwater and Coniston for purposes of
comparison.

OF THE ENGLISH LAKE

DISTRICT.

421

Tasie 4.—Composition of the Plankton of Ennerdale, Wastwater, and

Coniston.
The figures give numbers of nisin als a earch shecies Bey cent. of whole number Gonmatieel,
Surface. ||At Night—Surface., Deep Water.
| ; 9 | 5 || a |
Weis | Se £
| =) x a) = oi] = |) |
| IBQlSSi A |= |g s\— |
ENNERDALE WATER. IS si62| a] e wee] =!
Sept. 27-30, 1922. Ely all = |= |e ale |
ei gies Pe las = ||
a} ale | a || =I tea |
Ns pean | =
| |
LL ay Sh a oe | 1D | alo]
j = |= |
| Siidle GRIME gob0c0bnc0cv0sncce — | — |] 42) 37 | 2:38) — | — |} —}— |] —
| Holopedium gibberum ..............) ‘7| — ||20:2 |26-1 |32-1 |33-6 | 12:6 19 6] D6 | 36 |
| |
Bythotrephes luongimanus .. 2.00... .+: | 34) 78 |) 15 10-4) 115/172) G4 30 4-67) 7:2
| | | |
| Bosmina obtustvostris . 1.0.00. eevee — aed =| 1:2
Diaptomus gracilis ...............- |69:4| 7:8 ||16:2 49-216 |28:1|/46:8 6-4 34-6 50:8
Limnocalanus macrurus 6.0.2... vee |23:8 [81-3 ||58-2 13 47-1 19 33:4 35 54:8 [36:9
|
WASTWATER. iy
Oct. 1, 1922. Surface. | 80 feet. 100 feet. 120 feet.
Bosmina obtusirostvis v0.1... eevee 525 | 40 40:2 48:6
Bythotrephes longimanus ..... 0.0.04. 7-05 4-2 | 5°88 9:6
Toeppeadhorr@, IRRHEC 1 o0000ccce0n0oaes } 64 52 ‘98 2:05
JO RGRHORMNGS GPRUCHOS oo 000c0cc0000%08 SEG | 25:2 28-4 27-4
Cydlaps C@OYESOVUMD —o502000000000000 } eg) {at 24:5 12:3
|
ConisToN. Surface. | Surface. an ie Below
Oct. 2, 1922. Fine Net. | Coarse Net. | 50-80 feet. 100 eet,
Dip HOG GMA 000000656 ca0c00ne 4-9 | 372 57-7 48:9
Bythotrephes longimanus ............ + | oat 1:86 2:5
(eentodorchkindtierwree rer teeter oF — 56 3°9
| ID RRO YPRGURS 0550 008208000000 93 | 47-2 2] | | sell
| |
Cyclops abyssorum ................ 2:09 14 11-4 | 16:4

422 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

Tt is clear from the table that the plankton is by no means uniform
throughout the lake either vertically or horizontally, neither is it the same
by night and by day.

The chief component of the plankton is Diaptomus gracilis, which is
distributed throughout the lake, but is relatively more abundant in the
western end than in the deep eastern trough, and has a distinct preference
for the surface. The figures given for the deep-water samples must be
discounted to some extent, owing to the fishing of the net going down and
coming up. It is curious that this species seemed to be present in absolutely
smaller numbers on the surface by night than by day.

Limnocalanus is also distributed all over the lake, but is found only in
small numbers in the shallow water. It abounds both on the surface and in
deep water in the eastern trough, and may greatly exceed J). gracilis in
numbers. Although common at all times on the surface, it is noticeably
more abundant during darkness.

Holopedium gibberum, is only exceptionally found on the surface by day,
but is common in deep water, and at night forms a large part of the surface
plankton.

Bosmina obtusivostris, though apparently fairly common at some seasons,
was almost absent from the lake at the time of my visit, the few specimens
taken being in deep-water samples.

A curious feature of the plankton was tle appearance of Sida crystallina
on the surface at night. It was not taken during the day in any collections
in open water, but occurred in three out of four plankton samples taken at
night. All these samples were taken at the surface over water 80-120 feet
deep, the only sample in which Sida was not found having been taken in
shallow water. The specimens taken were almost all males.

Tat WNromostraca OF THE TARNS.

The occurrence of /aptomus laticens in Hawes Water and in Goats
Water (as recorded by Brady) raised the expectation that the fauna of the
high tarns of the district might prove to include other species representative
of the arctic-alpine Hntomostraca which do not occur in the lakes, and, at
the end of September 1922, I visited some of these tarns and made
collections in them and in pools and bogs up to an elevation of about 2000
feet. The resuit did not fulfil expectation, since PD. laticeps was not found in
any tarn other than Goats Water. There was also a notable absence of the
northern forms of Harpaticid, with the exception of Canthocamptus cuspi-
datus, which seems to be comparatively common in Sphagnum in running
water where the temperature is low. Nevertheless I think the results are of
sufficient interest to be given in some detail, The complete list of species’is
shown in tabular form,

OF THE ENGLISH LAKE DISTRICT. 423,

1. Ploutern Tarn. This tarn, which is fairly deep and contains trout, lies,
at a height of about 1250 feet, just on the east of the watershed between
Ennerdale and Crummock valleys, in a deep hollow in the grassy moor. It
is on the edge of the area of the Skiddaw Slates, the granite hill of Great
Borne rising steeply on the south. All around the tarn is swampy ground,
with trickling springs and hollows filled with Sphagnum.

The water is dark and peaty, and the marginal vegetation consists of
Littorella (dominant) and Lobelia. The tarn drains into Crummock.
Seven species only of Hntomostraca were taken, Gosmina obtusirosiris being
the dominant species. A large proportion of the females were ephippial,
but only one male was seen. Very few females carried summer eggs, so that
the colony was probably at its maximum and about to die out. /raptomus
gracilis was abundant, a large proportion being males. All were of a striking
blue colour, but of quite typical structure.

2. Greendale Tarn lies, at a height of 1320 feet, on the Borrowdale
volcanic series at the head of a glen north of Wastwater. West of it isa
gently sloping moor, while east of it rises the bare rocky Middle Kell. The
tarn is very shallow, with clear water and stony bottom, the vegetation
consisting of Lobelia, Littorella, Callitriche, Myriophyllum, and Jsoetes.
While I was unfortunately unable to take the temperature, I can say that
the water was (Oct. 1st, 1922) so intensely cold that it was impossible to
wade out and collect in the open. LHleven species of Entomostraca were
taken, among which Diaphanosoma brachywrum was the most abundant.
Bosmina was common, but none were ephippial. /iaptomus gracilis was
abundant, and here of a fine red colour *, but the majority were immature.
Alonopsis elongata was abundant, but none were sexual.

3. Levers Water, altitude 1350 feet, lies on the Borrowdale volcanic series,
and is a rather large, nearly circular, tarn in a barren rocky corrie. It owes
its existence to a morainic dam, but its outlet is artificially controlled and its
level somewhat raised for the supply of water to the Coniston copper works.
The floor and sides of the tarn are rocky and without any vegetation, and
Entomostraca are almost absent. The only species found in quantity was
Alonopsis elongata, and, besides this species, only a single individual each of
Bosmina obtusirostris and Acroperus harpe were taken in spite of prolonged
search. On the other hand, B. obtwsirostris was found common in quite a
small pool near the tarn.

The lifelessness of this tarn is rather remarkable, and I can offer no
explanation.

4. Low Water, at a height of 1786 feet, lies in a circular corrie beneath
the crags of Coniston Old Man, and is, as Levers Water, partly dammed for

* Ward (1904) notes specially the red colour of copepods in elevated lakes.

424 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

theJservice of the mines. It also is rocky, deep, and without vegetation, the
water rather dark coloured. The fauna consisted of four species only, and
these in small numbers. Diaptomus gracilis was here of a rieh red-orange
colour. A curious feature of the fauna was the extreme blackness of the
Alonopsis elongata, which proved to be due to the retention of the old shell
at moulting. The majority of the individuals showed three old shell valves
adhering and forming “lines of growth.” In other tarns and in the lakes
similar individuals were occasionally seen, but the feature was pronounced
only in Low Water. Lilljeborg mentions the same thing as characteristic of
specimens of this species from the high north of Sweden, and it is apparently
a character connected in some way with the low temperature of the habitat.

5. Goats Water, 1646 feet, is a small tarn in a combe on the north-east
side of Coniston Old Man. It owes its existence to a dam of rocks fallen
from Dow Crags, and the water from it soaks through the dam and issues as
a stream some distance below. The water is deep and clear, the shore rocky
and entirely barren. The fauna was found to be very scanty, but six species
were taken, the most noticeable being Diaptomus laticeps, conspicuous from
its red colour.

6. Highlow Tarn, about 2 miles N.E. of Coniston, at a height of about
600 feet, is mainly of artificial origin, being due to the union of several small
shallow tarns by the damming of the stream. lying comparatively low,
with shelving banks, its vegetation is rich and includes Nymphea. Conse-
quently its fauna is quite different from that of high tarns. Cerodaphnaa,
Polyphemus, and Bosmina longirostris are common, and Volvor was present
in such quantity as to give the water a thick green colour.

7. Sphagnum pools and spring waters.

On the high ground of the Floutern Pass and round the foot of Coniston
Old Man there is much swampy ground, often with little rills of running
water, and occasional pools largely filled with Sphagnum. The water was
uniformly very cold.

By the outflow of Greendale Tarn there are a number of small pools
generally margined with Sphagnum, but one of them, which was of some
depth, was choked with Callitriche and contained some larvee of the Newt
Molge cristatus. These larvee were kindly identified for me by Miss J. B.
Procter, who informs me that they were either newly hatched or only a few
days old. This very late hatching is very unusual, and no doubt due to the
coldness of the water.

The Sphagnum in running water was commonly tenanted by Canthocamptus
cuspidatus, a slow-moving pink-coloured Harpacticid which is a characteristic
northern species. In pools and springs the moss contained either C. cuspi-

datus, OC. eschokkei, or C. pygmaeus, but never more than one species in one

OF THE ENGLISH LAKE DISTRICT. 425

place. Several species of Cyclops were taken, of which the most interesting
is C. venustus, which was common in pools on Floutern Pass and was also
found by Greendale Tarn. It was somewhat unexpected to find C. nanus in
pools on Floutern Pass since I regard this species as a warm-water form.

It is worth noticing that Acantholeberis eurvirostris was only found in two
high-lying pools. This is a species found in lime-free bog waters, but it is
evidently not a cold-water northern form. Alona rustica and Chydorus piger
are species of similar preference, and were found rather infrequently on the
high ground.

Taste 5.—The Crustacea of the Tarns.

|
|
= q | 3 | “so “2 a
So | Belts 2) ae Fs) a
| SE (SE | EE) oe |e | Se
ms | 52 | oe | eS | ES | eS
[= a let |) Sie Ne i
| Diaphanosoma brachyurum ........ + Oe (tits oe |
Certiodaphnia pulchella .,.......... + |
DOT WHHO UGOSMUS.“sonaoocsooceos + |
: : |
ees ODEOROSIHE ge000000000008 S10 =f + + oo ar
ELurycercus lamellatus .......... pera||l Soon at +
Acroperus harpe ..... Soaosdoccdona 6 + Bic 4 “Ks +
Allonopsis elongata ..........+.--+- “f + ck ar =e ab
v |
AUIDTO CWOR ooccconneaoc Saeees eeaes ig a ar |
Alonella nana ........ Saati fentsbersores is +
| |
ie
“1 CLCISA) 30. poodoeDDa Ie a0 et Meet
Graptoleberis testudinaria ...... ea chal ||mens He +
}
Polyphemus pediculus .............-. + |
Cyclops leuckartt ........ O00 ABO ORS +
9 EGLS coovene Lestat ho Craeeener + + + j at +
Pi OL NUE etiolals: Rete cao os he ee 4 +
Fel) eUREU Sin oot Goede. on otc Gee + |
|
Dreptonrisy G7 AciUusm rare ie coco] ae + =a | sR
9 Taticeps ........---+---- og oc ge p09 £0 +
Canthocamptus zschokket .......... oe ag operate iceland Ame +

Highlow Tarn has all the characters of a lowland pool, but the remaining
tarns described are probably fairly representative of all the high-lying waters
of the district. Those examined fall into two groups, of the peaty and the
rocky type, the former represented by Floutern and Greendale Tarns and
the latter by Levers Water, Low Water, and Goats Water. Floutern and
Greendale Tarns have a relatively rich Entomostracan fauna characterized

LINN. JOURN.—ZOOLOGY, VOL. XXXY. 31

426 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

by Diaphanosoma, while the rocky tarms of Coniston have a very scanty
fauna, both of species and individuals. The brilliant colour, either red or
blue, of DY. gracilis in these high tarns is very striking.

The means of dispersal of the species is a problem of some obscurity. All
the tarns are of comparatively recent origin, owing their existence to
morainic dams or rock-falls, and each, as a rule, occupies the head of a
lateral valley.

Keilhack, studying high-lying waters in the Dauphiné Alps, concluded
that distribution of species was effected by migratory birds, and that wafer-
sheds rarely, if ever, exert any influence on distribution. While it is
impossible to offer any other satisfactory explanation, the agency of birds,
especially in the Lakeland area, seems to be of doubtful efficacy. A detailed
survey of the tarns of this compact and uniform district, with special
reference to the dissemination of species, would probably lead to results of
great general interest.

Notes on some of the Species.
LIMNOCGALANUS MACRURUS, Sars.

The occurrence of this fine Centropagid in Ennerdale is rather a startling
discovery, and raises problems of exceptional interest. It was first recognized
in a collection made by Dr. Pearsall on Sept. 23, 1921, the sample con-
taining five specimens. As a result of this discovery Dr. Pearsall arranged
for a series of collections to be made in the lake, and J also visited it myself
on Sept. 27, 1922, but, although I am able to give some account of its
distribution in the lake and its seasonal cycle, the facts are by no means
complete, and can only be ascertained by quantitative methods with suitable
apparatus. So far as information goes at present, the species is very abun-
dant in the lake and occurs in every part of it, at all events in autumn, which
is the period of maximum development ; but it is far more abundant in the
deep eastern trough, and occurs only in small numbers in the shallow western
end. All previous observations, by Hkman and others, agree in showing
that L. macrurus is only exceptionally found at the surface, and is almost
entirely confined to water below a depth of 5 metres. Further, the maximum
temperature of water in which it lives is 14° C., but even when the tempera-
ture is lower than this it does not come to the surface. In Ennerdale this is
not the case. The first collection in which it occurred was made on the
surface (in stormy weather it is true), the temperature of the water being
14-4° C., and my own collections between Sept. 27 and 29 proved that it lived
at that time in considerable numbers at the surface, although it certainly was
more abundant in deeper water, down to about 80 feet. It is impossible to
give exact figures, since I had no closing net at my disposal, but the figures
given in Table 2 (p. 417) will serve to show the distribution at that time
with, I believe, a sufficient approximation to the truth. I was unfortunately
unable to take temperature observations.

OF THE ENGLISH LAKE DISTRICT. ADT

Diurnal migration of the species was strongly marked at the time of my
visit. On three successive nights surface plankton samples were taken over
the shallow and medium (50-70 feet) depths, and in each case Limnocalanus
was found in much greater numbers than during the day. The figures in
the table do not bring out this fact so clearly as would have been the case if
absolute numbers could have been given, but the difference between day and
night plankton was quite obvious and striking on the spot.

So far as concerns seasonal distribution, the facts with regard to Limno-
calanus in Hnnerdale are somewhat uncertain. Ekman has shown that in
the Baltic Jakes there is only one breeding-period—in autumn. Young
appear first in March, and by May the first-hatched young are fully grown.
But there is then a period of about five months during which little or no
growth takes place and the genital organs remained undeveloped. Sexual
maturity is reached in autumn, and breeding begins in November at a
temperature of about 7° ©. The adults of the previous year die off in
spring and all have disappeared by May.

Dr. Pearsall’s collections may be regarded as indicating a similar life-cycle
in Ennerdale, but are by no means conclusive. Neither young nor adult
were taken on Feb. 18, 1922, but two adults and many young were found in
a sample from the shallow end of the lake on April 20, though, curiously
enough, none at all were taken in a deep-water collection on the same day.
In June and July no young, but a very few adults, were taken ; but in
October adults were common in 60 feet of water. It seems safe to say that
young are only found in spring, and that adults are abundant, and even the
commonest plankton species, in September and October.

The interest of this species lies more, however, in its geographical distri-
bution and its origin as a member of the fresh-water fauna, and it is necessary
to go into this question in some detail since its occurrence in Hnnerdale
raises a particular difficult geological problem.

Hkman has dealt exhaustively with the structure and distribution of
L. masrurus (1913), particularly with reference to the Baltic area, and his
results may be summarized as follows :-—

1. LZ. macrurus of fresh-water lakes is not specifically separable from
L. grimaldit, which inhabits brackish water in the Baltic and Caspian Seas
and along the coasts of Alaska and Siberia.

The differences are :—

(a) Shape of head, the dorsal contour being more swollen in L. macrurus.

(6) In L. grimaldit the last thoracic segment is rather triangular or
pointed, or may have a minute hook-like projection, whereas it
is rounded in L. macrurus.

(c) The antenne are longer in L. grimaldi.

(d) The fureal rami of L. grimaldii are longer than in L. macrurus.

While the two extremes are readily distinguished, they are united by
local varieties presenting every intermediate condition,

aie

428 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

2. In Hurope ZL. macrurus is only found in lakes which can be proved to
have been separated from the sea in post-Glacial times. It is therefore a
‘‘yelict” form in the strictest sense. It has not arrived in any of the lakes
it now inhabits by active migration.

3. The shape of the head in LZ. macrurus, while not at all variable among
individuals in any given lake, varies greatly according to the locality, and
it can be shown that those lakes (in Hurope) which contain the extreme
fresh-water form have been separated longest from the sea, whilst in more
recent lakes the Limnocalanus approaches more nearly to the grimaldi
form.

4. It is evident that Z. grimaldii is the parent form, and that LZ. macrurus
is a variant from it which has arisen independently in many places, the effect
of the change of medium being always in the same direction. |

5. The first step in the transformation occurred in the Baltic during the
* Ancylus Lake” period, and the Baltic lakes in which Limnocalanus occurs
are relicts of the Ancylus lake.

With the most interesting speculations of Ekman on the subject of the
origin of species founded on these facts we have no concern here.

Now, while it seems to me that Hkman’s conclusions with regard to the
Limnocalanus of the Baltic area and of the Caspian Sea are entirely well
founded, there are certain criticisms to be made, and it is difficult to extend
his explanation to the Limnocalanus of Hnnerdale and the lakes of North
America in which it occurs.

In the first place £ cannot unreservedly accept Ekman’s basis of measure-
ment of the head form on which much of his argument depends. Hkman
has adopted the following system of measurement :—

(1) Head-length—to the dorsal groove immediately in front of the mandi-

bular muscle. (See text-fig. 2.)

(2) Head-height—B—expressed as a percentage of half the head-length.

(3) Height of vertex above or below the dorsal line of the thorax expressed

as a percentage of half the head-length.

The last measurement is negative in all the grimaldii forms, and positive
only in the extreme macrurus forms where the dorsal contour of the head
swells up above tne dorsal line of the thorax.

While the first and second measurements are fairly definite and extra-
ordinarily constant, the third seems to me so uncertain as to be of little value,
since (as Hkman has himself pointed out) the selection of a dorsal horizontal
base-line is a matter of guess-work. I have been quite unable to find agree-
ment between his own figures for this character and the drawings of the
specimens from which the measurements were taken ; in fact, these measure-
ments cannot, as I believe, give a real definition of the shape of the head,
which can only be shown ina drawing. Still, for purposes of comparison,
I give, in Table 6, measurements taken hy Hkman’s methods for the
Hnnerdale and other forms.

OF THE ENGLISH LAKE DISTRICT. 429

It is evident that, if ZL. grimaldi be the parent form, those races which
approach it most nearly in shape of head and in other respects may be regarded
as most nearly related, and consequently more recently separated from. it.
It is therefore of some interest to apply Ekman’s system of measurement
to races other than those of the Baltic region, and to see if by that means
any clue to their age may be found.

Limnocalanus macrurus from Ennerdale, showing method of measurement.
A. Head-length. B. Head-height. ©. Crown-level.

TabiE 6.—Measurements of Limnocalanus macrurus and L. grimaldi.

Figures marked * are taken from Ekman, 1914.

| pee Head- | Head- | Crown-)
Length. percent! length. | height. | level.
of body. A. B. A&B.
ISMMNERAVS ..o0000000000% 2 208 | 13:1 229 157 — 26 eS loik
d 206 | 14
Sweden—Mijlaren ...... 2 234 | 16 32 126 —16 110
ic} 23 16
3 IAGO cocooece, QD | Bil || sy 33 Wey || Sede ox] Wile
U.S.A.—Green Take .... Q | 228 | 14 32 134 | —11-5 | 122'5
3 224 | 13:8
9 Take Hrie...... co) 2G | IGS | ey 140 | —17 123
Si BY 13:3 | | |
5 Lake Canandaigua 9 9:53 15 BY 13 | =1G ) 1g
3 Soll |) Ne 141 —15 128
Caspian Sea ...:........ co) 2°59 158 = 90* | —25* 65*
i

430 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

I possess a few specimens of LZ. grimaldii from the Caspian Sea which
were given to me by the late Canon Norman some years ago, and by the
kindness of Dr. C. Juday I have had the opportunity of examining
LI. macrurus from Green Lake, Lake Brie, and Lake Canandaigua (one of
the Finger Lakes east of Lake Ontario). Dr. S. Ekman also has heen good
enough to send me specimens from Lakes Miilaren and Insjén in Sweden.

Measurements of these races are given in Table 6.

So far as the numerical expression of the head-form goes, the Hnnerdale
race cannot be compared with any of the Baltic races, since in none of them
is so great a head-height aceempanied by so low a crown-level. The only
race with similar head-height (Lake Mjésen) has a dorsal swelling rising
above the horizontal dorsal line. On the other hand, the high, evenly-
rounded dorsal outline rather closely resembles Hkman’s figure of the race
from Unden in Sweden, though that race has a crown-height according to
his measurements of only —6. It seems that in respect of this character the
Ennerdale race stands somewhat apart, but that it must have been evolved
from the grimaldii-ty pe at a comparatively remote period.

The North American races differ but little in head-form, and conform more
closely to the Baltic type. Both have a posterior more or less horizontal part
with an anterior descending slope, and agree very closely in appearance with
the Malaren form, though the measurements do not agree very well with
those given for that race (Pl. 28. fig. 6). But, as compared with the Enner-
dale race (Pl. 28. fig. 1), specimens from North America certainly approach
far more nearly to the grimaldii type, and are consequently to be regarded as
of more recent origin.

This conclusion is strengthened by a consideration of the other characters
in which the two forms differ.

(1) The true grimaldii form of the Arctic Ocean (examples from Jana
River) measure over 3 mm., while the somewhat modified form of the Baltic
may also attain 3 mm. (Gulf of Finland). It is, on the other hand, roughly
true of the Baltic area that the oldest fresh-water races are the smallest
(Sommen 1°87 mm., Unden 1°72 mm.). Small size is therefore to some
extent evidence of prolonged sojourn in fresh water. In respect of this
character the Ennerdale race (2°18 mm.) holds a more or less middle position.
The race from Lake Hrie, on the other hand, is one of the largest known
(2°86 mm.), greatly exceeding any other fresh-water race and exceeded only
by those of North Siberia and the Gulf of Finland in salt water, The
Green Lake form, though similar in structare, is but little larger than that
of the Ennerdale race.

(2) The antenna in the true grimaldii form reaches to the base of the furca
or even further (PI. 23. fig. 5), whereas in typical L. macrurus it does not
extend further than the first abdominal segment. In this respect the Hnner-
dale race belongs to the extreme fresh-water type, while those from North

OF THE ENGLISH LAKE DISTRICT. 431

America have the antennz slightly longer. In the Miilaren race, which is
one of the most primitive forms, the antennz in the female may reach to the
base of the furca.

(3) The length of the furcal rami in proportion to the whole body differs
in different lakes, but the published information on this point is too scanty
to admit of a definite conclusion as to whether there is any connection
between the shortness of the furea and the length of sojourn in fresh water.
The fureal rami of the Ennerdale and the American forms are notably
shorter than those of the grimaldii form from the Caspian, but, on the other
hand, both the Milaren and the Insjén races have these rami as long as or
longer than those of my Caspian specimens.

(4) The pointed triangular form of the lateral expansion of the last thoracic
segment has been given by Prof. Sars as one of the distinctive characters of
L. grimaldii, since in all the fresh-water races this segment is simply rounded.
I find in my Caspian Sea specimens that this segment is produced laterally
into a very well-defined point (PI. 23. fig. 5), and it is important to note that
a similar point is to be found in some of the specimens from Green Lake and
from Lake Canandaigua, and very rarely also in those from Lake Hrie,
while the Ennerdale race has the segment simply rounded. This character,
small as it may appear, seems to be of importance in indicating relationship
with the parent form, and the North American race must for this reason also
be regarded as of comparatively recent origin. The primitive Milaren race
has this segment generally very slightly angular, and occasionally there is a
definite point as in LZ. grimaldi (PI. 28. fig. 8).

It is remarkable that, in spite of the considerable differences in general
body-form between the brackish and fresh-water races of Limnocalanus,
there is no appreciable difference to be found between any of the appendages
of the two forms, except as regards the length of the antennules. I have
compared the appendages of the seven races ayailable to me, and find the
agreement perfect in almost every detail. Thereis, however, one very slight
difference which may be peculiar to the Caspian race, namely the absence of
a spine on the outer distal angle of the first joint of the exopodite in the
right fifth foot of the male. Prof. Sars does not figure a spine in this
position, neither was it present in a specimen examined by me, but it is
present in all other races of which figures are published.

Having dealt, so far as I am able, with the biology and structure of the
Ennerdale and North American races of Limnocalanus in comparison with
others, there remains the consideration of the question as to whether these
races can be regarded as ‘“‘relicts”’ in the same sense as those of the Baltic
area.

It must be admitted that in neither case can any definite answer be given.

There can be little doubt that the whole of the Lakeland area was
completely covered by ice during the Glacial period, though the view that

432 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

the lakes themselves are glacially-eroded rock-basins is not universally
admitted. In any case, however, the fauna must have reached these lakes
after the withdrawal of the ice.

According to Clifton Ward (1876) the passing of the Glacial period was
immediately followed by a submergence of the whole district by as much as
2000 feet, which would leave only the highest ground projecting as a
number of small islands in the midst of an arctic sea. If a submergence to
this or to a much smaller extent can be admitted, then L. macrurus might
well be a relict of this sea left in Ennerdale. Unfortunately Ward’s
conclusions are not admitted by geologists of the present day, though I have
not been able to find in any literature available to me any serious discussion
of the question of post-Glacial changes of level in this district. The matter
is hardly referred to in Marr’s ‘Geology of the Lake District, but
apparently be does not admit a submergence of more than a few feet, which
could not have made any material difference to the distribution of land and sea.

If a submergence to an extent sufficient to bring the sea in the first
instance into direct connection with the Hnnerdale basin cannot be conceded,
there are only two alternatives. Hither the interpretation of the geological
facts is wrony, or a marine species has either been able to migrate for miles
up a rushing stream of fresh water or has been transported directly into the
lake. No biologist would for a moment consider such migration or trans-
ference to be even conceivable. It must be remembered that we are dealing
with a species which occurs in this one lake only in the British Isles and
cannot have been transported to its present habitat from any other lake in
Britain.

Even if Clifton Ward’s great submergence could be proved true, it is still
a little doubtful if it would satisfactorily explain the facts. Our knowledge
of marine relicts seems to point to the conclusion that transformation of
marine species into fresh-water relicts has rarely happened, and only in cases
where very large areas of water have been involved. Thus the L. macrurus
of the Baltic area owe their origin to the transformation of the Yoldia Sea
into the Ancylus Lake, rather than to the separation of lakes from the Yoldia
Sea itself. In other words, Z. grimaldii was transformed into a form of
L. macrurus in the Ancylus Lake and not, in the first instance, in the
existing inland lakes. Similarly, we should not expect that a race of
L. grimaldii could have survived in so smalla lake as Ennerdale itself even
if it could be shown that the lake had itself been cut off from the sea.

To explain the occurrence of this species in Ennerdale in the same way as
its oceurrence in the Baltic lakes, it is more reasonable to assume that the
Irish Sea itself has passed through a history somewhat similar to that of the
Baltic, having been changed from an arctic sea containing L. grimaldit to a
fresh or brackish water, and finally to its present condition, and that during
the second phase LZ. grimaldit became in part transformed in the direction of
L. macrurus. Further, that conditions were such that, on the renewed

OF THE ENGLISH LAKE DISTRICT. 4393

entrance of salt water, immigration into Hnnerdale was possible. I must
freely admit that such a supposition is not supported by the geological
evidence, but it does not appear to me to be an altogether impossible
assumption in view of the following facts.

There is evidence of a subsidence of land immediately after the withdrawal
of the ice, and of a later elevation at a time when the submerged peat-beds
and forests were formed. But the amount of the latter elevation is uncertain,
and there is no positive evidence of a rise of more than 60 feet. On the
other hand, the occurrence in Jreland of the Red Deer and Reindeer seem to
necessitate the supposition that there was some land connection between
England and Ireland in neolithic time, and, as Jukes-Browne has pointed out,
firstly it is possible that this elevation may have been greater than has been
supposed, and secondly it is very probable that the floor of the Irish Sea,
which was thickly covered by drift, has been greatly eroded and re-modelled.
It may consequently have been at so much higher a level that a comparatively
small elevation may have sufficed to effect a land connection with partial or
complete damming off of the sea.

Dr. Scharff has assumed the existence of a great lake occupying the
trough of the Irish Sea to explain the present distribution of the genus
Coregonus in Britain and Ireland, and if such a lake could have originated
by freshening of a Glacial sea, not only might Z. grimaldii have been therein
isolated and transformed, but also M/ysis oculata could in this way have been
changed into JL, relicta and have reached its present station in Lough Neagh.

There are two obvious objections, namely that JZ. relicta does not live in
Ennerdale and Z. macrurus does not occur in Lough Neagh. Further, that
conditions which would allow the latter to reach Ennerdale should also allow
it to reach Wastwater and perhaps other lakes. To these objections not
much importance need be attached until more is known of the conditions
which determine the distribution of fresh-water Crustacea. Not only does
the plankton of Lough Neagh differ radically from that of the English lakes
in general, but these lakes differ unaccountably among themselves, and
Wastwater and Ennerdale, alike as they seem to be in physical conditions,
contain quite different plankton. Hence it cannot be assumed that a species
would establish itself in every lake to which it had access.

It is equally difficult to explain the distribution of Limnocalanus in North
America. Its distribution follows rather closely that of Mysis relicta, as
shown in Table 7, page 434.

L. macrurus therefore occurs in all the great lakes and in a number of
lakes belonging to the same drainage system, but not, so far as I can find, in
any lakes outside this region.

So far as the origin of these species is concerned, the view once held that
the Great Lakes were themselves at one time invaded by the sea and that
M. relicta is a true relict within them has been generally abandoned, but at
the same time there appears to be no doubt that the sea did, at a late stage

434 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

in the post-Glacial history of the region, penetrate up the St. Lawrence
valley and flood the basins of Lake Ontario and of Lake Champlain. At
this time there existed an eastern outlet from Lake Huron through the
Ottawa valley. The marine deposits of this stage are found at a height of
400-500 feet by Lake Champlain and at 600 feet near the east end of Lake
Ontario. It would appear that we have here all the conditions necessary for
the isolation and modification of these two “relict” species, but there are

TABLE 7.
| Mysis Limnocalanus |
relicta. MacrUrus. |
BalcelOntariov ele P +
[ove “uMtichioanieneet ia zt + |
Superior sb age + ab, |
Pe UELTUR OM: Secretary yet _ +
SMS EMC Laine ees quite - +
privat) FOV Sats 3 ep ene tate aP | +
Apr GGUIVEN os doe a Adda + +
Gieentlialvome eee + +
The Finger Lakes :— |
Canandaigua...... + ah
Cayce ea: + | +
| INOW, “Soccocooee + |
| OWESCO sodgnovscne — + |
| SONA booanvdo00 + +
| Skaneateles ...... _ | ain |

two great difficulties in the way, namely the absence of Mysis relicta from
Lakes Ontario and Champlain and the existence at or before that time of the
Niagara Falls, which would be an effective bar to migration into the other
lakes. On the other hand, the absence of a species from a lake at the
present day does not necessarily prove that it has never existed there, and
the occurrence of both species in the Finger Lakes, which form part of the
drainage system of Ontario, seems to be good evidence that they have
reached these lakes from Ontario. Their penetration into the other Great
Lakes must, one must suppose, have taken place by way of the Ottawa con-
nection, and in any case we are forced to assume for L. macrurus a capacity
for active migration up rivers which is denied by Ekman.

* So far as I know, there is no published record of the occurrence of Limmnocalanus in
Lake Ontario, but Dr. C. Juday informs me that he has found it in a plankton sample from

this lake taken by Dr. N. A. Clement in October 1922. I am greatly indebted to Dr, Juday
for this information.

OF THE ENGLISH LAKE DISTRICT. 435

The recent discovery of another species of the genus (L. johansent, Marsh)
in a tundra pool in Alaska seems to raise a new problem altogether. But
Marsh’s description is very short, and it is not clear that this new species is
really distinet from L. grimaldit or L. macrurus, the only essential difference
mentioned being the shortness of the fureal rami. It is important that this
species should be fully described and its distribution ascertained.

Diapromus LATICEPS, Sur's.
Syn. D. hircus, Brady.

Brady’s species, J). hircus, was established in 1891 on specimens taken
from Goats Water, an elevated tarn on the slopes of Coniston Old Man.
There can be no doubt that his species is identical with J). laticeps, from
which it differs in no essential character.

It is an arctic and alpine species occurring mainly in Scandinavia, but
recorded from Akmolinsk in Russia, the Julian Alps, Serbia, and
Herzegovina. It has also been taken in several lakes in Scotland and in
Ireland.

The only lake in the Lake District in which the species is found is Hawes
Water, which is the highest of all (694 feet), and for that reason may have
a different temperature. It seemed likely that 1). daticeps might be common
in the high tarns of the district, but, as [ have said in my notes on these
tarns, this is not the case. On the other hand, our knowledge of the fauna of
these tarns is very incomplete, and it may prove to occur in the tarns of
Harter Fell and to have reached Hawes Waiter from thence.

CYCLOPS ABYSSORUM, Sars.

This species has been found in all the lakes examined with the exception
of Hnnerdale, Derwentwater, Bassenthwaite, and Esthwaite. In Hnnerdale
no limnetic species of Cyclops was seen in any of the samples taken either
by Dr. Pearsall or myself, but Brady has recorded the occurrence there of
C. vicinus. The occurrence of that species in the plankton of this district
seems most unlikely, and it is more probable that Brady had specimens of
C. abyssorum, in which case the species once inhabited Ennerdale but has
now disappeared.

Prof. Brady has recorded the occurrence of this species in Windermere
and Coniston in deep water in Aug. 1883, and Prof. Sars, in his ‘ Crustacea
of Norway,’ describes it as inhabiting only the deeper layers of water. My
own collections in Wastwater and Coniston show that, though it is found in
autumn on the surface, it is distinctly more abundant in deeper water, and
its absence from Derwentwater, Bassenthwaite, and Esthwaite is no doubt
largely due to their small depth. It is replaced in these lakes by C. leuckarti.
In 1911 a few specimens were taken by me in Bassenthwaite, but it is
evidently not an established member of the plankton in that lake.

436 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

The seasonal cycle of the species cannot be determined with certainty
from Dr. Pearsall’s collections, which were mostly taken on the surface, and
therefore are unreliable in the case of a deep-water species; but a careful
comparison of the records trom the different lakes leads to the conclusion
that there is only one period of breeding, namely in the autumn, and that
the adults die after breeding, the species passing the winter either in the
form of resting-eges or as nauplii. In all the lakes mature specimens were
rare or absent in spring and early summer, but were abundant in collections
made between August and November. Females bearing eggs were found in
Hawes Water in August, but in the other lakes only in September or
October. If this is a true statement of the breeding cycle, it resembles very
closely the cycle in L. macrurus—another cold-water northern form. On the

TABLE 8.—Measurements of Cyclops abyssorum.

| Furcal Sete. |

| Length | Furca. | ————— a” — - | Seta 3.

| (ineluding|Per cent.} Outer- | 5 | a 4 Per cent.

furca). | of body.| most. en ee of body.

| ij pow |

| Wastwater ...... 135mm.) 135 100 418 481 194 B44

| Crummock ...... 1:48 138 af 402 | 484 186 35°3

li@onistontieeee eee 1-44 13:7 394 | 485 190 36:3
Ullswater ........ 1-42 116 ol 388 | 456 190 33

| Hawes Water ....| 1:45 12°8 = 829 | 379 250 31

| Windermere ...... 1:77 TY») 5, 346 | 423 160) || Sil

| |

| Grasmere were 1-68 14'9 % 34] | 424 190 32°5

other hand, Scheffelt (1908) and Burckhardt (1900) found two breeding-

_ periods in the Cyclops of the strenwus-group studied by them in the Black

Forest and Swiss Lakes respectively. It is not, however, clear with what
race or species these authors were dealing, since the species of this groap
recognized by Sars and Lilljeborg as distinct have not been generally
admitted, and many writers have followed Schmeil in including all within
the species C. strenuus. It seems certain that the three species C. lacustris,
C. scutifer, and C. vicinus should be maintained. Whether C. abyssorum is
sufficiently distinct from C. strenuus as defined by Sars is rather more
uncertain, but the evidence goes to show that it is a limnetie and cold-water
representative of C. strenuus, which it is convenient to distinguish. It is
readily distinguished from typical C.strenuwus by the greater length of the
antennee, of the furcal rami, and of the furcal sete. The much greater

OF THE ENGLISH LAKE DISTRICT, 437

relative length of the innermost seta in C. abyssorum is perhaps the easiest
character by which the species may be separated, but Prof. Sars has
enumerated a number of details wherein the species differ. It is to be hoped
that Prof. Sars’ limitation of these species will be followed, since it is most
probable that they will be found to have different life-cycles and different
habitats and distribution. The general’ adoption of Schmeil’s definition of
C. strenuus has made comparison of statements with regard to these questions
unreliable and indeed valueless. So far as published information goes, it
seems that (. abyssorum is confined to northern lakes, and is absent from
Swiss lakes, where apparently a form resembling C. scutifer takes its place.

The Lake District species is undoubtedly C. abyssorum, though the form
of the receptaculum differs to some extent from Prof. Sars’ description and
agrees more closely with that of C. lacustris.

Those who (e.g. Graeter, 1903) accept the species C. strenwus in its wider
sense lay great stress on its variability, and it might be supposed, if their
view is correct, that great differences would be found between individuals
from different lakes. This is far from being the case. In general form of

body

particularly in the expansion of the last thoracic segment—there is,
so far as I can find, no appreciable variation. In size, in the length of the
furea and the relative length of the fureal sete there are differences, but
they are not great. In Table 8 I have given measurements of individuals
from seven of the lakes. hese are, in the case of the first four, averages
from ten specimens, but in the remainder of from two to six.

Tt will be seen that the smallest specimens are from Wastwater and the
largest from Windermere and Grasmere, which is hardly according to
expectation. Those from the “ primitive ” lakes agree in having the longest
sete, together with rather long furca, but the longest furca is found in
Grasmere. ‘The differences are, however, by no means sufficient to bridge
the gap between C. strenuus and C. abyssorum.

It is rather remarkable that the colour of the Lakeland individuals is
commonly a very pronounced yellow. The species has not, in fact, at all the
general appearance of a limnetic form, but rather of an inhabitant of ponds
and ditches.

CycLops veNustus, Vorman 5: Scott.
Syn. C. erinitus, Graeter, 1908.

Cyclops venustus was described in 1906* from specimens taken from
marshy ground on Exmoor. It has since been found by Mr. D. J. Scourfield
on Exmoor and Dartmoor, and by me in the Bog of Allen, near Newbridge
in Ireland. It appears to be a species confined to lime-free waters, and may
be included among what may be called the Sphagnophil HEntomostraca, such
as Acantholeberis curvirostris, Chydorus piger, Moraria brevipes, ete.

* ‘Crustacea of Devon and Cornwall,’ p. 189 (London, 1906).

438 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

The description given by Norman and Scott is not altogether complete,
inasmuch as no mention is made of the peculiar markings on the integument,
which are quite distinctive, and the allusion to the swimming-feet as being
similar to those of C. vernalis is misleading. It seems therefore advisable
to supplement their description to some extent.

The most characteristic feature of the species is the very pronounced
crenulation of the margins of the abdominal segments and (to a less extent)
of the last thoracic segment. The surface of the integument of the thorax
is marked by delicate strize which give it the appearance of being minutely
wrinkled, while the surface, both dorsal and ventral, of the abdominal seg-
ments is either striated or ornamented with lines of minute pits similar to
those found in C. diaphanus. These pits seem to run, at least on the dorsal
surface, on the crest of low ridges, which can be distinctly seen-in side view.
The fureal rami are provided, as in C. viridis, with cilia on their inner
margins, but these cilia are in groups and do not form a continuous
fringe. Usually the anal operculum is very prominent.

The swimming-legs differ markedly from those of C. vernalis not only in
their more robust form, but also in the number of spines and setee on the
terminal joint of the outer ramus (see Table 9). But C. vernalis is subject
to some variation in respect of the number of spines, and it is not unusual to
find individuals with a spine-like formula of 3.4.4. 4. as in C. venustus ; but in
these cases the number of setee is not increased. C. vernalis also resembles
C. venustus in frequently having the cuticle of the abdominal segments
pitted. The two species are, however, quite distinct, the form of the furca
alone sufficing to distinguish them, apart from the differences in the form of
the swimming-legs and the excessive pitting of the cuticle in C. venustus.

The resemblance to C. capillatus, Sars, is somewhat closer. The antenne
are 12-jointed in each species ; the form of the receptaculum and the 5th leg
is closely similar, and the spine- and seta-formula of the legs is identical.
On the other hand, the legs of C. capillatus are distinctly more slender, and
the fureal rami much more elongated and smooth on their inner edge.
Further, though the abdominal. segments show a slight toothing of the
posterior margins, this is not pronounced, and I have not been able to detect
any surface pitting in specimens from Norway which I collected many years
ago in the Romsdal.

There remains for comparison the species described by Graeter in 1908
under the name of C. crinitus. M. Chappuis has been kind enough to send
me a mounted specimen of this species, and an examination of this specimen
and of the published descriptions of Graeter and Chappuis convinces me that
the two species are identical. They agree in the following characters,
wherein they differ from most other species :—

(1) Antennule of 12 joints.

(2) Fureal rami short, slightly divergent, with groups of cilia on their

inner edge.

OF THE ENGLISH LAKE DISTRICT. 439

(8) Median seta of the furcal rami of unusual length.

(4) Outer seta of the furea of half (crinitus) * or scarcely more than half
(venustus) the length of the inner seta.

(5) Swimmineg-legs with the same spine- and seta-formula f —a formula
only found otherwise in C. capillatus, Sars.

Graeter states that the abdominal segments are jagged and that the surface
of the cuticle is pitted over the whole body, but I have not been able to see
this in the specimen, owing, no doubt, to its transparency.

Since Graeter’s name is later than that of Norman and Scott, I have
therefore regarded it as a synonym of C. venustus.

Tt is evident that this species and C. viridis are very closely related, as has
already been pointed out both by Graeter and by Chappuis, but they are
certainly distinct.

This species was found to be not uncommon in Sphagnum pools near
Greendale Tarn and on Floutern Pass.

TaBLE 9.—Showing spines and setee on last joint of the exopodite
in species of the C. viridis-group.

jLeg 1 2 3 4
Ce | aN
3. | enn :
a Rn n nD
=i|aifel]/e@)sias|ale
We | | | ee es |] ee
[2 | Bla lela|n | a |»
| | | | |
| Cyclops venustus, Norm. &. Scott ..| 3 | iS || ab 5 i) A 5
A G@rupicoky (Entice) Geese unnes eel Aso es 4 | 4 5
|
|
DIN-WLSy NSCMeT eas ye sci Dat gi a 3 a 3
ap CMR, SENS 5.6 bln boo 0c | BS Pe Oey ee
| | |
een |
oy DEACHAS IMISEMER . 5.4 obo d006 ier ss 25) 83 Veale ees!
|
| |
5 CGHAGES WE os aes esc sno Geet y a rae dba bell dl
|
FOUN, SHS cco coHon ands VS By eae eal) Ab abo dt
|

Sipa CRYSTALLINA, O. F’, Muell.

S. erystallina is a littoral species not properly, or at least generally,
belonging to the plankton, though frequently recorded as occurring in open
water. In the Lake of Geneva, Lake Como, Lake Maggiore, and other

* Jn the specimen which I have examined this seta is more than half the length of the
inner seta, and so agrees completely with C. venustus.

t In the specimen at my disposal it is impossible to count the spines and sete with
certainty on all legs. I haye given in the table only those which were beyond doubt.

440 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

large Swiss lakes, however, it appears to be a regular member of the
plankton, and Burckhardt has described the form there found as a distinet
species—S. limnetica, characterized chiefly by the absence of a nuchal organ.

In the Lake District it was found in small numbers in the majority of the
collections from Derwentwater, so that it may probably be regarded as really
limnetic there, whereas the few records of it from Bassenthwaite may be due
simply to migration from the littoral region. The same is the case with
Esthwaite and Grasmere. On the other hand, my experience of night
collecting in Ennerdale seems to show that this species, though remaining
in the littoral region during the day, may take to the open water at night.
(See p. 422.)

All the specimens from the Lake District were of the typical form, with
well-developed nuchal organ.

HoLopEDIUM GIBBERUM, Zaddach.

The distribution of Holopedium in the Lake District is somewhat peculiar.
It is recorded by Beck from Hasedale Tarn (915 feet) and from Windermere,
and it also oecurs in Grasmere (P. A. Buxton, Sept. 1912) and Rydal Water
(D. J. Scourfield). In Ullswater it was found in Sept. 1912 and on Aug. 20,
1920, but not in any other collection made by Dr. Pearsall. In Ennerdale
it was present in very small numbers in three out of seven collections taken
by him, but was abundant in my own collections in Sept. 1922. So far as
is known at present it does not oceur in any other lake in the district.

Hasedale Tarn, Grasmere, Rydal, and Windermere form part of the same
drainage system. The fact that it has so rarely been taken in Windermere
may possibly be due to its being occasionally carried into the lake, but not
becoming established there ; but it is more probable that it is not uniformly
distributed in the lake, and also that, being a deep-water form, it is not
taken in surface plankton. In Ennerdale it is fairly evenly distributed in
both shallow and deep parts, but it is rare on the surface during the day-
time, preferring deep water. At night, however, it is abundant on the surface.

This preference for the deeper layers of water does not appear to be
generally characteristic of the species, since Scheffelt (1908) states that it is
found at the surface at all times; but he also found it to be unevenly dis-
tributed and to occur in swarms. Lilljeborg notes the same thing in
Sweden.

It is generally regarded asa “ Glacial relict,” being confined to northern
regions or to cold water.

The genus DAPHNIA.

The inerease of knowledge of local and seasonal variation among the

limnetic Daphnias has led to the general recognition that any division into

well-defined species is impossible. All are therefore regarded as belonging
to one species, 1). longispina, within which an almost infinite number of

OF THE ENGLISH LAKRA DISTRICT. 44].

“subspecies,” ‘‘ varieties,’ and “forms” have been described. I do not
propose to discuss the question of the systematic arrangement of these
Daphnias, but it seems to me that the enlargement of the significance of the
name J). longispina in this sense has serious disadvantages, and that the
suggestion of Langhans that a new name, VD. variabilis, should be introduced
to include the whole series of these variable Daphnias might with advantage
be adopted. The question can, however, hardly be regarded as finally
settled, and, for the present, it is more satisfactory to accept Lilljeborg’s
longispina, hyalina, and cucullata, whilst fully recognising that the three are
united by transitional forms.

All the limnetic Daphnias of the lakes belong to the “species” hyalina
with its “subspecies” galeata and lacustris. Beck has recorded the occur-
rence of DP. cucullata (under the name Hyalodaphnia berolinensis), but this
species has been seen in the district by no one else, and it is probable that
he was mistaken. Daphnia is absent altogether from Wastwater and
Ennerdale Water, and apparently also normally from Bassenthwaite. There
are none from the latter lake either in Dr. Pearsall’s collections or my own,
and Mr. Scourfield informs me that he also failed to find it there. On the
other hand, Miss Pratt records both D. pulex and D. longispina as occurring
in April 1898, so that apparently some species occurs there at times. As
Bassenthwaite receives the overflow from Derwentwater which contains an
abundance of Daphnia, it is very remarkable that the same form should not
establish itself in Bassenthwaite.

The distribution of the different forms of Daphnia in the district seems to
bear little or no relation to the drainage system. While Buttermere and
Crummock, which were originally united into one lake, both contain the
typical D. hyalina, which occurs in these lakes only, the Daphnia of
Esthwaite is quite different from that of Windermere, into which Hsthwaite
drains, and the same form, D). galeata, inhabits Windermere and Coniston,
which are not connected in any way.

DAPHNIA HYALINA s. str., in Buttermere and Crummock.

In the form from these two lakes the front of the head is usually very
nearly straight or slightly concave (text-fig. 3, A): but it is very variable,
and may occasionally be distinctly convex as in )). pellucida. In a collection
made on Aug. 27, 1920, several specimens were seen with the front of the
form shown in text-fig. 3,B. An exactly similar variety is illustrated by
Lilljeborg (1900, pl. xv. fig. 4) as “ forma estivalis monstrosa.” It is
characteristic of the Crummock Daphnias that the shell-spine forms a con-
siderable angle with the body axis—from 20°-40°. Although I regard this
form of Daphnia as properly referable to D. hyalina s. str., Lilljeborg, in
some respects the population is transitional on the one hand to D. lacustris
and on the other to D. pellucida.

LINN. JOURN.—ZOOLOGY, VOL, XXXY. 3%

442 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

In a plankton sample of Sept. 1912 a single specimen was met with which
obviously belongs to ). galeata (text-fig. 3,C). I do not think that this
specimen can be regarded as a sport or mutation from D. hyalina, but rather
as an instance of the transport of an ephippium from Coniston or Winder-
mere. If this be so it goes to show that these lakes are not so isolated as
would appear, and that interchange of species is not infrequent. If, in spite
of such interchange, the Daphnias inhabiting the different lakes remain
distinct, it is evident that the local races are established by right of survival,
and it leads one to suppose that, difficult as they may be to separate, yet they
may represent fixed species. Our conception of species is determined mainly
by our ability to discriminate individuals by fixed and measurable standards ;
but not only are physiological differences largely ignored or are unmeasurable,
but two races separated only by minimal mean differences may be just us
real specific units as those distinguished by obvious characters in all indi-
viduals. The evidence goes to show that a single lake as a rule is inhabited
by one race of limnetic Daphnia only, and, although within such a race
extreme forms are commonly found, it is obviously of no use to give these
variants names, as has frequently been done in the past. But, with that
exception, it seems advisable in the present state of knowledge to recognize
all well-marked races as distinct rather than to treat all as variants of one
collective species. The question of real specific distinction cannot, I think,
be determined by laboratory experiment, neither is the fact that the winter
forms of a number of Daphnias can be shown to be morphologically
identical prove anything more than close relationship. If the many
forms of Daphnia are to be regarded as variants of one species originating
under the influence of external conditions, the reappearance in widely-
separated areas and under conditions which can hardly be imagined to be
identical of precisely the same form—e. g. /). galeata in Windermere and in
Central Asia—is difficult or impossible to understand. It must be admitted
that the whole problem of specific validity is raised by the genus Daphnia in
its acutest form, and is far from being solved.

DAPHNIA LACUSTRIS, Sars. a

1. Esthwaite and Hawes Water.

The Daphnias of Esthwaite illustrate remarkably well the impossibility of
drawing any definite distinction between the various forms now grouped
within the species D. longispina. While some individuals taken in Esthwaite
in July 1921 were indistinguishable from /). longispina forma rosea, having
the crest interrupted over the eye and the general form of that species, the
usual type of Daphnia present in that and in other collections helongs to
7). hyalina var. lacustris. J was at first inclined to suppose that two distinct
races were present, but the two forms grade so completely one into the other
that they must be regarded as one. The outline of the head may be
completely rounded, or may show a scarcely noticeable dorsal angle, while in

OF THE ENGLISH LAK® DISTRICT. 443,

others, and especially in young specimens, this angle is produced into a
small papilliform projection. The crest is generally continuous over the eye,
hut is often so inconspicuous as to be seen with difficulty, while sometimes it
is interrupted as in DJ. longispina.

An exactly similar Daphnia occurs in Hawes Water, but in such small
numbers that I have not had much material for examination. Neatly all the

AL Daphnia hyalina from Crummock, 29.9.22, Typical form for the lake.

B, Daphnia hyalina trom Crummock, 27.8.20, Head of female of Lilljeborg's
forma estivalis monstrosa.

U. Daphnia galeata from Crummock, Sept. 1912.

D. Bosmina obtusirostris from Wastwater, 30,9.22.

specimens seen have the head quite rounded, but occasionally there is a
slight trace of the dorsal angle so often seen in Hsthwaite specimens.

This form seems to be the same as that which is found in Loch Leven and
Loch Lomond in Scotland, and was deseribed by Richard (1896, p. 307) as
D), lacustris var. vicina, with the exception that the posterior spine in the
Lake District form is usually direeted more or less dorsally.

444 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

Burekhardt’s PD. hyalina forma lucernensis from the Lake of Lucerne is
evidently the same variety. In one individual from Hsthwaite the dorsal
projection of the head is denticulate as figured by Burckhardt for his forma
jurassica.

Derwentwater.

The Daphnias from Derwentwater rather closely resemble ). hgalina s. str.,
but should, I think, rather be included in D. lacustris by reason of the
concave front and rather higher head. The dorsal angle characteristic of
the form from Esthwaite and Hawes Water is occasionally, but very rarely,
seen in Derwentwater.

DAPHNIA GALBATA, Sars.

The typical form of D. galeata oceurs throughout the year in Coniston,
Windermere, and Ullswater, being most abundant in the former. Although
Dr. Pearsall’s samples cover every season in the year, I have not found
ephippial females in any of them, so that the species appears to be acyclic.
The same appears to be the case with the D. lacustris of Hsthwaite, Hawes
Water, and Derwentwater, but in Crummock and Buttermere there is a
distinct sexual phase in D. hyalina in October, though, even here, the species
persists through the winter.

Daphnia—Seasonal Variation.

The material at my disposal does not admit of any positive statement with
regard to seasonal variation of form, but such variation is very slight if it
occurs at all. With regard to J. galeata, in which it could most easily be -
detected, I have found on the one hand crests equally as high in winter as in
summer, and individuals with a quite round head in July and August. I[
have no doubt that a small variation of head-form, and of size of hody, could
be shown statistically with sufficient material, but the variation is certainly
not so pronounced as to be appreciable to the eye.

In this respect also the plankton of these lakes differs markedly from the
Danish and Central European lakes, and approaches the northern type.

BosMINA OBTUSIROSTRIS, Sars.

The identity of the northern races of B. coregoni with these occurring in
the alpine lakes is now established, and, aceording to Riihe, the united species
should be known under the name of B. coregoni-longispina. As, however,
the name B, longispina has generally been used in a more restricted sense,
and is so used by Lilljeborg, and as the name B. obtusirostris has been
universally used for the northern forms with short mucro, I prefer to retain
it in Lilljeborg’s sense.

In the Lake District this species occurs, often in great abundance, in nearly
all the lakes. Its apparent absence from Buttermere may be due to the fact
that the collections were made from the shore only ; but, although a single

OF THE ENGLISH LAKE DISTRICT. 445

specimen was found in Bassenthwaite in October 1921, it is certainly not
established there.

It would be of interest to ascertain if there are any constant differences
between the Bosminas of the different lakes, but this can only be done satis-
factorily by’measurements of large numbers of specimens carried out on the
system laid down by Burckhardt, and my material does not admit of such
treatment, since the necessary number of well-preserved specimens is not
available from all the lakes at the same seasons. Since there is very great
variability within the population of a lake with regard to length of antenna
and of mucro, the selection of single individuals for measurement or
illustration is likely to be misleading, but it is not entirely without value,

Tapie 10.—Measurements of single individuals of
Bosmina obtusirostris.

| Shell. Length | Length |
Date. | Length. | —- “a” — ot of
Length.’ Width. |antenna.; mucro.
Windermere ...... | 2I). vil. 63 743 784 310 | 81
Wsthwaite ........ Bix, | 726 (ales 928. | 96 |
Derwentwater ....| 17. v. “64 703 750 390 93
Ullswater ........ Tewviie |) 65 753 758 430 92 |
Hawes Water ....| ll. vi. | 6 716 783 SOOM OO
Coniston ........ 5.vii. | 68 | 706 | 647 | 404 | lio
Ennerdale ........ i8,i || CB | M2 | We | sees | es
Wastwater........ 30. ix. | 92 717 706 543 | 152
|Crummock........ 12. vil. | 89 707 752 505 | 123
|Floutern Tarn ....| 27. ix. 52 711 7ll ayy || 8s
Goats Water...... Sin |) aS 754. me | 28) |) 1) |
| | | |

and a table of measurements of individuals chosen as being so far as could
be judged typical of the population is here given. The example given from
Esthwaite is above and that from Ennerdale below the average size, but
typical of form. This table shows that the Bosminas of the two primitive
lakes Crummock and Wastwater are not only very much larger than all
others, but have longer mucro and antenna (text-fig. 3, D). On the other
hand, those from Ennerdale, which might be expected to be similar to those
from Wastwater, are, in fact, conspicuous for their very short antenna and
mucro and are not particularly large. Herein is another strong point of
difference between these two apparently similar lakes. In the remainder
of the lakes the differences shown between the Bosminas are not, I think,
outside the limits of their ordinary variation within a single population.
The specimens from Floutern Tarn and Goats Water are not only the
smallest, but lave the shortest mucro.

446 MR. R. GURNEY ON THE CRUSTACEAN PLANKTON

For the reasons given above I have also been unable to study the seasonal
variation. I have, it is true, made an attempt to do so with material from
Coniston, but the numbers available and sufficiently well preserved are not
enough to give a satisfactory result. They tend to show that there is a
variation, but that it is very small.

The species is found at all times of the year, and, except in Wastwater
and the tarns, there seem to be no pronounced sexual periods. Ephippial
females have been seen in collections from Windermere (Sept. and Oct. 1921),
Crummock (Oct. 1921), Derwentwater (Nov. 1921), and Wastwater (Sept.
and Oct. 1921 and 1922). In Wastwater alone do ephippial females form
any considerable proportion of the population. On Sept. 30, 1922, 31 per
cent. of females were ephippial, but not a single male was seen. In Coniston
and Ullswater there seems to be no sexual phase.

Bosmina Lonerrosrris, O. F. Muell.

This species onty occurs in Hsthwaite and Grasmere, both being lakes of
the “evolved” type; but, whereas it appears to be the only species of
Bosmina in Grasmere, in Esthwaite it is found in company with B. obtusi-
rostris, an unusual association which is also found in Lough Neagh and has
beeu commented on by Dakin and Latarche.

Miss Pratt states that it was common in Bassenthwaite in 1898, but it wa
not found in any of Dr. Pearsall’s collections from that Jake. Miss Baie
list of Crustacea from Bassenthwaite differs in so many respects from my
own that one can only conclude that conditions must have changed very
greatly since 1898.

The Esthwaite form is quite typical, but in Sept. 1921 the antenne of the
adults were found to be unusually short, and many were even of the form
described by Lilljeborg under the name of B. brevicornis Hellich. It
appeared in the collections first in April and increased in numbers till June.
In July numbers decreased, and in August it was very rare. In September
it was again, present in abundance, and large numbers of females bore
ephippia. It was not found in collections made in October and November.

LIVERATURE REFERRED ‘LO.

Brcx, C.—On some new Cladocera of the English Lakes, Journ. R. Mic. Soc. (2) 11, 1885,
Pandas

Brren, HE. A.—Plankton Studies on Lake Mendota. If. Trans. Wisc. Acad. xi. 1895,
p: 274.

Brapvy, G. S.—Notes on the Crustacean fauna of the English Lakes. Intellectual Observer,
xi. 1868, p. 416 *.

Brapy, G. S.—British species of Entomostraca belonging to Daphnia and other allied

genera. Nat. Hist. Trans. Northd. and Durham, xiii. 1898, p. 217.

* T have not been able to consult this paper.

2
at >

te

Lg G7, S— = = ——
=

CRUSTACEAN PLANKTON

\
yeTHN

\
—

FROM THE ENGLISH LAKES.

JouRN. LINN. Soc., ZOOL VOL. XXXV. PL. 23.

y/o S
eS es va
MAF j fy x
l Hy
} iif AA }
= = S=s
; C S| |
i“—~=s
vn ——<0))
WM ;
\
: —_\\ ez
—p
=i \\ N ~S

OF THE ENGLISH LAKE DISTRIC'. 447

Bourckuardt, G.—Faunistische und systematische Studien iiber das Zooplankton der
erdsseren Seen der Schweiz und ihres Grenzgebietes. Rev. Suisse Zool. vii. 1899.

Daxin, W. J., and Lararcun, M.—-The Plankton of Lough Neagh. Proc. R. Irish Acad.
xxx. 1913.

NkMAN, S.—Studien iiber die marinen Relikte der nordeuropdischen Binnengewiisser.—
II. Die Variation der Kopfform bei Zimnocalanus grimaldii (De Guerne) und
ZL. macrurus, Sars. Int. Rey. Ges. Hydrob. und Hydroz. vi. 1914,)p. 335,

GraeEteEr, A.—Die Copepoden der Umgebuny yon Basel. Rev. Suisse Zool. xi. 1903.

Grater, H.—Hin neuer Hohlencopepode Cyclops crinttus, n. sp. Zool. Anz. xxxiii. 1908,
p. 46.

Marr, J. .—The Geology of the Lake District. Cambridge, 1916.

Mri. H. R.—Bathymetrical Survey of the English Lakes. Geog. Journ. iv. 1894, p. 237 ;
vi. 1895, pp. 46 & 135.

Naumann, E.—Ueber die natiirliche Nahrung des limnischen Zooplanktons. Lunds Univ.
Arsskrift, N.F. Avd. 2, Bd. 14, no. 31, 1918.

PrarsaLt, W. H.—The aquatic vegetation of the English Lakes. Journ. of Ecology, viii.
1920, p. 163.

PrarsaLL, W. H.—The development of vegetation in the English Lakes considered in
relation to the general evolution of Glacial lakes and rock-basins. Proc. Roy. Soc.
ser. B, xcii. 1921, p. 259.

Pratt, HK. M.—The Entomostraca of Lake Bassenthwaite. Ann. Mag. Nat. Hist. (7) ii.
1898, p. 46.

RicHarp, J.—Revision des Cladocéres. Ann. Sci. Nat. Zool. (8) vy. 1896.

ScuEerrert, KE. Die Copepoden und Cladoceren des siidlichen Schwarzwaldes. Arch. f.
Ilydrob. iv. 1908.

ScovurFieLD, D. J.—Synopsis of the known species of British fresh-water Entomostraca.
Journ, Quek. Mic. Club, (2) viii. 1903, p. 531.

Warp, J. Crirron—The Geology of the northern part of the English Lake District. Mem.
Geol. Sury. London, 1876.

Warp, H. B.—A biological reconnaissance of some elevated lakes in the Sierras and
Rockies. Trans. Amer. Mic. Soe. xxv. 1904.

Worrrrecx, R.—Die natiirliche Nahrung pelagischen Cladoceren und die Rolle des Zentri-_
fugenplanktons in Siisswasser. Int. Rev. ges. Hydrob. und Hydroz. i. 1908,
p. 87l.

EXPLANATION OF PLATE 23.
Limnocalanus macrurus, Sars.

. Side view of female from Ennerdale Water.

. Fifth leg of female.

Fifth leg of male.

Outer branch of right fifth lee of male.

Side view of female from the Caspian Sea (LZ. grimaldii).

Side view of male from Lale Erie.

. Head of female from Insjin, Sweden.

Side view of last thoracic segment of male from Lake Milaren, Sweden.

J
oF wi

SIO

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Pi GEN NaAN. SOCIETY,

Vou. XXXV. ZOOLOGY. No. 237.

CONTENTS.

Page
I. Some Protozoa from the Soils and Mosses of Spitsbergen.

Results of the Oxford University Expedition to Spitsbergen,
No.27. By A. Sanpoy. (Communicated by D. Warp CutLeEr,
WelbS)) Cela 2H anniel ( Wexarsayerinest)) 5 nsdnoecsanee avasdaaseoc 449

II. On a Collection of Sponges from the Abrolhos Islands, Western
Australia. By Arraur Denpy, D.Se., F.R.S., F.L.S., Professor
of Zoology in the University of London (King’s College), and
Lestrze M. Frepertcn, M.Sc., Harold Row Student in the
Zoological Department, King’s College. (Plates 25-26.) ...... AT7

ILI. Report on Opisthobranchiata from the Abrolhos Islands, Western
Australia, with Description of a new parasitie Copepod, By
Cras. H. O’Donocuus, D.Sc., F.R.S.C., Professor of Zoology,
University of Manitoba, Canada. (Communicated by Prof.
W- dp Darin, DS. IIS.) (Blais Bir es),) Geccdaceascuscasne SIL

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PROTOZOA FROM SOILS AND MOSSES OF SPITSBERGEN, 449

Some Protozoa from the Soils and Mosses of Spitsbergen. Results
of the Oxford University Hxpedition to Spitsbergen, No. 27.
By H. Sanpon. (Communicated by D. Warp Curtmr, F.L.S.)

[From the Protozoology Department, Rothamsted.]
(Prater 24 and 6 Text-fieures.)
[Read 7th June, 1923.]

Tae protozoal fauna of Spitsbergen and of the neighbouring lands is in
some respects already fairly well known. As far back as 1869, Ehren-
berg (14) identified four species in material brought back from there, and
more recently Scourfield (29) and Penard (28) have published extensive
lists of collections from Advent Bay, Green Harbour, and Amsterdam Is.
The establishment of a biological station on the Murmansk coast enabled
Awerinzew (2-4) to carry out investigations of the protozoa of that region,
and his collections extended as far as Bear Island, the southernmost island
of the Spitsbergen Archipelago ; while Mereschkowski (19) and Levander (18)
have also identified numerous species from the northern parts of Finland,
Russia, and Siberia. Scourfield, Penard, and Awerinzew, however, confined
their attention entirely to the inhabitants of mosses, and in particular to the
testaceous rhizopods, and none of the authors since Ehrenberg have investi-
gated the soil-dwelling forms. It has recently been shown that in temperate
lands protozoa inhabit the soil in such large numbers that they must play an
important part in the activities of the soil (see especially 12); and a
knowledge of the soil protozoa from localities subject to extreme climatic
conditions therefore assumes considerable interest, not only in relation to the
general ecological problems of those regions, but also because of the light it
may be expected to throw on the influence of climatic conditions on the soil
organisms.

The material kindly placed at my disposal by the members of the Oxford
University Hxpedition consisted of three samples of mud from the brackish
pools near Bruce City described by Walton (83), five samples of soils from
various localities, and fifteen samples of mosses; and the supplementary
expedition of 1922 also brought back three samples of soil. The soil samples
were all packed in tightly-fitting tins which had previously been sterilised
by heat, and the mosses had been dried and sealed up in envelopes.

ZOOLOGY, VOL, XXXV. 33

LINN. JOURN.

450 MR. H. SANDON ON PROTOZOA FROM THE

Merrnops.

Cultures were made from all the samples on plates of nutrient agar, in
sterile tap-water, and dilute hay infusion, and were examined at intervals for
three or four weeks. For detecting rhizopod tests, soil or moss was taken
from the tap-water cultures after a few days, spread out on a glass slide
and examined with a low power, the mosses being first teased apart with
a needle.

RESULTS.
1. Mun.

These samples gave very disappointing results, possibly owing to excessive
heat having been employed during drying. Heteromita globosa was found
in all of them, and Satnowron mikroteron (un. g., n. sp.) (see Appendix) in the
mud from pond 5, the least saline of the three. Since the members of the ex-
pedition recorded the presence of peridinians in some of these ponds, cultures
in Miquel’s solution (1) were made at the kind suggestion of Dr. H. J. Allen
in the hopes of obtaining growths of these organisms, but, again, only the
ubiquitous Heteromita globosa and some diatoms were found.

In his notes made on the spot Mr. Elton records that Uroglena volvow was
very abundant in all the ponds in this district, and that there were also found
flagellates belonging to the genera Huglena and Glenodiniwm, and ciliates
probably belonging to the genera Vorticella, Carchesium, Dileptus, Prorodon,
and Colpoda. He makes the further interesting observation that whereas in
the Bruce City ponds, which were fed with drainage water from limestone
hills, Uroglena was very abundant, at Cape Bohemian, where the pools were in
sandstone, this species was absent, and its place was taken by large numbers
of Synura uvella and of peridinians. From the latter region his records also
include Glenodinium sp., Euglena sp., and three ciliates, probably Spathedium,

Nassula, and Ophrydium.

2. SorLs. |

As will be seen from the accompanying table (pp. 456-457), the 1921 soil
samples form an interesting series. Apart from rhizopod shells, soil 1
yielded only two species, and in fact three out of the six cultures made from
this sample were completely sterile. The number of species recorded from
sample 5, on the other hand, is quite comparable to that which would be
found in any British farm or garden soil, while the other samples fall between
these two extremes. Soil 1, however, contained great numbers of empty
rhizopod tests belonging to six different species, whereas in the other samples
these organisms were much less abundant.

An attempt was made to connect these striking differences with other
characteristics of the soils. The bacterial numbers, counted as they were

SOILS AND MOSSES OF SPITSBERGEN. 451

after the soil had been sealed in tins for some months, are of course no
indication of the actual number of bacteria in the soils at the time when the
samples were taken, but for purely comparative purposes they no doubt
correspond to real differences in the bacterial population of the soils. The
chemical and physical properties of the soils, however, throw no light on
these differences. The high numbers of organisms found in soil 5 may be
attributed to the sea-birds’ droppings with which it was manured; but, on the
other hand, soil 4 was similarly manured, and yet yielded fewer organisms
than soil 3, which was practically free from humus. The abundance of
rhizopod tests in soil 1 corresponds closely with what is found in peaty soils
elsewhere (20). It is possible that this does not really indicate a larger
active population of these organisms than in other soils, but is due simply to
the fact that in a soil, where microbial activity is small, such tests, coming
either from animals living in the soil, or being brought down from the
mosses ete. growing in it, disintegrate more slowly than in other soils, and
so accumulate in considerable numhers.

The 1922 samples were examined in a fresher condition than those of the
previous year, and on the whole yielded a considerably larger number of
living species. The good results obtained from sample 5 (1921), however,
seem to indicate that these earlier samples had not suffered appreciably as a
result of storage, and the difference in the results obtained from the two sets
of samples probably therefore corresponds with an actual difference in the
population of the soils im situ. Sample 3 (1922) was taken with a view to
confirming the results obtained from sample 1 (1921), in which both bacteria
and protozoa were so exceptionally few. Unfortunately it was not possible
to bring soil from the identical spot, and although the localities from which
the two samples were obtained appeared very similar and supported a similar
type of. vegetation, the analysis given in the table (p. 456) shows that the two
soils are not really comparable.

The abundance of protozoa in samples 1 and 3 (1922) is of particular
interest, in view of the high acidity of these soils. Such results, however,
are not unique, for an active protozoal population has been found by
Mlle. Perey in the soil of one of the Rothamsted experimental plots for
which the pH is 365; and S. M. Nasir (in some unpublished work also
done in this laboratory) has found that in artificial cultures active flagellates
ean tolerate a pH of 4:5, active ameebe a pH of 3:9, while ciliates can
grow even when the acidity is as high as that represented by a pH value
of 3:5.

3. Mosszs.

Twelve of the samples came from Klaas Billen Bay, and were composed of
the following mosses :——/Typnum (5 spp.), Cinclidium stygium, Orthothecium
chryseum, Swartzia montana, Bryum spp., Camptothecium nitens, Grimmvia

33"

452 MR. H. SANDON ON PROTOZOA FROM THE

commutata, and Brachythecium salebrosum, Most of the samples were com-
posed of a mixture of two or three of these mosses. The remaining three
samples came from Prince Charles Foreland, and were composed of
Rhacomitrium lanuginosum, R. canescens var. ericoides, and Dicranum

groenlandicum.

Flagellata.

Cercomonas ecrassicauda Alexeieff. In 8 samples from Klaas| and in 0 samples from Prince
Billen Bay, Charles Foreland.

bo

Orcomonas termo Kent. .:.... Site

” ” ” ” 1 ” 19 ”

Heteronuta globosa Stein ...... eal 53 » Selle 99 pe
‘5 lens Muller ........ See t | A A Pe 5 0 - ms »
Spiromonas angusta (Duj.) Kent. ,, 1 i ‘ =A 5 0 A rs os
Helkesimastix fecicola Wood-,, 1 3 7 * Piatt) ; 91 on

cock & Lapage

Phyllomitus sp.........-0--0+0 gd % 90 6 ” yo»
Anisonema minus (n. sp.) .-..+. peel 5 on o ea) ' 09
Entosiphon sulcatum (Duj.) Stein ,, 1 . ” - AO) ie 5 as
Sainvuron oru (0. g.,D.8p.) ..++ 5, 2 > 9% ra wo LG) 5 mp
Allantion tachyploon (n.g.,0. sp.) ,, 2 3 %p 99 3 0 » 3 i

Most of these species are identical with those found in the soil samples.
This is perhaps not surprising, as most of the mosses had a certain amount of
soil clinging to them, and no attempt was made to remove this before making
the cultures. Hence it is impossible to say whether the organisms found
naturally inhabited the surface of the moss plants or the soil in which the
latter were growing.

Rhizopoda.

Hartmanella hyalina Alexeieft .. In 3 samples from Klaas| and in 0 samples from Prince
Billen Bay, { Charles Foreland.

Ameba limicola Rhumbler .... ,, 1 4 40 = % (0) . 5 &

Sp.a(Cutler,Cramp,& Sandon) ,, 1 ; : 4 0) ae sieve ae

Amoeba vespertilio Penard ...... on Ll ” ” ’ NE) ys ”

FMC Cii7;ZCOLCN ORCC LNEy ran tane doe HY 5 Ay He bytes) * ie

a Gara) EMRE, 6.5.0.0.000 P(e) ; ; Ma) 90 . ”

, actinophora Auerb. .... ,, 1 si - a 0 =) < ys

Corycia flava Greef .......... Pawn 5 ‘ 5 a iL » 90 oh

» coronata var. simpler ,, 1 6 4 - e 0 Fr *

[ Penard

Diffugia pyriformis Perty ...... nie 5 9 ” ” 0 ; 4 99

PN Glace onan Wiener Pebtih i a op > 1 5 oe 5)

5 Gordie ID 506660008 a nG, i A os 5 3 oy » m

6 (Mendig IPE, So a oon rink al - 96 ip i 0 - “p 50

», constricta Khrenb....... 9 Wil : i % np ” ” 6

Centropyxis aculeata Stein ....,, 5 ” ” ” ” 0 ” ” ”

4 levigata Penard.... ,, 5 A ss 5 cs 0 ss of 5

Arcella vulgaris Whrenb, ...... Pe atS 09 5 % » 0 9 9 A

” arenaria Greeiattneree ” q ” ” ” ” 1 ” ” ”

SOILS AND MOSSES OF SPITSBERGEN. 453

Nebelu collaris Leidy .......... In 6 samples from Klaas | and in 0 samples from Prince
Billen Bay, { Charles Foreland.
Quadrula irregularis Aycher.... ,, 8 , 3 i * 0 5 re 5
Euglypha ciliata Ehrenb, ...... ite we “4 ae 5 Wi 0 “ “ 44
a strigosa Leidy........ aye vill i aa _ 5 0 rH ;
” ” var. glabra ” 1 ” ” ” ’ 2 ” ” ’
[ Wailes
i UPS JY oho ddp ace. 9 KES) 4 ‘ e - il “ o mA
Bi tuberculata Duj....... » O “ ass i. i iL i 0 p
Assulina muscorum Greef .... ,, O 0 " 0 7) 2 » ” ”
Cyphoderia sp.,Schlumb. ...... » @ in ‘ Fs i 0
Campascus triqueter Penard .... ,, 3 ns ms A 0) ; :
Trinema enchelys Whrenb....... BD . Fy 0) A
- » va GalemD . Il 0 55 5 5
[ Penard
s complanatum Penard.. ,, 1 a 7 we a 2) x ;
» Umeare Penard ...... » @ 90 90 re mm Al ” » ,
Corythion dubium Taranek...... yr) “ a es i 2 99 on
Chlamydophrys stercorea Cienk... ,, 1 i : 5 il ; *

Ciliata.

Of the samples from Klaas Billen Bay, six contained Colpoda cucullus
O. F. Muell., two contained Balantiophorus minutus Schew., and the following
forms occurred in one sample each :—Balantiophorus elongatus Schew.,
Blepharisma laterita Ehrenb., Cyclidium glaucoma Ehrenb., Enchelys sp..
Oxytricha sp., Stylonychia sp. In the soils from Prince Charles Foreland
only Colpoda cucullus O. F. Muell. was found.

Discussion.

(1) The Influence of Arctic Conditions on Protozoa.

Previous investigators have differed considerably as to the influence of
arctic conditions on protozoa. Although the ubiquity of most of the com-
moner fresh-water and moss-dwelling species is well known, it is rather
difficult to believe that the same species can live in arctic as in temperate
and even tropical lands without showing some local peculiarities. It was
perhaps this consideration which led Ehrenberg (15) to create new specific
names for the rhizopod shells obtained from Greenland, which names have
since apparently all been discarded. Scourfield (29) remarks on the numbers
of abnormal tests (especially in the species Euglypha ciliata and Nebela col-
laris) found in his Spitsbergen material, and Penard (26) similarly comments
on the tendency to produce deviations from the type in the rhizopods from
the still more extreme regions of the antarctic. On the other hand, whereas
Penard’s antarctic specimens were mostly smaller than the average (espe-
cially in the genera Assulina, Corycia, Corythion, and Euglypha}, Awerinzew (2)

454. MR. H. SANDON ON PROTOZOA FROM THE

found that tests from north polar regions were on the whole larger than the
average.

The present investigation throws little light on this question. Several
species (e.g. Arcella arenaria, Trinema complanatum) included some
exceptionally big specimens, but others were on the average rather below
the normal size (Centropyais aculeata, C. levigata, Nebela collaris). Obser-
vations of this kind are, however, very inconclusive unless based on statistical
data ; and as no such data from temperate regions are available for com-
parison, there was no point in devoting time to obtaining them from the
present material. It seems, however, fairly certain that there is ‘no general
tendency to either a larger or a smaller size in the Spitsbergen rhizopods,
and that none of the species found formed any distinctive local varieties.

(2) Species recorded.

The rhizopods commonly found living on mosses have been divided by
Penard (24) into two groups which he calls respectively “formes banales ”
(i.e. forms commonly found in mosses but not especially adapted to such
a life) and “formes caractéristiques” (¢.e. forms specially adapted to
living among mosses and rarely found elsewhere). The former group
contains 26 species, of which 21 have now been recorded from Spitsbergen ;
but of the second group, only 2 out of 17 have so far been found there, and
one of these is Amaba terricola, which seems ont of place in this classifica-
tion, since, so far from being specially adapted to living among mosses, it
occurs frequently, as its name implies, in soil. The absence of the “ formes
varactéristiques ”? is probably due in part to the fact that they include
a number of naked amcebee which do not normally form cysts, and which
therefore may have been present in the moss when it was collected, but died
before the samples were examined.

Levander (18) and Heinis (16) have already drawn attention to the close
similarity existing between the rhizopod fauna of the high latitudes and
that of the high altitudes of the European Alps, as recorded by Zschokke
(37), Daday (13), and Heinis (46). The present investigation completely
confirms this analogy, but unfortunately the data available do not appear to
be sufficiently complete to extend it to other groups of protozoa. The
generalisation that Heinis makes with respect to the high Alpine protozoa,
however, seems to apply equally to the Spitsbergen records: namely, that
the severity of the climate does not result in the occurrence of local species,
but simply eliminates the less adaptable forms, with the result that the
population is composed of the most cosmopolitan forms. The possibility of
the existence of true aretic species is, however, not entirely ruled out by these
considerations (except in the testaceous forms), since such species, if they
exist, may have died out completely before the material could be examined,

SOILS AND MOSSES OF SPITSBERGEN. 4

One new flagellate is described which does not seem to have been recorded
elsewhere, and it is possible that some of the organisms which occurred in
the cultures in too small numbers to be identifiable may have represented
other new species.

The ubiquity of these moss- and soil-dwelling protozoa has an additional
significance, since they have no means of avoiding the extremes of tempera-
ture or of moisture to which they are exposed: Planktonic organisms can
avoid extreme heat or cold by sinking to greater depths, and worms, many
insects, etc., can similarly burrow to more equable layers if the surface of
the soil becomes too hot or too cold except, of course, in regions where
the sub-soil is frozen. But such vertical migrations of the land-living
protozoa, if they take place at all, must be very much more restricted,
and the wide distribution of these organisms must therefore indicate
a wide range of tolerance of external conditions on the part of the
organisms themselves or of their cysts (though resistant cysts are not at
present known for all the creatures in question). There remains the possi-
bility of the existence of physiological strains differing solely in their powers
of resisting extreme conditions. This is, in fact, very probable, since many
unpublished observations made in this laboratory have demonstrated the
presence in tropical soils (Egypt, Hast Africa, West Indies, etc.) of species
identical with some occurring in Spitsbergen, which therefore in natural
conditions tolerate a wider range of temperatures than is possible in
laboratory cultures.

The identity of the protozoal fauna of Spitsbergen with that of temperate
lands is strikingly paralleled by the observations made by Barthel (5) on the
soil bacteria of Greenland. An examination of soils collected by the
Rasmussen Expedition from Disko Is. (Lat. 70° N.), from Cape York (Lat.
76°. N.), and from the extreme northern coast showed that these soils, just
like ordinary Huropean soils, are capable of initiating the processes of
nitrification, denitrification, ammonification, and fermentation of urea ; and
though nitrogen fixation was not definitely proved, there was some evidence
of its occurrence. These results would no doubt apply equally in the
somewhat milder climate of Spitsbergen. ‘Thus all the evidence at present
available points to the qualitative identity of the microbiological activity
of soils from Spitsbergen and from temperate lands. As to the quantitative
aspects of the question, observations made on the spot would be essential.

MR. H. SANDON ON PROYOZOA FROM THE
of Aye % Nin ¥, loss
Sample. Locality. Description. fe dry dry on
matter. Gu eata
matter. ignition.
1921.1. [Low hill near |Black peat contain- 901 1:53 51:9
coast, Prince | ing roots, etc. Dry
Charles tundra _-vegeta- |
Foreland. tion (Silene, Salia,
mosses, etc.). |
2. |Klaas Billen Fine black loam | 94:9 0°75 27-9
Bay. from roots .of | |
| Andromeda. |
| |
3. Klaas Billen Stony calcareous soil 98°9 0-152 32°8
Bay. from roots of Dryas;
practically nohumus,
but much dry plant-|
remains. |
4. |Moss plain be-Damp mossy peat, | 88:4 2°85 69°5
lowVogel Hook) much manured by,
(Pr. Charles | sea-birds’droppings. |
Foreland).
5. | Vogel Hook. [Fine brown peat, 91:2 2°47 A473
much manured by
sea-birds’droppings.
1922. 1. |Grassy slopes |The grass here is 48°6 1-485 88:94
on bird cliffs, more luxuriant than !
Green Mt., Ice} anywhere else on
Fiord. the island.
2. |Bogey patch on| Vegetation 44e1. 0:4.40 22°42,
hillside at | Sphagnum ete.
Advent Valley.
Ice Fiord.
3. (Dry tundra at Vegetation Dryas, | 79°71 0:378 13:82
foot of hills, | Salix polaris,Silene,|
Advent Valley.| ete. Well-rotted |
mould.
|
|

SOILS AND MOSSES OF SPITSBERGEN.

No. of | |
pH. nngiew2 FLAGELLATA. CILIATA. RHIZOPODA. |
per gramme |
(millions). |
lots) | 13 Heteromita lens. Colpoda cucullus. ‘Difilugia globulus, D. con-|
stricta, EHuglypha levis,|
| Assulina muscorwn,
| Hyalosphenia minuta.
72 4°87 Oicomonas termo, |Balantiophorus mi-/Ameba radiosa, Difflugia,
Heteromita globosa,| nwtus, Uroleptus | constricta, Centropywis
Cercomonas sp. mobilis, and an |! levigata, Quadrula
| unidentified species.| inregularis. |
8:0 12°65 Heteromita globosa,\Colpoda cucullus, |Difflugia constricta, |
Cryptomonas Cyclidium glaw- Difflugia sp. |
i) obovoides. coma, Plewrotricha |
sp., Vorticella
microstoma.
66 6°25 Heteromita globosa, |Colpoda cucullus, |Ameba limaz, A. radiosa,
H. lens. Gonostomum affine.| A. terricola, Difflugia
globulus, D. constricta.
6:0 545 ‘\Oicomonas termo, |Colpoda cucullus, |Dimastigameba gruberi,
Heteromita globosa,) Uronema sp., Oxy-| Hartmanella hyalina,
H. lens, Cercomonas}| tricha pellionella,| A. diploidea, A. terri-
crassicauda, Helke-| Blepharisma cola, A. sp., Nuclearia
simastia fecicola, | laterita, and one | simplex, Chlamydophrys
Copromonas sp., unidentified species.) stercorea, Difilugia glo-
Cladomonas fruti- bulus, D. constricta,
cosa. Centropyxis levigata,
| Buglypha strigosa.
i
39 — \Oicomonas termo, |Chilodon cucullulus,|Ameba limaz, A. radiosa,
Heteromita globosa,| Enchelys (probably| A. diploidea, A. terricola,
' by lens, E. farcimen), Vorti-| A.actinophora, Nuclearia
| Cercomonas sp. cella microstoma, | simplex, Chlamydophrys
i Oxytricha pellion- | stercorea, Trinema
ella, and 3 unidenti-| lineare, Diflugia con-
fied species. stricta, D. globulus,
D. fallax, Nebela lageni-
formis, EBuglypha sp.,
Assulina muscorum.
6:0 = Oicomonas termo, |Enchelys (probably |Ameba limax, Trinema
Heteromita lens, E. farcimen). | lineare, T. enchelys,
Cercomonas sp., Diflugia constricta,
Sp. 6. D. globulus, Euglypha
| strigosa (forma glabra).
52 = Oicomonas termo, |Cyclidiwm glaucoma|Dimastigameba grubert,

Heteromita globosa,
H. lens, Cercomonas
sp., Phalanstertwm'
solitarium, Spiro-
monas angusta,

| Petalomonas an-
gusta, Allantion
tachyploon, <Ani-
sonema minus.

and 2 unidentified
species.

A. radiosa, Actinophrys
sp., A. diploidea, Tri-
nema Vineare, Difflugia
constricta, Huglypha
levis, Assulina mus-
corum.

457

458 MR. H. SANDON ON PROTOZOA FROM THE

APPENDIX.

Noves ON THE SPHCIES FOUND.
1. Flagellata.

Cercomonas.—This is one of the commonest of soil Flagellates. The usual
species is C. crassticauda Alexeieff, but the specific characters are not very
well defined, and other species such as C. longicauda may also have been
present in some of the cultures.

Heteromita.—This genus is extremely abundant in all soils, commonly
occurring in two forms which agree closely with the description of H. lens
Miller and H. globosa Stein (see especially 35). The former is a small
organism, lenticular in shape and from 4 to 6m in length, though some-
times very small globular individuals occur only 2 or 3m in diameter. A
pair of flagella arise at the anterior end, one directed forwards being about
twice the body-length, and the other, which is normally trailed passively,
being a little longer. The nucleus is not visible in the living animal, and
the contractile vacuole is also not usually apparent, but when present lies
just behind the middle. When swimming the body is turned at a sharp
angle to the direction of motion, and the anterior flagellum lashes vigorously
through a wide amplitude. At times the tip of the flagellum adheres to the
glass slide, and both flagella wave actively, so that the animal jerks vigorously
from side to side without progressing. When stained with hematoxylin a
typical vesicular nucleus appears surrounded by a dense mass of peripheral
chromatin, especially at the posterior side. The cyst is small (diameter about
4 yw), thin-walled, and usually homogeneous in structure, though sometimes
the protoplasm appears concentrated round the wall, leaving a vacuole in the
centre, to one side of which there may be a small refractile granule.

Fleteromita globosa is larger (length 8 to 12), and more variable in
shape. Usually the anterior end is rounded and the posterior end slightly
narrower, but sometimes this is reversed. At times the protoplasm becomes
quite plastic at the posterior end, which may then adhere to the glass slide
while the anterior part moves round slowly in circles owing to the move-
ments of the anterior flagellum. In this condition the hinder part may
become drawn out into a long filament before the animal breaks free. The
flagella arise together just behind the anterior extremity, the shorter one
being about equal to the body-length and directed forwards, while the other,
which is usually trailed passively, is about 1} times as long. A nucleus is
clearly visible near the anterior end, and when stained shows much less
peripheral chromatin than in //. lens. The contractile vacuole is central.
The movements of the body are vibratory; it is not turned outwards
so sharply as in //. lens, and the anterior flagellum lashes through a smaller
amplitude. The cyst has a thin outer wall which is usually raised a little
from the thicker inner wall. The protoplasm of the cyst is hyaline, and con-
tains a large very refractile granule against which is a big vacuole. This

SOILS AND MOSSES OF SPITSBERGEN. 459

cyst is well figured by Puschkarew (27), but the two walls are always separable,
and the granule and vacuole are sometimes poorly developed or even absent.

Though these two species seem quite distinct, apparently intermediate
forms both of the active and eneysted stages are frequently found, the identi-
fication of which is very difficult, and in cultures of either species a few
abnormally large or small individuals are usually found which might be
mistaken for members of the other species. Until conclusive observations in
pure cultures have been carried out, there consequently remains a doubt
whether we are actually dealing with two true species or with two extreme
forms of a single very variable species.

SALNOURON MIKROTERON* (n. gen.,n. sp.). (Pl. 24. figs. 1-6.)

A brief description of this organism under the name of Sp. y has already
been published (12). It is a small flagellate from 3 w to 6 in length with
a single flagellum, slightly longer than the body, which arises just behind the
anterior end and is directed backwards. The anterior end is rounded, and
usually the body is slightly crescent-shaped (fig. 2) (the side from which the
flagellum arises beine slightly coneave) and tapers somewhat towards the
posterior end. In these specimens the maximum width is about half ithe
length or less, but sometimes individuals occur in which the posterior end is
swollen, and the side from which the flagellum arises is sigmoid in outline,
the concavity being limited to the anterior end. In normal cultures the
shape is quite persistent, but in the deeper parts of hanging-drop cultures
plastic individuals sometimes occur adhering to the ylass slides (figs. 4, 5).
The plastic individuals contain ingested bacteria, which are absent in the
normal free-swimming form, the nutrition of the latter being apparently
wholly saprophytic. The nucleus is not visible in the living animal, probably
owing to its small size, and no contractile vacuole has ever been observed.

The movements are very like those of a small Heteromita, since the
flagellum is trailed quite passively while the animal progresses at a con-
siderable speed with a rapid vibratory motion, the body being turned at an
angle of about 20° to the direction of motion. The progression is in a
straight line (not spiral), and the animal does not turn on its axis as do most
organisms which possess an anterior flagellum.

When stained, a large vesicular nucleus becomes visible in the anterior
part. The basal granule varies considerably in size and position, at times
lying against the anterior side of the nuclear membrane and at other times
being quite free from it and apparently not connected to it by any rhizo-
plast. Most characteristic of this species, however, is a large mass of
plastin, often visible in the living animal as a refringent body, which lies
just behind the nucleus or at times forms a ring completely enclosing it
(figs. 1-3). In a dividing animal this body divides immediately after

* caivw, to wag.

460 MR. H. SANDON ON PROTOZOA FROM THE

the basal granule, but its size and shape seem to depend on the general
metabolism of the organism rather than on the phase of its life-cycle, for
dividing specimens can be found in which two nuclei are present, each
entirely surrounded by a ring of plastin, whereas in other cases the plastin
divides to form two centrally-placed spherical masses. The changes which
this mass undergoes have unfortunately up to the present obscured the
division changes of the karyosome, the difficulty of observing which is ren-
dered greater both by the extremely small size of the organism and by the
fact that the karyosome itself varies considerably in size and at times appears
to be entirely lost (fig. 3), as in the case of Herpetomonas (Leptomonas)
jaculans as described by Berliner (7).

In old cultures thin-walled spherical cysts rarely reaching as much as 3
in diameter appear (fig. 6). The protoplasm is uniform and shows no charac-
teristic features, but tends to be denser round the periphery of the cyst
and vacuolated in the centre. Staining reveals a simple excentrically-
situated nucleus, the mass of plastin apparently having been lost before
encystment.

This species is very widely distributed. It is common in the Rothamsted
soil, and has also been found in samples of soil from Africa, the West Indies,
Azores, ete.

SAINOURON OXU (n. sp.). (PI. 24. figs. 7-10.)

This species resembles the foregoing very closely. It is, however, con-
siderably larger, being from 10 or 12 in length ina young culture, but
diminishing to 6 w or 7 in old cultures. The anterior end, instead of being
rounded, is generally drawn out into a curved beak, at the base of which the
flagellum arises (fig. 7). This beak is occasionally bent or fattened (fig. 8),
and the whole body is in fact more plastic than in S. mkroteron. A large
vesicular nucleus lies in the anterior part of the body, and is generally sur-
rounded by a complete peripheral ring of chromatin (fig. 8), but frequently
the ring is broken and in its place one or more deeply-staining masses
(recalling the plastin masses of S. mikroteron) lie against the posterior part
of the nuclear membrane (fig. 7). The basal granule from which the flagellum
arises is situated at the base of the beak when the latter is present, or slightly
to one side of the anterior extremity when the beak is not present. Usually
the granule lies too close against the nuclear membrane to be distinguishable
(fig. 9), but at times there is a considerable space between them (figs. 7, 8).
In spite of careful examinations of cultures of all ages, no contractile vacuole
has been found. Nutrition appears to be saprophytic as in S. mikroteron,
which species S. ovw also resembles very closely in its movements. It shows,
however, one interesting peculiarity in its behaviour in that when a drop of
a culture is put on a slide and covered with a coverslip for examination, the
organisms remain perfectly quiescent for some time. ‘lhe first signs of
activity usually appear about one minute after the preparation of the slide,

SOILS AND MOSSES OF SPITSBERGEN. 461

but even after four minutes there are usually still a considerable number
that have not resumed their movements.

The cysts (fig. 10) are thin-walled, spherical, and from 4 to 5p in
diameter. ‘Ihe contents are granular, the granules usually being arranged
in aring. When stained, a nucleus with karyosome and peripheral blocks
much as in the active stages is visible.

It has not been found outside Spitshergen, where it occurred in two samples
of moss, both from Klaas Billen Bay.

ALLANTION TACHYPLOON * (n. gen., n. sp.). (PI. 24. figs. 11, 12.)

This organism, though it resembles Sainowron in having a single flagellum
which arises just behind the anterior end and is directed backwards, has such
a characteristic mode of progression that one feels constrained to create a
new genus for it. In size it varies from 7 w to 14 (usually 8 u-10u), and
the width is about half the length. The shape is very constant, being
equally rounded at the two ends and usually slightly crescentic. It is
enclosed in a firm pellicle, and no trace of pseudopodia can ever be
found ; in fact, the “only deviations from the typical shape are the
occurrence of shortened, almost spherical individuals, or of organisms which
have their concave side (7. e. the side from which the flagellum arises) com-
pressed, appearing in consequence more transparent than the rest of the
organism when they are seen in side view, and causing the animals to
become triangular in cross-section instead of circular, as is more normal.
Both these forms are commonest in rather old cultures. There is a simple
vesicular nucleus with some peripheral chromatin in the anterior half of the
body, and one or more deeply-staining spherical masses, presumably meta-
bolic products of some kind, are often found in various parts of the body
behind the nucleus. There is no contractile vacuole. The flagellum is about
half as long again as the body, and is very much stouter than in such flagel-
lates as Sainowron or Heteromita. It is invariably directed backwards.
When the animal is progressing, the flagellum is applied for its whole length
to the substratum, while the body itself is turned upwards at a sharp angle.
Neither the flagellum nor the body of the animal shows any trace of vibration
or movement of any kind, and yet the animal glides rapidly forwards, covering
a distance of two or three times its own length per second. The organism
whose movements resemble this most nearly is Helkesimastia Woodcock &
Lapage (86), from which the present species differs in being quite without
an anterior flagellum (the presence of which in Helkes/mastix I have been able
to confirm, but which plays no appreciable part in the animal’s locomotion).
It also differs in the absence of a contractile vacuole, and in the presence of
a firm pellicle as a result of which the flagellum is quite free except at its
point of insertion instead of adhering to the body as it does in Helkesimastia.
This mode of progression, depending presumably on some surface action

* d\dayriov, a little sausage ; raynmAoor, swiftly-sailing.

462 MR, H. SANDON ON PROTOZOA FROM THE

between the flagellum and the surface to which it is applied, may however
occur in other organisms having what has so far been regarded as a passive
trailing flagellum. It is impossible, for instance, to watch a Cercomonas for
long without concluding that its characteristic gliding motion is not produced
by the tractellar action of the anterior flagellum, for the motion continues
without a break even when this flagellum is temporarily motionless or
recovering from its streke ; and, further, the mean position of the flagellum
often forms a wide angle with the direction of motion of the organism, which,
on the other hand, is always in a line with the posterior flagellum. The
movements of Cerecomonas and Helkesimastiv are in fact so similar (in a
casual examination it is easy to mistake Helkesimastia for a small Cerco-
monas) that it seems probable that they are both produced in the same
way and that the motive power in both, as in Allantion tachyploon, is really
the so-called trailing flagellum.

When Allantion is not moving in this way, it adheres by the tip of the
flagellum to the slide and jerks violently from side to side without progressing
as a result of repeated flexions and extensions of the flagellum.

Remains of ingested bacteria are sometimes found, so that nutrition is
apparently holozoic though the actual method of feeding has not been
observed. No cysts of this species have been identified.

Though this organism does not appear to have been described before, it
occurs very commonly in soils both from Hurope and from the tropics.

ANISONEMA MINUS (n. sp.). (PI. 24. fig. 13.)

This species is quite typical of the genus in all except size. It is usually
from 6» to 7p in length and never exceeds 9 uw, whereas the lowest limit for
any Anisonema given by Senn (80) is 11. It is strongly flattened and has
a longittidinal groove on the under side. In outline it is egg-shaped, but
slightly asymmetrical at the anterior end. The anterior flagellum is stout
and about equal to the body-length, while the posterior flagellum is rather
more slender and about twice as long. They arise from separate basal
granules a little way behind the anterior end. The contractile vacuole is
central, but a pharynx has not been definitely discerned. There is a large
elliptical vesicular nucleus lying to one side of the body, containing a karyo-
some, which though usually spherical is sometimes irregular in shape, and
a ring of peripheral chromatin. The movements are slow and very steady,
being produced by the movements of the anterior flagellum, which lashes
through a wide amplitude.

This species occurred in one sample only. I have, however, found it in
soils from the West Indies, so that although apparently never abundant, it
seems to be a widely-distributed soil form.

Cladomonas sp.—This occurred in only one culture from soil sample 5
(1921). The monads are spherical to ellipsoidal in shape, from 5 y to 7 w in
length, and with two forwardly-directed flagella, each about 15 to 1? times the

SOILS AND MOSSES OF SPITSBERGEN. 463

body-length, inserted at the anterior end. Hach monad is situated at the end
of a tube usually with only the flagella projecting, but sometimes the cell
itself protrudes to some extent. The tube is colourless and transparent, and
is quite empty except for the organism at its tip; in fact, but for the cocci
adhering to it, it would easily pass unnoticed. Its composition was not
investigated, but it is flexible and yields to the movements of the organism.
Occasionally one finds an unbranched piece of tube, 50 ~ or more in length,
with a single flagellate at its end, but more usually the tube branches, though
not with the regular dichotomy of C. fruticosa Stein. Usually, however, the
distal parts of the tube are surrounded with a dense mass of bacteria, pro-
tozoal cysts, ete., so that its ramifications cannot be fully followed out. The
flagella are held at an angle of about 20° to one another and vibrate rigidly
without bending. These movements are so rapid that the flagella can then
only be detected by the characteristic movements of the surrounding bacteria,
etc., one or twoof which usually adhere to the tip of each flagellum, and are
seen to be whirling round in a small circle. Occasionally the motion stops,
and the animal remains at rest for a few seconds. ‘The effect of the whirling
of the flagella is to drive small bacteria towards the mouth of the tube, where
they adhere both to the walls of the tube itself and to the body of the animal,
which is consequently always the centre of a small mass of the organisms.
Occasionally a single monad is found adhering to the glass slide without any
tube. In this condition the flagella vibrate just as usual, causing cocci to
come against the body, to which they then adhere.

This organism differs from C. fruticosa Stein only in the less regular
branching of the tube, a feature which hardly justifies the creation of a new

specific name.

PHALANSTERIUM SOLITARIUM (n.sp.). (PI. 24. figs. 14,15 & text-figs. 1, 2.)

This organism differs. in several respects from the two established
species P. consociatum Fresenius and P. digitatum Stein. It usually appears
in cultures in the form of free-swimming monads 10m to 15m in length
(text-fiv. 1) and more or less spherical in shape, though sometimes rather
elongated, and often distorted by the presence of large vacuoles. At the
anterior end is a single flagellum, three or four times as long as the body,
which is surrounded at its base by a narrow “collar,” into which a eyto-
pharynx opens. The flagellum is very characteristic, the proximal part
(about one-fourth of the whole length) being stout and fairly rigid, while
the distal part tapers to a very fine and flexible whip. Unlike the flagellum
of the Craspedomonads, which is always a pulsellum, this flagellum acts as a
tractellum, so that the creature swims with its collar directed forwards.
A reversal of this mode of swimming has never been observed. The mode
of swimming enables the species to he easily recognised, since the whole
organism vibrates rapidly as it were about a pivot situated near the hinder
end of the body. Oftena few cocci adhere to the rigid part of the flagellum,

464 MR. H. SANDON ON PROTOZOA FROM THE

and these are consequently seen vibrating vigorously a short distance in
front of the animal itself. The progression is rapid and in a straight line
without any tendency on the part of the organism to turn on its axis. It
follows that the movement of the flagellum (which is too rapid to be

Fie. 1. Fic. 2.

contr vac

Phalansterium solitarium, X 2500.

1. Normal living form with collar.
2. Hlongated form from an old culture.

visible) must consist of a lashing from side to side, and is not the typical
spiral movement of the Craspedomonads. ‘The contractile vacuole

is not
usually visible, but is situated just behind the centre of the body.

SOILS AND MOSSES OF SPITSBERGEN. 465

In old cultures the organisms often lose their spherical shape and become
very narrow, tapering particularly at the anterior end (text-fig. 2). In this
condition the collar is difficult to observe, and often appears to be completely
merged in the cell protoplasm. EHyen in the more typical spherical form
the collar is occasionally invisible, and at death it is invariably completely
retracted.

In the living organisms the nucleus, when it is visible, is composed of a
single spherical karyosome surrounded by a narrow clear zone. Unfor-
tunately it has only been possible to obtain a single stained preparation in
which the karyosome was often vacuolated (Pl. 24. fig. 14) or distorted by the
pressure of large cytoplasmic vacuoles (PI. 24. fig. 15). The fixation of these
specimens appeared to be satisfactory, but until further preparations have
been obtained, it is impossible to say whether or not these appearances are
abnormal.

In its sedentary phase the organism becomes surrounded by a thick
gelatinous layer. Unlike P. consocratum and P. digitatum it does not form
any definite colonies, though frequently a number of these gelatinous
capsules are formed side by side. HWrequently also two or more cells occur
in a single capsule, but the process of multiplication has not been observed.
The flagellum projects through the wall of the capsule, which has no well-
defined opening, and its fine whip-like distal portion lashes vigorously with
a spiral movement, which causes cocci etc. to be whirled towards its base,
where they accumulate in a mass which quivers slightly with the movements
of the flagellum. The whole gelatinous capsule, in fact, becomes encrusted
with cocci etc. so that the details of the organism inside are not easily
distinguished. The collar appears to be rather smaller than in the free-
swimming form, and may frequently be entirely lost. Neither in the sessile
nor in the free-swimming phases lave any bacteria been observed inside the
collar, and the mode of nutrition is therefore unknown. While in this
gelatinous capsule, the organism loses its collar and flagellum and becomes
Boetecah in a thick wall, forming a rather angular cyst, which, however, has
none of the ridges showed in Ghankkometts Beis ( (11).

This species ie also been found in svils from Rothamsted and from the
West Indies.

Of the other flagellates recorded, Otcomonas termo Hhrenb. is one of the
most numerous and most ubiquitous soil organisms. Spiromonas angusta(Duj.)
Kent, Helkesimastiv fecicola Woodcock & Lapage, and Phyllomitus sp.,
though much less abundant, are not uncommon in soils from other regions.
Petalomonas angusta (Klebs) Lemm., found in soil sample 3 (1922), does
not seem to have been found in the soil before, though the nearly related
Copromonas subtilis Dobell is not uncommon.

There were in addition a small number of unfamiliar forms which appeared
in one or more of the cultures, but in such small numbers that a satisfactory

LINN. JOURN.—ZOOLOGY, VOL. XXXV. a4

466 MR. H. SANDON ON PROTOZOA FROM THE

examination was not practicable, and the attempts to obtain stronger sub-
cultures failed. It is possible that some of these may have been new
species. Of these forms the following two deserve mention, since they
have each been found in one or more other localities ; and although it has not
so far been possible to obtain the stained preparations necessary for an
adequate description, the appearance of the living organisms is quite

Fie, 3. Fie 4,

aw
contr: Vac. \: /

I

contr. vac

Sp. 6, X 26500; freehand drawings from life.

3. Free-swimming form with anterior pseudopodium.
4, Ameeboid form,

characteristic. It is hoped that a satisfactory identification or deseription:
may be published later.

Sp. 6. (Text-figs. 3, 4.)

In the flagellate condition (text-fig. 3) this organism is very plastic, and
its shape is therefore rather indefinable, but typically the anterior end is

rounded, and posterior end tapers to a point. Quite spherical individuals,

SOILS AND MOSSES OF SPITSBERGEN. 467

however, are not uncommon. TFinger-like pseudopodia may be formed
at all parts of the body, but they are commonest at the posterior end.
Two flagella arise in a slight pit at the anterior end, of which one is
directed forwards and is between 1 and 14 times the length of the
animal, and the other, which is a little longer, is directed backwards or
slightly to one side, but is quite free from the body except at its point
of insertion, thus differing from the corresponding flagellum in Cercomonas.
In small specimens the relative lengths of the flagella are often greater,
up to three times the body-length. A vesicular nucleus is usually clearly
visible in living specimens at or just in front of the middle of the organism,
and a single contractile vacuole can occasionally be observed near the hinder
end. The usual length is about 15, though occasionally small specimens of
6 to 10u occur. The movements are usually jerky, both the flagella being
held more rigidly than in most flagellates, the anterior one vibrating
vigorously.

The ameeboid stage (text-fig. 4) is less commonly observed. One flagellum
is retained which is directed sideways and continues to vibrate vigorously
while the animal moves forward with a single broad pseudopodium, the
posterior end being rounded. As in the swimming stage, the nucleus is more
or less centrally placed and is easily visible, but the contractile vacuole now
lies anteriorly between the ecto- and ento-plasms.

This species resembles the genera Cercobodo Krassilstchik and Dimastig-
ameba Blochmann rather closely in many respects. From the former,
however, it is distinguished by the true amoeboid movements of the sessile
form, while the fine radiating pseudopodia which Blochmann mentions as
being sometimes formed by Dimastigameba have not been observed.

It occurred in only one of the present soil samples, but has been found on
several occasions in small numbers in soil from Harpenden and from the
Azores, W. Indies, and other localities.

Sp.e¢. (Text-figs. 5, 6.)

Tn its general appearance and movements this organism is somewhat like
Allantion tachyploon (see p. 461). The length is usually about 18 «, though
individuals as small as 10 or as large as 24 mw occasionally occur. The
width is about half the length. The sides are straight and the ends both
rounded. In section it is circular or slightly flattened ‘“ dorso-ventrally.”
The shape is persistent and the body not flexible, but finger-like pseudo-
podia may occur at either end. These are usually about 4 pu to 6 w long, but
occasionally a much longer one is formed. A stout flagellum, slightly
longer than the body, arises at the extreme anterior end, and is directed
backwards as in Allantion. In most individuals a very short second flagellum,
about 24 to3p in length, can be detected. Near the base of the flagella
are two prominent contractile vacuoles, but the nucleus is usually indis-
distinguishable in the living organisms, and, as already mentioned, no

34*

468 MR. H. SANDON ON PROTOZOA FROM THE

observations of its cytology have been made. The hind end of the body
often contains a considerable number of granules, apparently the remains
of food, though the actual ingestion of bacteria has not been observed. The
movements are very similar to those of Allantion, the long flagellum being
applied closely to the glass slide, and the whole animal gliding quickly and
steadily forwards. When the short second flagellum was visible it moved
vigorously from side to side, but its movements were not sufficiently powerful
to impart any vibration to the animal as a whole. The characteristic
quiescence in a newly-prepared slide, noted in the case of Sainowron oxu
(p. 460), is shown to an even more marked degree by this species, as much as
ten minutes sometimes elapsing before movements are resumed. If the drop
of culture fluid is spread out on a slide but not covered with a coverslip,
activity reeommences much more quickly.

Fre, 6.

2. Ciliata.

None of these require any comment except Uronema sp., which was found
in soil sample 5 (1921). The numerous specimens found appeared identical
with those obtained in cultures of sea-weeds from Plymouth, and it is
possible that they were really marine organisms introduced into the soil in
the droppings of sea-birds. Unfortunately at the time I was not aware of
Schewiakoff’s and Buddenbrock’s (8) work on this genus, and not having
made drawings, cannot say whether the organisms found were U. mari-

num Duj. or the very similar species U. schewiakofi Buddenbrock, which
appears to be a true fresh-water form.

3. Rhizopoda.

Amoeba limav.—The identification of the different species of “imax” amosbe
is usually only practicable when the complete life-history is known, or at
least when both the active and the encysted stages are available. In the

SOILS AND MOSSES OF SPITSBERGEN. 469

cultures in which only active individuals could be found the identification is
consequently incomplete. The common soil species Nwgleria (Dimastigameba)
grubert (Schardinger) Alexeieff was found in two of the soil samples, and
the very similar form Hartmanella hyalina Alexeieft in three of the mosses.
A characteristic small species described in a recent paper (12) as sp. a also
occurred in one of the moss samples. Unfortunately this organism does
not grow very well in cultures, and consequently it is still impossible to
complete the description of it, or to name it satisfactorily.

Amaba limicola Rhumbler.—This species occurred in only one culture and
was smaller than usual, being only about 20 w in diameter when rounded.

Ameba vespertilio Penard.—Penard records this species as being present
“en assez grand abondance ” in his Spitsbergen material, but in the present
investigation it was found in only one moss sample. The specimens examined
were nearly all in the radiate form when first observed, but passed over into
the more active non-radiate form when transferred to a slide for closer.
examination. in the latter condition the average length was about 60 p.

One sample of moss from Prince Vharles Foreland gave a few specimens
of a smaller amoeba which may belong to this species. When active this
animal is constantly and rapidly changing its shape, but does not progress.
The pseudopodia are numerous and are of two kinds

narrow finger-like ones
reaching about 20 in length, and other very fine ones which are eyen
longer. At times the amabee become rounded and motionless, with very
short finger-like pseudopodia projecting all over the body. The diameter is
then from 18 to 30. There is apparently only one contractile vacuole,
which is situated near the middle of the body and at times reaches a very
considerable size.

Ameba terricola Greet occurred in three soil samples and in one moss.
They were all perfectly typical except in size, in which respect they mostly
agreed with those found by Penard. The specimens from the mosses were
usually about 50 ~ in diameter and those from the soils even smaller (20 1).
Soil sample 4 (1922) contained a few full-sized ones of 120 in diameter,
with a nucleus of 354x18p and a contractile vacuole which at tinies
reached a diameter of 50. Though encystment is unknown in the European
members of this species, it apparently does sometimes occur, for the speci-
mens found in the material brought back by the British Antarctic Hxpedition
of 1907-1909 were all encysted (25). No cysts were found, however, in the
material.

Ameba striata Penard.—This beautiful amoeba occurred in its typical form
in one of the moss samples. When progressing steadily its dimensions were
about 60% by 364. The large contractile vacuole (diameter 12 wz) is charac-
teristic of the species, but in the present case it was almost invariably
single. Once only was it observed being formed by the coalescence of three
smaller vacuoles.

A single small specimen (length 30) was found in another sample trom
the same locality.

470 MR. H. SANDON ON PROTOZOA FROM THE

Ameba actinophora Auerb.—This organism, as it appeared in the cultures,
agreed closely with the figures and description of A. actenophora given by
Cash, but the envelope was much less well defined than in those figured by
Penard (22) under the name of Cochliopodium actinophorum. The usual
appearance is of a small granular amoeba completely surrounded by a wide
ectoplasmal border except during the extrusion of feecal matter, when the
granular endoplasm extends to the surface at the hinder end. In this con-
dition it changes shape quickly or moves forward slowly, the appearance
being somewhat like that of a broad limax except for the wide clear border
of ectoplasm. The anterior end is wider than the posterior, and the con-
tractile vacuole is anterior. At other times progression ceases, and long,
pointed pseudopodia arise from the endoplasm, which projects right through
the ectoplasm. The length of the amaba varies from 24 « to 36, and the
pseudopodia may reach about 15 yw in length.

Corycia flava Greef.—Though empty, rolled-up tests probably belonging
to this species were observed in many of the moss samples, living specimens
occurred in only two (one from each locality). They were quite typical
and the size was about 135 p.

Several specimens of Corycia coronata var. simplex (Renard), which is now
generally regarded as a variety of C. flava ($2), occurred in one of the other
samples from Klaas Billen Bay. Diameter of shell 105 w to 120 p.

Digugia.— As in Scourfield’s and Penard’s samples, 1). constricta Khrenb.
was much the commonest testaceous rhizopod found. The numerous speci-
mens recorded as 1). globulus Duj. were all empty tests, and it is therefore
impossible to be sure whether they belonged to this species or to the genus
Phryganella Penard, as the two are distinguishable only by the form of the
pseudopodia. The diameter was usually between 33 and 50, but in one
sample very large specimens (diameter 120 ~) were found which were more
probably tests of Phryganella nidulus Penard. D. fallax Penard was also
very common. The size was generally about 60 w in length by 54 w in greatest
width, with an aperture of between 256 and 30. The smallest individual
recorded was 30 long and the largest 72 p.

D. lwida Penard, the scarcest of the Difflugias found, occurred in only
one sample. The size was usually about 54 w in length by 27 in greatest
width. One specimen reached a length of 80 w.

D. pyriformis Perty was not common, and varied from 87 to 105 u in
length.

Centropyais.—Although the two species of this genus were about equally
widely distributed among the samples, the individuals of C. devigata Penard
were considerably the more numerous. They were mostly rather small speci-
mens, the diameter of the shell usually lying between 60 and 80 y, with a
height of 45 to 60 and an aperture of from 25m to 45 w in diameter.
A single very large specimen (144 in diameter and 72 in height) was

SOILS AND MOSSES OF SPITSBERGEN. 471

found which differed from the smaller ones in having a transparent yellowish-
brown chitinous test free from sand-particles.

The specimens of C. aculeata Stein were also on the average slightly
smaller than usual, varying from $4 to 120 in diameter and ila aper-
tures of from 20 to 38. ‘This species showed considerable variation in
form. The commonest type agrees closely with Cash’s var. spinosa, as the
spines were relatively long and the mouth distinctly lobed. This type included
the largest specimens found (diameter 114 to 120). In others the mouth
was circular and the spines very short, often in fact reduced to mere knobs.
The smallest specimens found, those with a diameter of from 85 p to 95 p,
were almost invariably spineless, and no doubt belong to the C. aculeata var.
ecornis of Leidy.

Arcella.—Typical tests of A. urenaria Greef varying in diameter from
724 to 180 were by no means uncommon. ‘The tests of A. vulgaris Ehrenb.,
though clearly differentiated from the others by the perfectly regular domed
appearance without any border when seen in side view, were decidedly
larger than usual, being between 100m and 190, in diameter.

Hyalosphenia minuta Cash.—A very few specimens of this species were
found in one of the soils. Average dimensions: length 34, greatest
breadth 20, mouth 10,.

Nebela collaris Leidy.—Length 84 to 135, greatest width 48 to 72 p,
aperture 184 to 30u. A broken test was also found which apparently
belonged to this species, and which when complete must have attained a
length of about 160. In structure the tests all resembled that figured by
Leidy (pl. xxiv. fig. 11). Scourfield also found some pitted like Hyalosphenia
or like Leidy’s pl. xxiv. fig. 12.

Nebela lagenifor mis Samar —This is deseribed by Cash (10) as a definitely
sub-alpine species. Typical but rather small specimens were found in one of
the soil samples, the dimensions being: length 103m to 105, breadth
48 to 544, aperture 21 to 25, length of noel 33 to 36p.

Quadrula irregularis Archer.—This species was abundant in most of the
mosses from Klaas Billen Bay, and showed considerable variation both in
size and shape. They mostly belonged to the variety globulosa Penard, and
had a diameter of from 25 to 45 and an aperture of from 9p to ld5yu. But
several were more elongated and frequently not quite symmetrical, belonging
therefore to Penard’s s variety oblonga.

Euglypha—The specimens found were all typical and require little
comment.

E. ciliata Ehrenb. was the commonest species in the mosses from Klaas
Billen Bay, and varied from 70 to 90 in length and from 51 to 60m in
greatest width.

LH. strigosa Leidy was also common. Most of the specimens belonged to
the variety glabra Wailes, and varied in length from 63 to 96, in greatest

472 MR. H. SANDON ON PROTOZOA FROM THE

width from 38 to 66m, and with apertures between 15pm and 24m wide.
Two ciliated specimens only were found with the dimensions: length 56m
and 65, greatest width 33 and 43, and apertures 15 and 31p.
ti. tuberculata Duj. occurred in one sample only. The dimensions were :
length 56, greatest width 27 u, aperture 12 4, body-scales (elliptical) 7» long.
This is no doubt the . alveolata found by Penard and Scourfield.

FE. levis Perty.—This species was not at all common and most of the
specimens found were damaged, so that the identification was in many cases
not very satisfactory. The dimensions of those found in the mosses were :
length 39 to 60, greatest width 24 to 40m, and aperture Ilp to l5p.
The specimens found in the soils were smaller, some being only 18 long.

Assulina muscorum Greet (A. minor Penard and A. seminulum (pars)
Leidy).—Typical specimens varying from 45 to 54 in length were abun-
dant in two of the moss samples from Prince Charles Foreland; in fact, their
presence constituted the most striking difference between these mosses and
those from the other locality. They were also quite numerous in some
of the soils, but these specimens were rather more variable in size (length
33 to 53m) and in shape, which was frequently asymmetrical. The soil
specimens were also quite colourless, but as they were all dead it was
impossible to decide whether this was due to the usual brown membrane
having been destroyed by the soil solution or whether they really belonged
to a colourless variety.

Cyphoderia ampulla var. vitrea Wailes.—Only a few individuals of this
species were found, and in most cases it was impossible to distinguish the
structure of the test, which appeared quite homogeneous. In one case,
however, regular polygonal scales could be detected. The specimens were
uniformly small, the length varying from 50 to 56 instead of the normal
100 to 120 (22), with a greatest width of 254 to 27 and an aperture of
10u to 12m. The usual yellowish tint was also absent and the tests (which
were all empty) were perfectly colourless.

In each of three moss samples a single specimen of a larger yellowish
retort-shaped test was found. Two of these measured 75, in length and the
third was nearly 170u. These tests showed no trace of any structure and
were all flexible, being in fact dented and creased when found. The section
appeared to be circular, but it was impossible to be quite certain of this
owing to the damage that they had suffered. They probably belonged to the
species Campascus triqueter Penard, which is confined elsewhere to lakes of
glacial origin and is consequently regarded (24) as a relict of a true arctic
fauna. The occurrence therefore of this otherwise lacustrine form in arctic
mosses is of particular interest, and it is unfortunate that no undamaged
specimens were found.

Trinema.—T. enchelys Hhrenb. and 7. lineare Penard were both quite
common, though the former species was not recorded from Prince Charles

SOILS AND MOSSES OF SPITSBERGEN. 473

Foreland. ‘The former varied from 33 to 65 in length and the latter
from 16m to 30. Trinema complanatum Penard was also common in two of
the moss samples from Prince Charles Foreland, and is no doubt the form
observed by Scourfield (1/400” long) which he compares with Leidy (pl. xxxix.
figs. 41, 54, 61, 63). They were mostly between 45 and 50 in length by
27 to 30m in width, but a few were considerably larger, ranging up to
75 long. One sample from Klaas Billen Bay contained, in addition
to all these three species, a few large specimens nearly 70 in length of
T. enchelys var. galeata Penard, a form very like 7. complanatum, but
distinguished by the presence of lines on the ventral surface curving
backwards from the aperture to the sides of the test.

Corythion dubium Taranek.—This was very scarce even in the two samples
in which it was present. They were mostly from 404 to 50 in length, but
one reached 65.

Chlamydophrys stercorea Cienk.—The classification of the Chamydophrys
group of rlizopoda has recently been revised by Belar (6), but, as his
ditferentiation is based on the details of nuclear division, it could not be
followed here, so the older and wider name C. stercorea is therefore retained.
Judging from the size (length between 24» to 27m by 19m to 21), the
specimens found probably belong to the species C. schaudinni Schussler.

Nuclearia simplex Cienk—The individuals found were all colourless, and
appeared identical with those found in English soils.

Actinophrys sp.—Only a single specimen was found, and unfortunately a
detailed examination was not possible. The diameter was 21. Penard
obtained from Spitsbergen a small new species of Raphidiophrys (LR. bruniz)
of similar dimensions, to which this specimen may have belonged. The two
genera are distinguished by the presence of minute spicules in the gelatinous
envelope of Raphidiophrys which are absent in the case of Actinophrys, but
as they are not usually visible in the living animal, it is impossible to say
whether they were present or not in the specimen found.

SUMMARY.

The protozoa contained in 3 samples of mud, 8 samples of soil, and 14
samples of mosses from Spitsbergen have been investigated. An abundant
fauna was found, most of which was identical with that occurring in the
soils and mosses of temperate lands.

Seven new species of flagellates are described, of which however five have
also been found in soils from non-arctic regions.

In conclusion, I wish to thank Mr. G. C. Sawyer for the soil analyses,
Mr. HE. M. Crowther for the pH estimations, Miss L. M. Crump for sug-
gestions of names for the new species described, and especially Mr. D. W.
Cutler, under whose supervision the whole of the work has been carried out.

AT4 MR. H. SANDON ON PROTOZOA FROM THE

LITERATURE QUOTED.
(1) Auten, E. J. 1910.—Q.J. M.S. lv. part 2.
(2) Awrrinzew, 8. 1906.—Trav. Soc. Imp. Nat. St. Pétersh. xxxvi. (Abstracted in
J. Roy. Mier. Soc. 1907, p. 684.)

(3) 1907.—Ueber einige Stisswasser-Protozoen der Bireninsel. Zool. Anzeig.
Xxxi. p. 245.
(4) 1907.—Ueber die Siisswasserprotozoen der Insel Waigatsch. Zool. Anzeic.

xxxl. p. 306,
(5) Barruen, Cur. 1922.—Recherches hactériologiques. Meddelelser om Grgnland
Ixiv. Copenhagen.
(6) Berar, K. 1922.—Untersuchungen tiber Thecambben der Chlamydophrys-Gruppe.
Arch. f. Prot. xlii.
(7) Breuer, B. 1909.—Flagellaten-Studien. Arch. f. Prot, xv.
(8) BupprnsBrock, W. von. 1920.—Beobachtungen iiber einige neue oder wenig bekannte
marine Infusorien. Arch. f. Prot. xl.
(9) Burscuur, O. 1883-1887.—Protozoa, in Bronn’s ‘ Klassen und Ordnungen des Thier-
reichs”’
(10) Casu, J., & Watnes, G. H. 1905-1921,—The British Fresh-water Rhizopoda and
Heliozoa. Ray Soc., London,
(11) CimnKowsky, L. 8. 1870.—Arch. f. mikr. Anat. vi.
(12) Curter, D. W., Crumr, L. M., & Sanpon, H. 1922.—A Quantitative Investigation
of the Bacterial and Protozoan Population of the Soil. Phil. Trans. B,
vol. 211.
(13) Dapay, J. 1897.—Beitrage zur Kenntnis der Mikrofauna der Tatra-Seen. Termész.
Fizetek, xx. .
(14) Eurunspere, C.G. 1869,—Die mikroskopische Lebensyerhiltnisse auf der Oberflache
der Insel Spitsbergen. Monatsh. Akad. Berlin.
(15) ——— 1874.—Die zweite deutsche Nordpolarfahrt in den Jahren 1869 und 1870,
vol. ii. Leipzig.
(16) Hernts, F, 1910.—Systematik und Biologie der moosebewohnenden Rhizopoden . ..
der Umgebung von Basel. Archiv f. Hydrobiol. und Planktonkunde, v.
(17) Kent, W.SavittE. 1880.—A Manual of the Infusoria.
(18) Levanpsr, K. M. 1901.—Beitrage zur Fauna und Algenflora der siissen Gewasser
an der Murmankuste. Acta Soc. Faun. Fenn. xx., viii.
(19) Merescukowsk1, C. von. 1878.—Studien iiber Protozoen des nordlichen Russlands.
Arch, f. miky, Anat. xvi.
(20) Mixture, P. E. 1887.—Studien tiber die natiirlichen Humusformen. Berlin.
(21) Pascuur, A., & Lemmurmann, HE. 1918, 1914.—Flagellata, in Die Siisswasserflora
Deutschlands. Jena.
(22) PenarD, E, 1902.—La Faune Rhizopodique du bassin de Leman. Geneva.

(23) 1903.—Notice sur les Rhizopodes du Spitsbergen, Arch. f. Prot. ii.

(24) 1909.—Sur quelques Rhizopodes des mousses. Arch. f. Prot. ix.

(25) 1911.—Rhizopodes d’eaux douce. Report on Scientific Investigations of the
British Antarctic xpedition, 1907-1909. I. London.

(26) ——— 191].—Rhizopodes d’eaux douce. Deux. expédition Antarctique frangaise.

Paris.

(27) PuscHKarEw, B. M. 1913.—Ueber die Verbreitune der Siisswasserprotozoen durch
die Luft. Arch. f. Prot. xxviii.

(28) Scurwiakorr, W. 1893. Ueber die geographische Verbreitung der Sitisswasser-
protozoen. Mem. Acad. St. Pétersb. xli.

Sandon. Journ. LINN. Soc, ZOOL. VOL XXXV. PL. 24

SY
PHighley Lith C Hodges & Son. imp

PROTOZOA MROM SPT SIDING

SOILS AND MOSSES OF SPITSBERGEN. 475

(29) Scourrietp, D. J. 1897.—Contributions to the Non-marine Fauna of Spitsbergen.
I. Rhizopoda. Proc. Zool. Soe. xlv.

(30) Senn, G. 1900.—Flagellata in Engler and Prantl’s ‘Die natiirlichen Pflanzenfamilien,’
I. Abt. 10. Leipzig.

(31) Stern, F. von. 1859-1883.—Der Organismus der Infusionsthiere. Leipzig.

(32) Waites, G. H. 1912.—Freshwater Rhizopoda and Heliozoa from the States of New
York, ete. J. Linn. Soe., Zool. xxxii.

(33) Waxron, J. 1922.—A Spitsbergen Salt-marsh. Journ. Ecol. x.

(34) Winson, C. W. 1916.—On the Life-history of a Soil Ameba. Univ. Cal. Publ.,
Zool. vol. xvi.

(35) Woopcock, H. M. 1896.—Observations on Coprozoic Flagellates. Phil. Trans. B,
vol. 207.

(36) Woopcocg, H. M., & Lapace,G. 1915.—Observations on the Life-cycle of a new
Flagellate. Proc. Roy. Soc. B, vol. [xxxviii. ;

(37) Zscuockr, F. 1900.—Die Tierwelt der Hochgebirgsseen. Neue Denksch. d. allg.
schweiz. Geselly., 1900.

EXPLANATION OF PLATE 24.

All the specimens were fixed in Schaudinn’s alcoholic sublimate + 5 per cent. acetic
acid and stained in Heidenhain’s hematoxylin. The figures were drawn with the camera
lucida and a Zeiss 1/12" objective and are x 5000, except fig. 15 which is x 2500.

Figs. 1-8. Sainowron mikroteron, typical forms.
4,5 as s sessile forms with ingested food bodies.
Fig. 6 » cyst.
7. Sainouron oxu, typical active form.
8 0 » showing plastic beak.
9. - 5, beak withdrawn.
10. D 9 cyst.
Figs. 11, 12. Adlantion tachyploon.
Fig. 13. Anisonema minus.

Figs. 14,15. Phalanstertum solitarium.

In fig. 15 only the thickened basal part of the flagellum is shown.

SPONGES FROM THE ABROLHOS ISLANDS. 477

On a Collection of Sponges from the Abrolhos Islands, Western Australia.
By Artaur Denpy, D.Sce., F.R.S., F.L.S., Professor of Zoology in the
University of London (King’s College), and Lustizm M. Freverick, M.Sc.,
Harold Row Student in the Zoological Department, King’s College.

(PLATES 25 & 26.)
[Read 21st June, 1923. ]

INTRODUCTION.

Tue collection of Sponges made by Professor Dakin at the Abrolhos
Islands is a very rich and, in some respects, a very remarkable one. It
comprises 48 determinable species, of which 12 are regarded as new. The
collection is especially rich in Calcarea, and includes a number of fine
specimens of that extremely rare and interesting sponge Grantiopsis cylin-
drica; a new species of the no less rare and remarkable genus Lelapia, and
a new genus of Leucascide. ‘The Tetraxonida, as usual, form the bulk of
the collection, but do not include any very striking novelties ; there are,
however, two rather remarkable new species of Huceratosa.
The following is a complete list :—

Order CALCAREA.

Family Homocmiip&.
1. Leweosolenia grisea n. sp.

bo

Leucosolenia protogenes Haeckel.

Family Levcascipa.
Ascoleucetta compressa n. gen. et sp.
Leucetta chagosensis Dendy.
Leucetta microraphis Haeckel.

or H CF

Family LevcaLrips.
6. Leucaltis clathria Haeckel.

Family Sycerrip”.
7. Sycon gelatinosum de Blainville.

Family HErERoPip”.

8. Vosmaeropsis mackinnoni n. sp.

Family GRANTIIDA.
9. Grantiopsis cylindrica Dendy.
9a. Grantiopsis cylindrica Dendy var. fruticosa nov.

478 PROF. A. DENDY AND MISS L. M. FREDERICK ON

Family LELApnp&.
10. Lelapia antiqua n. sp.

Order TETRAXONIDA.
Sub-order HoMoOSCLEROPHORA.

Family PLaKINIDé.
11. Dercitopsis minor Dendy.
12. Dercitopsis mammillaris (lendenfeld).

Sub-order ASTROTETRAXONIDA,

Family STELLErTIDa.

13. Stelletta brevis Hentschel.

14. Stelletta debilis Thiele.

15. Stelletta sigmatriena Lendenfeld.
16. Ancorina australiensis (Carter).
17. Ancorina brevidens n. sp.

18. Aurora row: Dendy.

19. Asteropus simplex (Carter).

Family Eryiipa.
20. Erylus procimus Dendy.

Family Donatup”.
21. Donatia robusta (Bowerbank).
22. Donatia multistella (Lendenfeld).

Family Coonprostps.
23. Chondrilla australiensis Carter.

Sub-order SIGMATOTETRAXONIDA.

Family HapLoscLeripa&.

24. Reniera aqueductus Schmidt.
25. Reniera cribricutis Dendy.
26. Reniera permollis (Bowerbank).

teniera spp.
27. Petrosia dura (Nardo) Vosmaer.
28. Halichondria phakellioides n. sp.
29. Chalina palmata (iamarck).

Chalina sp.

Pachychalina sp.
30. Ceraochalina multiformis var. manaarensis Dendy.
31. Phlaodictyon abrolhosensis n. sp.

SPONGES FROM THE ABROLHOS ISLANDS. 479

Family DesMACIDONID2.
32. Pseudoesperia carteri n. sp.
33. Pseudoesperia trichophora n. sp.
34. Hsperella plumosa Carter.
35. Biemna tubulata Dendy.
36. Eehinodictyum bilamellatum (Lamarck) Ridley.
37. Anchinoé fictrtioides n. sp.
38. Dendoricella schmidti (Ridley).

Family CLAVULIDA.
39. Trachycladus levispirulifer Carter.
40. Stgmosceptrella fibrosa Dendy.
Al. Spirastrella vagabunda Ridley.
42. Aaptos aaptos (Schmidt).
43. Polymastia mammullaris (O. F. Miiller) Bowerbank.

Order EUCERATOSA.
Family DarRwiNeLLip”&.
44, Megalopastes arenisibrosa n. sp.

Family SPoNGELIID®.
45. Spongelia dakini n. sp.
46. Psammopemma CPASSUNE Carter var.

Family Sponciipaz.
47. HHippospongia intestinalis (uamarck).
48. Coscinoderma pyriforme Lendenfeld var. «.
48 a. Coscinoderma pyriforme Lendenfeld var. @.

As might be expected from the position of the Abrolhos Islands, the
sponge fauna is mainly intermediate in character between that of the more
westerly Indian Ocean and that of the more easterly Australian coasts, but it
contains a small element apparently derived from the North, viz. Lelapia
antiqua, nearly related to the Japanese L. nipponica ; Stelletta debilis, pre-
viously known only from Ternate, and Ancorina brevidens, very closely related
to Ancorina amboinensis from Amboina ; while Eehinodictyum bilamellatum,
though widely spread on the eastern coasts of Australia, seems to be a

_characteristically tropical or sub-tropical species.

We have to record our indebtedness to Professor W. J. Dakin, D.Sc., for
the opportunity of studying this valuable collection; to Mr. M. Burton, M.Sc.,
for determining the two species of Donatia, a genus to which he has devoted
special attention; to Mr. Charles Biddolph, the skilled assistant in the
Zoological Department of King’s College, for the photographic illustrations,
and to Mr. J. C. Dendy for help in section-cutting.

480 PROF. A. DENDY AND MISS L. M. FREDERICK ON

1. LeucosoLenia GRISEA n. sp. (PI. 25. fig. 1; Pl. 26. fig. 1.)

There are two specimens in the collection, the larger (R.N. VI. 21), which
1s attached to a rock by its base, is plicate, consisting of vertical lamellee
which fold and branch in such a way as to lie parallel to one another
(Pl. 25. fig. 1). The total height of the specimen is 30 mm., the total width
25 mm., and the average thickness of the lamelle 1°5 mm. The second
specimen (R.N. LV. 3a) consists of a curved lamella, forming an imperfect
funnel 20 mm. in height and 10 mm. in diameter at the top.

The surface is smooth, even and sieve-like in appearance, owing to the
pseudopores which are thickly and evenly scattered all over it. The oscula,
which are marginal in position, are about 1 mm. in diameter and havea slightly
prominent margin. Colour in spirit grey ; texture firm and compact.

‘Lhe pseudopores, which are nearly circular and measure about 0°26 mm.
in diameter, pierce the pseudoderm, which is about 0:07 mm. in thickness,
and lead into wide, irregular interspaces lying between the branching and
anastomosing Ascon tubes. The prosopyles are accompanied by groups of
black pigment granules which doubtless indicate the porocytes, as in the
case of Leucosolenia coriacea as described by Minchin [1900], and in that
of Leucosolenia cavata Dendy [1891]; it is the presence of this jet-black
pigment which gives the sponge its characteristic grey colour. The granules
are not confined to the porocytes, but are also found scattered in the flat
epithelium. The secondary Ascon tubes open into the main exhalant canals,
which are also lined by basinucleate collared cells, and open to the exterior
by true oscula.

The canal system conforms to type B. (Dendy [1891].)

The skeleton consists of triradiate and quadriradiate spicules arranged
irregularly in one or more layers in the walls of the Ascon tubes; the tri-
radiates are more densely packed in the thick pseudoderm.

Spicules :—(1) Triradiates (Pl. 26. fig. 1a); equiangular ; rays measuring
about 0°14 mm. by 0:013 mm.; straight, conical, rather bluntly pointed ;
usually equiradiate, but the basal sometimes a little longer than the oral rays.

(2) Quadriradiates (Pl. 26. fig. 106); rather scarce, like the triradiates
but with the addition of a long, slender, slightly curved and sharply pointed
apical ray which projects into the gastral cavity.

This species is readily recognised by its characteristic external form and
its grey colour, due to the black pigment granules.

Register Nos. and Localities. 1V. 3a, Wooded Isle ; VI. 21, Sandy Isle.

2. LmuCOSOLENIA PROTOGENES Haeckel. (Pl. 25. fig. 2.)
(For discussion and synonymy vide Dendy [1891 ].)
There is in the collection a single small specimen (PI. 25. fig. 2) of this
common Australian species.

Register No. and Locality. VILL. 4, Sandy Isle.

SPONGES FROM THE ABROLHOS ISLANDS. 481

Genus ASCOLEUCEITA n. gen.

Leucascidee with a well-developed dermal cortex densely packed with large
and small triradiates and pierced by weil-defined inhalant apertures guarded
each by a fringe of hair-like oxea projecting radially inwards from
within the margin. The elongated and much branched flagellate chambers
are more or less radially arranged around the wide exhalant canals, which
open by numerous oscula.

3. ASCOLEUCETTA COMPRESSA n. sp. (PI. 25. fig. 3; Pl. 26. figs. 2,3, 4,5.)

The sponge, which measures 25 mm. in height, 22 mm. in greatest width,
and hasan average thickness of 4 mm., forms an erect, compressed, irregular,
more or less lobulated mass. The surface is smooth but uneven, and sieve-
like in appearance owing to the presence of numerous small, circular, inhalant
apertures, about 0-2 mm. in diameter, which are closely and uniformly scattered
all over it. The oscula, which measure about 1 mm. in diameter, are arranged
singly and are for the most part marginal. Colour in spirit light greyish
brown; texture firm and compact, rather hard.

The inhalant apertures (Pl. 26. fig. 5) pierce the well-developed dermal
cortex, which is about 0°13 mm. in thickness, and lead into large, well-
defined inhalant canals which run in between the flagellate chambers. The
latter are much elongated and copiously branched ; they tend to be radially
arranged round the larger exhalant canals, with their blind ends directed
more or less at right angles towards the dermal surface (PI. 26. figs. 3, 4).
The chambers, whose walls are pierced by numerous prosopyles, open into
wide exhalant canals which lead up to the vents. The collared cells are basi-
nucleate and confined to the radial chambers, being absent from the exhalant
canals.

The skeleton consists of triradiates and quadriradiates, for the most part
equiangular and equiradiate. The dermal cortex (Pl. 26. fig. 5) contains
numerous large triradiates arranged tangentially, intermingled with small
triradiates which are packed in a dense feltwork around the margins of the
inhalant apertures. ‘These apertures are guarded by a fringe of hair-like
oxea projecting radially inwards from the margin in small groups (PI. 26.
fig. 5). There are no large spicules in the interior of the sponge, small tri-
and quadriradiates being scattered closely but irregularly in the walls of the
radial chambers and exhalant canals, the facial rays lying tangentially,
while the slender apical rays of the quadriradiates project into the chambers
and into the exhalant canals.

Spicules :—(1) Large triradiates (PI. 26. fig. 2a) ; with conical, straight
or slightly crooked, gradually or sometimes rather abruptly sharp-pointed
rays, measuring about 0°35 by 0:052 mm.

(2) Small triradiates (Pl. 26. fig. 26); with straight, conical, bluntly
pointed rays, measuring up to 0:14 by 0:03 mm. All the triradiates are

LINN. JOURN.— ZOOLOGY, VOL. XXXV. 35

482 PROF. A. DENDY AND MISS L. M. FREDERICK ON

equiangular and more or less equiradiate, but sometimes the rays become
slightly unequal.

(3) Quadriradiates of three kinds: (a) (Pl. 26. fig. 2c), regular, very
abundant, like the small triradiates with the addition of a slender, curved,
often crooked, apical ray ; the facial rays measure about 0°15 by 0:035 mm. :
(b) (Pl. 26. fig. 2d), sagittal, with a wide oral angle and a slender, back-
wardly directed, very slightly curved apical ray ; the oral rays (which
measure up to 0°13 by 0°02 mm.) are sometimes slightly curved, the basal
ray is slightly shorter than the oral and straight and conical ; these spicules
are not very abundant and are probably confined to the exhalant canals and
the oscular margin: (c) (Pl. 26. fig. 2e), sagittal, small, very irregular,
usually with curved, crooked rays, measuring about 0-1 by 0-019 mm.
(b) and (c) have only been found in boiled-out preparations.

(4) Trichoxea (Pl. 26. fig. 5, tr.) very slender, sharp-pointed, measuring
about 0°16 mm. in length.

Register No. and Locality. III. 12, Wooded Isle.

4, LeUCETTA CHAGOSENSIS Dendy.
Leucetta chayosensis Dendy {1913}.

There are two specimens in the collection. One (R.N.II.4), which measures
32 mm. in length, 16 mm. in height, and has an average thickness of 8 mm.,
is compressed and irregular, proliferating into short, nodular processes ; the
vents, which have slightly prominent margins, are scattered and vary in size
between 1 and 2mm. The second specimen (R.N. II. 5 6) consists of three
small lobose fragments which have been broken off a larger specimen.

The Abrolhos specimens agree closely with the type except that the
ectosome is thinner and the subdermal cavities slightly smaller. Numerous
embryos are present, the structure of which indicates clearly that the larva
of Leucetta is a parenchymula.

Previously known Distribution. Chagos Archipelago, Indian Ocean (Dendy).

Register Nos. and Locality. I. 4, LU. 56, Wooded Isle.

5. Leucerra microRAPHIS Haeckel.

Leucetta primigenta var. microraphis Haeckel { 1872).
Leuconia dura Poléjaeff (1883).

Leucetta primigenia var, microrrhaphis Ridley [1884].
Leucetta microrhaphis Lendenfeld [1885 |.

Leucandra microrhaphis Dendy [1892].

Leucetta microraphis Dendy & Row {1913}.

The single specimen consists of a small, oblong fragment measuring about
22 by 10 mm. ; at one end there is a small slit-like vent measuring about
2by 1mm. Colour in spirit light brown ; texture very hard and coarse.

The skeleton consists of triradiate spicules only; these are of two sizes and
scattered irregularly throughout the sponge except at the surface, where

SPONGES FROM THE ABROLHOS ISLANDS. 483

they are arranged tangentially. The rays of the large triradiates measure up
to 1-1 by 0:23 mm., those of the smaller 0-18 by 0°013 mm.

Previously known Distribution. Bermudas (Poléjaef’) ; Australia (Poléjaef,
Ridley, Lendenfeld, Dendy).

Register No. and Locality. V1. 17 6, Sandy Isle.

6. LeucaLtis cLaTaria Haeckel.
(For literature and synonymy vide Dendy and Row [1913].)
There are two very fine specimens of characteristic external form, each
measuring about 80 mm. in diameter.
Previously known Distribution. See Dendy [1913].
Register No. and Locality. IIL. 3, 1V. 2, Wooded Isle.

7. SYCON GELATINOSUM de Blainville.

(Por literature and synonymy vide Dendy and Row [1913].)

There are six specimens of this common Australian species in the collection,
usually forming colonies of several or many persons, but varying considerably
in the size of person and in the way in which the colony branches; the
largest person measures 25 mm. in height and 12 mm. in greatest diameter.

The colour in spirit varies from light yellowish brown to dull greyish
‘brown. ‘The oscula are all provided with a fringe of spicules.

There is one specimen (R.N. VII. 1d) consisting of a single person only ;
it is 16 mm. in height and has an average diameter of 7 mm. ; the colour in
spirit is almost white.

The internal structure and spiculation of all the specimens are typical.

Previously known Distribution. Australia (various authors and collections) ;
Java (Haeckel).

Register Nos. and Localities. I. 2, Il. 5a, III. 2, Ii. 134, III. 136,
Wooded Isle ; VII. 1d, Sandy Isle. . '

8. VOSMAEROPSIS MACKINNONI n. sp. (PI. 25. fig. 4; Pl. 26. fig. 6.)

Sponge (Pl. 25. fig. 4) colonial, consisting of a number of short, thick,
subeylindrical individuals uuited together in an irregular manner along a
greater or lesser portion of their length. Hach individual has a circular or
oval osculum at its summit; in perfect individuals a beautiful oseular fringe
is present, formed of very long, hair-like oxea; in others this fringe is broken
off short. The individuals vary rather in size owing to their peculiar
branching and colonial habit ; an average full-grown person is 15 mm. in
height and 3 mm. in diameter, the thickness of the wall being 13mm. The
outer surface is rough and uneven, and large oxea can be seen projecting
irregularly from it; these oxea are more or less absent from the basal portions
of the sponge, where the surface is much more even and only slightly
roughened. Colour in spirit pale brown ; texture rather fragile.

DR *

00”

484. PROF. A. DENDY AND MISS L. M. FREDERICK ON

Both dermal and gastral cortices are well developed, the former about 0°18
and the latter about 0'l mm. thick. Between these lies the chamber layer
with a thickness of about 1 mm.

The canal system is ‘“‘sylleibid,” the elongated flagellate chambers, up to
0-4 mm. in length, opening into wide exhalant canals, which interdigitate
with the main inhalant canals running in from beneath the dermal cortex.
Small, scattered dermal pores lead into much narrower inhalant canals, which
pierce the dermal cortex to open into the wide outer ends of these main inhalant
canals.

The collared cells are apicinucleate.

The skeleton of the dermal cortex consists of tangentially placed triradiates
of various sizes, beneath which lie the short rays of the subdermal pseudo-
sagittal triradiates.

The skeleton of the gastral cortex consists exclusively of rather slender
triradiates arranged tangentially, which become strongly sagittal (alate)
towards the osculum, with the oral arms extended parallel with the margin of
the latter.

The skeleton of the chamber layer consists of (1) the contripetally directed
rays of the large subdermal pseudosagittal triradiates, (2) very large and
stout subgastral sagittal triradiates, (3) similar triradiates whose paired rays
lie at a variable distance beneath the gastral cortex, (4) the inner portions of
the large oxea, whose outer portions project through the dermal cortex.

Spicules:—(1) Triradiates of the dermal cortex (PI. 26. fig. 6d) ; approxi-
mately regular, with conical, gradually tapering, sharply-pointed rays,
measuring from about 0°23 by 0:026 to 0-4 by 0:04 mm.

(2) Triradiates of the gastral cortex (PI. 26. fig. 6¢); approximately
regular and more or less strongly sagittal; much smaller on the average than
those of the dermal cortex, with slender, gradually tapering, sharply-pointed
rays measuring about 0°21 by 0°013 mm.

(3) Subdermal pseudosagittal triradiates (Pl. 26. fig. 6c); the three rays
are all different ; the true basal ray, now forming a false pair with one of the
orals, is the shortest of the three and straight or nearly so, conical and
gradually sharp-pointed, measuring, say, 0°21 by 0:°026 mm. ; the oral ray
which forms an apparent pair with the basal is rather longer, measuring, say,
0:26 by 0:026 mm., and more or less crooked; the other oral ray, now
centripetally directed, is much longer and more slender, measuring, say, 0-5
by 0°02 mm., perfectly straight and gradually sharp-pointed.

(4) Subgastral sagittal triradiates (Pl. 26. fig. 6b); very large and stout.
rays conical, gradually and sharply pointed, oral rays often slightly curved or
crooked, basal ray (centrifugally directed) longer than orals; oral rays
measuring, say, 0°41 by 0:04 mm., with basal 0°52 by 0:05 mm., but variable.

(5) More distal triradiates of the chamber layer (PI. 26. fig. 6); not
sharply distinguishable from (4), but usually with shorter basal rays and
straighter orals,

SPONGES FROM THE ABROLHOS ISLANDS. 485

(6) Large oxea (PI. 26, fig. 6a), echinating the outer surface of the
sponge ; stout, curved, often crooked, especially the outer part, the inner
portion being straighter and tapering more gradually ; fairly sharply pointed
at each end; measuring up to 1:2 by 0:06 mm.

(7) Oxea of the peristomial fringe; very long and slender, hair-like,
usually broken off ; measuring up to 2°3 by 0-009 mm.; outer ends hastate,
very sharply pointed.

It is quite possible that this species may prove to be identical with one or
other of two species in the Hamburg South-west Australian collection for
which the late Mr. Row proposed the names Vosmaeropsis dendyi and
V. primitiva. These names are given in Dendy and Row’s “ Classification and
Phylogeny of the Calcareous Sponges” [1913], but no descriptions have yet
been published. Beyond the fact that the species in question belong to the
same section of the genus as V’. mackinnoni, it is impossible at the present time
to say anything about them, as the specimens have been sent to Japan for
further investigation by Dr. Sanji Hozawa.

Register No. and Locality. IV. 1, Wooded Isle.

9. GRANTIOPSIS CYLINDRICA Dendy. (PI. 25. figs. 5, 6, 7,5; Pl. 26. fig. 7.)
Hypograntia infrequens Carter [1885-6].
Grantiopsis cylindrica Dendy [1892, 1893 Aj.
Grantiopsis cylindrica Dendy & Row {1913}.
Grantiopsis infrequens Dendy & Row [1915].

There are five specimens in the collection, which vary slightly in size and
appearance ; each consists of a single person only.

R.N. I. 1 (Pl. 25. fig. 8) is typical ; it is attached toa rock by its base, and
forms a slightly curved, cylindrical tube, 20 mm. long and 5 mm. in diameter,
with a slightly constricted, terminal osculum, almost without fringe ; the wall
of the tube is 1 mm. in thickness. The surface is almost smooth, with a
characteristic glistening appearance due to the presence of the large
tangential triradiates. Colour in spirit almost white; texture firm and
compact, but brittle. Other typical specimens are shown in figs. 5-7.

The skeleton arrangement, the form and size of the spicules, and the canal
system agree closely with those of the type of the species as described by
Dendy [1892], and a detailed description is therefore not necessary. As,
however, the spicules have not as yet been figured separately, but only in the
transverse section given by Dendy [1893 A], we take this opportunity of
supplementing his description with the necessary illustrations (Pl. 26.
fig. 7, a-h; wide also Description of Plates).

In their “Classification and Phylogeny of the Calcareous Sponges” [1913],
Dendy and Row have pointed out that Carter’s Hypograntia infrequens is
undoubtedly a species of Grantiopsis, but, according to Mr. Row’s observations
on material collected by the Hamburg South-west Australian Expedition,

be
R

PROF. A. DENDY AND MISS L. M. FREDERICK ON

different specifically from G. cylindrica. We are now in a position, however,
as the result of microscopical examination of the very small, fragmentary type
in the British Museum, to state that Hypograntia infrequens and Girantiopsis
cylindrica are specifically identical. This being the case, it might be argued
that both the generic and the specific names proposed by Carter should take
priority.

As to the generic name Hypograntia, it must be pointed out, in the first place,
that the diagnosis of this genus contains nothing that is distinctive, and is
founded upon an obvious error with regard to the structure of the canal-
system, and, in the second place, that although Hypograntia infrequens is the
first of six species described in the same paper, it is said to be “‘incertw sedis,”
so that it cannot reasonably be regarded as the type of the genus, and as it is
quite distinct generically from the five remaining species, it seems advisable
to retain the generic name G'rantiopsis (originally regarded as a subgenus of
Grantia).

As to the specific name infrequens, it must be pointed out that Carter’s
description of the species is quite unrecognisable. He altogether omits the
most characteristic feature of the sponge, viz. the reduced tubar triradiates,
and deseribes the subgastral quadriradiates as triradiates. It was only the
fact that we happened to have access to Mr. Carter’s manuscript drawings that
put us on the track of the identification. Even in these drawings the reduced
tubar triradiates are not shown. Had the identification becn established
before the publication of the name Grantiopsis cylindrica, it would doubtiess
have been better to call the species Grantiopsis infrequens, but as the former
name has now become established in the literature and as Mr. Uarter’s species
was not recognisably described, we see no sufficient reason for reverting to
his name.

Previously known Distribution. Port Phillip Heads (Dendy, Carter).

Register No. and Locality. I. 1, VI. 17¢, VII. la, VII. 16, VIE. Le,
Sandy Isle.

9a. GRANTIOPSIS CYLINDRICA Dendy var. FRUTICOSA nov. (Pl. 25. fig. 9.)

The specimen, which is colonial, branched and bushy, consists of about 14
persons united together at their bases by short connections ; each person
forms a slightly curved cylindrical tube with a circular osculum at the
summit without projecting fringe; the persons average only 12 mm. in
length, with a nearly uniform diameter of 3°5 mm.; they have an almost
smooth surface, with a characteristic glistening appearance due to the
presence of the large tangential triradiates. Colour in spirit very pale
coffee-brown ; texture compact and firm. The skeleton arrangement, canal
system and spiculation agree so closely with those of the typical form that
no detailed description is necessary.

The variety differs from the type of the species chiefly in its bushy habit
and the smaller size of the individual persons, but also in that the dermal

SPONGES FROM THE ABROLHOS ISLANDS. 487

cortex forms only about one-third of the total thickness of the tube as com-
pared with about half in the typical form, while the spicules generally are
slightly smaller and the rays of the large dermal triradiates somewhat
shorter and blunter.

Register No. and Locality. II. 4, Wooded Isle.

10. LELAPIA ANTIQUA n. sp. (PI. 25. fig. 10; Pl. 26. fig. 8.)

The single specimen, which is attached by its lower end to a piece of
calcareous débris, measures 20 mm. in height and 10 mm. in greatest
diameter. It is erect, club-shaped, somewhat curved upon itself and slightly
flattened in one plane. The single oval osculum, which measures 3 by 2 mm.,
is nearly terminal in position and has a well-developed, collar-like peristome.
The surface of the lower two-thirds of the spongs is smooth; that of the
upper third is coarsely and unevenly hispid, owing to large projecting
brushes of long, slender oxea. Colour in spirit light yellowish grey ;
texture firm.

The dermal cortex is about 0-42 mm. thick, the gastral cortex about
0-13 mm., and the chamber layer, between the two, about 1°95 mm.

There appear to be numerous small dermal pores scattered over the surface,
but now very difficult to recognize. Short and narrow canals lead from the
surface into large and very irregular crypts in and beneath the dermal
cortex, from which the rather wide inhalant canals run radially inwards.
The canal system is typically leuconoid. The flagellate chambers, thickly
scattered in the clear, gelatinous mesogloa, are oval and about 0°1 mm. in
longer diameter. ‘The main exhalant canals open radially into the large
central gastral cavity.

The collared cells are apicinucleate.

The skeleton of the dermal cortex consists of two very distinet parts:
(1) several layers of slender-rayed, normal sagittal triradiates, arranged
tangentially, the whole about 0:085 mm. thick; (2) a deeper and thicker
part, consisting of longitudinal bundles of huge oxea, the whole about
0:34 mm. thick. The dense brushes of slender oxea that project from the
surface may also be included with the skeleton of the dermal cortex.

The skeleton of the chamber layer consists of (1) a confused interlacement
of huge oxea, lying in all directions ; (2) radially or subradially arranged
fibres or bundles of tuning-fork spicules, with the unpaired rays directed
outwards ; (3) the basal rays of subgastral sagittal triradiates.

The skeleton of the gastral cortex consists of several layers of alate
triradiates lying tangentially but otherwise without definite arrangement,
backed by the oral rays of the subgastral sagittal triradiates.

The skeleton of the peristome is rather remarkable ; there is no fringe of
freely projecting spicules, but a thin peristomial membrane supported by a
dense but approximately single layer of huge oxea, which are really a
continuation of the deeper skeleton of the dermal cortex, lined by a layer of

488 PROF. A. DENDY AND MISS L. M. FREDERICK ON

alate quadriradiates with short apical rays, forming a continuation of the
gastral skeleton, but with the paired rays arranged more regularly, parallel
with the oscular margin.

Spicules :-—(1) Normal sagittal triradiates of the dermal cortex (PI. 26.
fig. 8c) ; with slender, gradually sharp-pointed rays ; the oral angle is very
wide and the oral rays measure about 0°23 by 0:014 mm. ; the straight basal
ray is longer than the orals and measures, say, 0°4 by 0:013 mm.

(2) Subgastral sagittal triradiates (Pl. 26. fig. 8c’) ; these are like the
normal sagittal triradiates except that the rays are stouter and the orals more
distinctly recurved ; the orals measure up to 0°3 by 0:026 mm., with the
basal 0°5 by 0:03 mm.

(3) Laterally extended (alate) triradiates of the gastral cortex (Pl. 26.
fig. 8d) ; the oral rays are straight or very slightly recurved, gradually
sharp-pointed, measuring up to 0°31 by 0-016 mm., and the oral angle is so
wide that the rays are almost in the same straight line ; the basal ray is very
short, straight and conical, measuring about 0°04 by 0:01 mm.

(4) Alate quadriradiates of the peristome (Pl. 26. fig. 8) ; these are
similar to the alate triradiates, with the addition of a very short, straight,
sharp-pointed apical ray.

(5) “Tuning-fork” spicules (PI. 26. fig. 8/); with straight, slender,
gradually sharp-pointed rays; the basal ray is the longest and stoutest,
measuring about 0:4 by 0-009 mm.; the two oral rays, which are slightly
unequal in length, run straight forward parallel to one another ; the longer
of the two measures about 0°21 by 0:006 mm.

(6) Large stout oxea (Pl. 26. fig. 8a); fusiform, slightly curved, some-
times a little irregular in shape and diameter, tapering gradually to a sharp
point at each end ; full-grown spicules measure up to 2:0 by 0:09 mm.

(7) Slender oxea of the dermal brushes (PI. 26, fig. 85) ; straight or very
slightly curved, gradually sharp-pointed, sometimes slightly hastate,
measuring up to 1°6 by 0:007 mm., but often less.

This species closely resembles Lelapia australis Carter, as described and
figured by Dendy [1893 B], but differs in the absence of microxea (‘ mortar
spicules”) and in the presence of a definite layer of large, longitudinally
placed oxea in the deeper part of the dermal cortex. In the latter respect it
occupies an intermediate position between Lelapia australis and Lelapia
nipponica, a remarkable Japanese species described by Hara [1894]. We
have been unable to obtain access to the description of this species, but,
thanks to the kindness of Dr. Hézawa, have been able to examine a number
of transverse sections *. Jt is obvious that Lelapia nipponica is a much more
primitive species than Z. australis. The bundles of “ tuning-fork ” spicules
are arranged almost strictly radially, with the unpaired rays centrifugally
directed, and clearly represent the articular tubar skeleton, though the canal

* Since this paper was read, Dr. Hozawa [1923] has published an illustrated description
of the Japanese species and proposed for it the new genus Paralelana.

SPONGES FROM THE ABROLHOS ISLANDS. 489

system is already of the leuconoid type. The large oxea are confined to the
dermal cortex, where they form a well-defined layer, and have not yet invaded
the chamber layer as in LZ. australis and, to a less extent, in L. antiqua.
There are other minor differences distinguishing L. nipponica from either of
the Australian species, but the material in our possession is not sufficient to
elucidate all these.

It is clear that Lelapia antiqua is intermediate in structure between the
more primitive JL. népponica and the less primitive L. australis, and it is
interesting to note that it also vecupies an intermediate position geographi-
cally, Z. australis having been found so far only off the Victorian coast.

Register No. and Locality. VI. 1e, Sandy Isle.

11. Dercrropsis minor Dendy [1916 B].

There is one specimen in the collection. The sponge, which measures
42 mm. in length, 25 mm. in breadth, and has an average thickness of 7 mm.,
forms an irregular, oval, cushion-like mass, concave below, and probably grew
on the back of a crab. The margins are broadly rounded. The surface is
smooth and minutely punctate. The vents are small and few in number,
scattered singly over the upper surface. Inhalant pores are closely scattered
over the surface. Colour of surface in spirit varying from light brown to
dark slate-grey, internally dull yellow ; texture firm and compact.

The skeleton consists of a dense feltwork of loose spicules iirregularly
arranged except at the surface, where there is a dermal layer of perpendicu-
larly arranged oxea. The spiculation agrees closely with that of the type.

The ectosome and choanosome are not sharply differentiated, but the
former contains many more spicules. There is no fibrous tissue in the ecto-
some. The mesogloea of the choanosome is finely and uniformly granular.
The dermal pores lead into short inhalant canals, which penetrate the ecto-
some and lead into large crypts from which the inhalant canals of the
choanosome originate. The canal system is diplodal, not eurypylous as
described for Dercitopsis ceylonica Dendy. The flagellate chambers are large
and pouch-like, with a short wide aphodus and a short narrow prosodus ;
they measure about 0:04 mm. in lenger diameter. The exhalant and inhalant
canaliculi of the chambers are lined by flattened epithelium, the nuclei of
which can be seen. The collared cells are apicinucleate. Small round
testes are scattered in the deeper parts of the choanosome, in which
spermatogenesis is clearly shown. ‘Tailed spermatozoa are present. ‘There
is no trace of ova, so the species is probably dicecious.

Previously known Distribution. Indian Ocean (Dendy).

Register No. and Locality. Il. 1, Wooded Isle.

12. DERCITOPSIS MAMMILLARIS (Lendenfeld).
Plakinastrella mammillaris Lendenfeld [1906].
Dercitopsis mammillaris Dendy [1916 B}.
The single specimen is of depressed cushion-like form, and has apparently
been attached by a broad base to the vertical side of a rock, for two large

490 PROF. A. DENDY AND MISS L. M. FREDEKICK ON

oval vents, 3°5 mm. in diameter, with slightly prominent margins, oceur on
one edge. Microscopic inhalant pores are thickly scattered over the smooth
surface. Colour in spirit light greyish brown ; texture firm and compact.

The skeleton consists of a thick feltwork of loose, irregularly arranged
spicules. ‘There is a dense layer of oxea at the surface, largely but not
entirely radially arranged.

Spicules :—(1) Tetracts ; with smooth, sharp-pointed rays measuring from
0:093 to 0°2 mm. in length and from 0:018 to 0:04 mm. in thickness.

(2) Diacts (oxea) ; smooth, fusiform, slender, slightly curved, gradually
sharp-pointed, sometimes with a kink or an enlargement near the middle,
measuring from 0:06 to 0-14 mm. in length and from 0:004 to 0°005 mm. in
thickness. The oxea are more numerous than the tetracts. Triacts are absent.

The dermal pores lead into short, narrow, inhalant canals piercing the
densely spiculated ectosome, beneath which they unite to form sub-cortical
erypts from which the inhalant canals of the choanosome originate. The
canal system is diplodal, the aphodi and prosodi being longer and narrower
than in Dercitopsis minor Dendy. The flagellate chambers are sub-spherical,
measuring about 0°04 mm. in diameter. The narrow exhalant canals gradually
pass into wider ones, which in turn open into the wide oscular tubes. The
main inhalant canals are surrounded by collenchymatous mesogloa devoid of
spicules ; the main exhalant canais are probably like the inhalant, but as no
sections have been taken through a vent this cannot be proved. The meso-
elaea between the chambers is finely granular.

Testes are irregularly scattered in the choanosome ; they are rounded sacs
lined by an epithelium and containing various stages in the development of
spermatozoa. Segmenting embryosarealso present ; the sponge is therefore
hermaphrodite and presumably protogynous.

Previously known Distribution. West coast of Australia (Lendenjeld).

Register No. and Locality. II. 13, Wooded Isle.

13. STELLETTA BREVIS Hentschel [1909].

The single specimen, a cushion-like mass which has probably been torn off
a rock, measures 38 mm. in length, 25 mm. in breadth, and 15 mm. in
thickness. Surface smooth but finely granular. Oscula not visible. The.
surface colouring in spirit varies from a very light brown to a violet-grey,
that of the interior is light brown.

The sponge possesses a well-developed cortex. Immediately beneath the
surface is a thin layer of strongylasters, then comes a non-fibrous portion
filled with minute brown pigment granules and containing inhalant chones ;
below this is a light-coloured, non-pigmented, fibrous layer. Between the
cortex and the choanosome are large sub-cortical crypts. The choanosome,
in which oxyasters are thickly scattered, contains narrow inhalant
canals.

SPONGES FROM THE ABROLHOS ISLANDS. AQT

The skeleton, which agrees closely with that of Hentschel’s South-West
Australian specimen, consists of radially arranged bundles of plagiotrizenes
and oxea ; in the deeper parts there is lessregularity. The spicules resemble
closely those of the type in size and appearance.

Previously known Distribution. Sharks Bay, S.W. Australia (Hentschel).

Register No. and Locality. V1. 8, Sandy Isle.

14. SreLLerra DEBILIS Thiele.
Stelletta debihs Vhiele { 1900).
Stelletta debilis Lendenfeld [1903].

The single complete specimen is sub-spherical in shape, measuring 15 mm.
in diameter, There is a single oval osculum, 2 mm. long and 1 mm. broad,
near the point of attachment of the sponge to a piece of rock. Inhalant
pores are thickly seattered all over the slightly roughened surface between
the cladi of the triznes. Colour in spirit light brown; texture firm and
compact, but compressible.

There is a slightly fibrous cortex, and the skeleton is radially arranged and
quite typical.

Spicules :—(1) Orthotriznes ; shaft straight or nearly straiglit, tapering
gradually to a sharp point, measuring 1°35 by 0:026 mm.; cladi sharply
pointed, recurving at once so as to extend at right angles to the shaft,
measuring 0°16 mm. in length.

(2) Anatriznes ; these frequently project beyond the surface of the
sponge ; shaft long, slender, slightly curved, measuring 1°3 by 0-015 mm. ;
cladi well developed and markedly recurved, measuring 0:052 mm. in length.

(3) Oxea; fusiform, usually curved, sharp-pointed, measuring 1-1 by
0:02 mm.; these are most abundant in the deeper parts of the sponge.

(4) Oxyasters ; with 6 to 10 slender, sharp-pointed rays and no distinct
centrum; diameter 0°017 mm.; these are scattered sparsely throughout
choanosome and ectosome.

Previously known Distribution. Ternate (Thiele).

Register No. and Locality. 11. 16, Wooded Isle.

15. STELLETTA SIGMATRIZNA Lendenfeld [1906].

There are two small, complete specimens, sub-spherical in shape, 12 and
8 mm. in diameter respectively. Inhalant pores, visible with a lens, are
scattered evenly over the smooth surface. Hach specimen has one minute
oval osculum lying in a slight depression. Colour in spirit buff.

The skeleton consists of radially arranged oxea and trizenes, not concen-
trated into bundles. Just beneath the surface the cladi of the trieenes form
a sharply-defined layer; their shafts pierce the region of the chones, which
is cladome-free. In the choanosome are radially placed anatriznes,
orthotrienes in various stages of development, and less regularly arranged
oxea.

492 PROF. A. DENDY AND MISS L. M. FREDERICK ON

Spicules :—(1) Orthotrizenes ; shaft straight, gradually pointed, measur-
ing about 1:1 by 0:015 mm.; the cladi, which are about 0°3 mm. long, at
first form with the shaft an angle of about 120°, but somewhere along their
length they abruptly turn back so that their distal ends are at right angles to
the shaft.

(2) Anatrizenes ; shaft straight, gradually tapering to a sharp point,
measuring about 1:1 by 0°03 mm.; the sharply-pointed cladi, which are
about 0°15 mm. long, curve outwards almost at right angles to the shaft and
then run back parallel to it.

Our specimen contains only a few of the irregular orthotrizenes and of the
sigma-like anatriznes figured by Lendenfeld in his ‘ Valdivia’ report.

(3) Oxea ; straight or slightly curved, bluntly pointed, measuring about
11 by 0:015 mm.

(4) Oxyasters ; of two kinds—(a) large, with 2 to 6 fairly stout, roughened,
bluntly pointed rays, total diameter about 0:02 mm.; these are most abun-
dant just beneath the inhalant chones: (6) small, with 6 to 10 thin, very
minutely roughened rays; total diameter 0:014 mm. These are scatiered
through the choanosome, but not abundantly.

Previously known Distribution. Dirk Hartog, W. Australia (Lendenfeld).

Register Nos. and Localities. LV. 8, Wooded Isle ; VI. 18c, Sandy Isle.

16. ANCORINA AUSTRALIENSIS (Carter). (Pl. 25. fig. 11.)
Stelletta australiensis Carter | 1883].
Ecionema australiense Sollas [1888].
Ancorina australiensis Lendenfeld [1903].

There are four specimens in the collection. The largest (R.N. VI. 6a;
Pl. 25. fig. 11) forms an incomplete cone, 40 mm. in height, with a base
measuring 60 by 70 mm. ; the apex is truncated, and over the flattened area
thus formed, which measures 40 by 25 mm., are scattered numerous open
vents which vary in diameter from 0°25 to 1 mm. Part of the side of the
cone is encrusted by Spongelia dakinin.sp. The second specimen (R.N. I. 3)
is irregularly massive and measures 80 by 60 by 50 mm.; its upper surface
is almost entirely covered by an encrusting Aentera and a mass of other
débris. The third specimen (R.N. IL. 6) measures 40 by 45 by 25 mm., and
is largely overgrown by amass of other organisms. The fourth (R.N. III. 6)
an irregular, cake-shaped mass, measures 85 by 50 by 40. mm.; its upper
surface is uneven and slightly wrinkled.

No vents are visible except in the figured specimen; inhalant pores,
visible with a pocket lens, are scattered in groups of two or three over the
smooth upper surface of all the specimens; they are most marked in the
figured specimen. Colour in spirit purplish brown on outside, light yellowish
brown inside ; texture firm and compact.

The skeleton consists of radially arranged trieenes and oxea, with a layer
of small ectosomal oxea vertical to the surface. The microrhabds are

SPONGES FROM THE ABROLHOS ISLANDS. 493

thickly scattered through the ectosome and choanosome, as well as forming
a well-developed dermal layer. Large brown pigment granules are
scattered throughout all four specimens.

The form and size of the spicules agree closely with Sollas’s description
of Carter’s specimen.

Previously known Distribution. Fremantle, West Australia (Carter).

Register Nos. and Localities. VI..6 a, Sandy Isle; I. 3, Turtle Bay;
Il. 6, III. 6, Wooded Isle.

17. ANCORINA BREVIDENS n. sp. (PI. 26. fig. 9.)

The single specimen, which is about 60 mm. long, 40 mm. wide, and
20 mm. thick, is massive and potato-shaped ; at one end is a group of
about twenty small, close-set oscula, which average about 1 mm, in diameter.
The surface is smooth, a large portion of it being covered with foreign matter
of varying kind. Colour in spirit purplish grey; texture hard, compact,
slightly compressible.

The ectosome forms a cortex about 0:5 mm. thick, with numerous
cortical crypts in its deeper portion; the cortex appears to be only slightly
fibrous, but contains numerous brown granular pigment cells, which occur also
in the outer portion of the choanosome.

The main skeleton consists of close-packed, radial bundles of oxea and
tricenes ; the cladi of the trizenes occur at various levels in and just below
the cortex, but chiefly just beneath the surface. The outer part of the cortex
contains numerous comparatively small, slender, radially arranged oxea.
There is a dense dermal layer of microrhabds, and both microrhabds and
tylasters are abundantly scattered through ectosome and choanosome. __

Spicules :—(1) Orthotrienes (Pl. 26. fig. 9 a) ; shaft long, straight or
slightly curved, tapering gradually to a fine point (sometimes blunted),
measuring about 2°2 by 0°05 mm.; cladi gradually but bluntly pointed,
measuring about 0°16 by 0:04 mm.

(2) Anatrizenes (PI. 26, fig. 9 6) ; shaft very long, slender, tapering very
gradually to a finely pointed or blunted extremity, measuring about 3:0 by
0:02 mm.; cladi very much reduced in length, bluntly pointed, measuring
about 0°026 by 0°017 mm.

(3) Oxea of two sizes; the larger (Pl. 26. fig. 9 ¢) fusiform, curved,
gradually and sharply pointed, measuring about 2°3 by 0:04 mm. ; the
smaller (Pl. 26. fig. 9 c’) found only in the cortex, straight, sharp-pointed,
measuring about 0:34 by 0:009 mm.

(4) Microrhabds (PI. 26. fig. 9 d) ; roughened, slightly swollen in centre,
measuring about 0:011 by 0:003 mm.

(5) Tylasters (PI. 26. fig. 9 e) ; some have 6 to 8 slender rays with very
small heads and no centrum ; others, which are slightly smaller, have more
rays and a small centrum ; the average diameter of the entire spicule is

0:013 mm,

494 PROF. A. DENDY AND MISS L. M. FREDERICK ON

This species is evidently very closely related to Stelletta truncata Kieschnick
[1898] = Ancorina amboinensis Lendenfeld [1902], from Amboina, in the
Malay Archipelago, differing in the great abbreviation of the cladi of the
trizenes, especially of the anatrieenes.

Register No. and Locality. VI. 16 6, Long Island.

18. AurorA Row! Dendy [1916 B].

There are three lobose pieces which have probably been attached to rock ;
the largest measures 40 mm. in length, 20 mm. in breadth and 15 mm. in
thickness ; it hasa bit of branching coral growing throughit. The surface is
sub-glabrous and crumpled, with irregular grooves of varying depth running
across it. The oseula, which are few in number and single, lie at the apices
of small rounded prominences ; they vary in size, the largest being 2 mm. in
diameter. Inhalant pores are scattered singly over the surface. Colour in
spirit chocolate-brown ; texture compressible but fairly compact.

The skeleton arrangement and spiculation agree with those of the type
specimen, the only difference being the occasional occurrence of irregular
branching of the cladi of the orthotrizenes ; this is not sufficient, however,
to justify a specific distinction.

Previously known Distribution. Seychelles (Dendy).

Register No. and Locality. 1. 8, Wooded Isle.

19. ASTEROPUS SIMPLEX [ Carter ].

(For literature and synonymy vide Dendy [1916 B}.)

The single specimen is an irregular, cake-shaped mass, about 80 mm. in
maximum diameter. The upper surface is uneven, somewhat ridged and
depressed at intervals; a number of open vents, 1 mm. in diameter, are
seattered over it. Colour in spirit purplish brown ; texture firm, compact,
coarse.

The skeleton arrangement and spiculation agree with those of the ‘ Sealark ’
specimen, and need not be described.

Previously known Distribution. Fremantle and Port Phillip Heads,
Australia; Hayti (Carter); S.W. Australia (Hentschel) ; Indian Ocean
(Dendy)..

Register No. and Locality. V1.5, Sandy Isle.

20. Hryius proximus Dendy [1916 B].

The single specimen, an irregular lobate mass, is 28 mm. in length, 22 mm.
in breadth and 15 mm. in thickness. The surface is uneven and punctate, in
parts covered with caleareous débris. Scattered singly over an irregular
area, near the end where the sponge was probably attached, are many minute,
widely open, dermal pores, while over the major portion of the rest of the
sponge are numerous white specks, giving the surface its punctate appearance;
these are probably closed inhalant pores, Ona prominent part of the upper

SPONGES FROM THE ABROLHOS ISLANDS. 495

surface is a single, round, open vent, 1:5 mm. in diameter. Colour in spirit
light greyish brown ; texture hard and compact.

The arrangement, form and size of the spicules are similar to those of the
type specimen.

Previously known Distribution. Cargados, Indian Ocean (Dendy).

Register No. and Locality. V1. 12a, Sandy Isle.

21. Donatra Ropusta (Bowerbank).
Tethea robusta Bowerbank [1873].
Tethya globostellata Lendenfeld [1897].
Tethya clifton’ Ridley [1884].

Tethya ingalli Kirkpatrick [1900].
Donatia arabica Topsent [1906}.
Donatia japonica vax. albanensis Hentschel {1909}.
Tethya lyncurium Row [1911].
Donatia ingalli Dendy [1916 B).
Donatia globostellata Topsent {1918}.
(See M. Burton [1925] MS.)

There are three specimens in the collection, greyish white in colour,
ranging from 20 to 26 mm. in diameter. The surface of each is only slightly
tessellated.

The skeleton consists of radially arranged bundles of typical strongyloxea
which spread out beneath the surface in brushes; loose, radially arranged
mevascleres lie between the bundles.

Spicules :—(1) Strongyloxea ; measuring about 1°62 by 0:028 mm.

(2) Spherasters; these form a dense cortical layer; they have a large
centrum and short conical rays ; diameter about 0°1 mm.

(3) Small asters ; with short rays which may either bear a crown -of
short spines or be beset with numerous small spines ; the total diameter is from
0-012 to 0°016 mm. These are found in the cortex and the choanosome.

(4) Small asters ; with long rays which may either end in a sharp point
or be spined terminally. These are confined to the choanosome; but there
are many intermediate forms scattered through the sponge.

This species is undoubtedly very closely related to Donatia lyncurium

(Linneeus), from which it differs in the large size of the spherasters and the
ereater differentiation of the small asters.

Previously known Distribution. Red Sea (Row, Topsent) ; Indian Ocean
(Dendy, Lendenfeld, Ridley, Kirkpatrick) ; Australia (Hentschel).

Register Nos. and Localities. 11. 11, Wooded Isle; VI. 7, VI. 19, Sandy
Isle.

22. DoNATIA MULTISTELLA (Lendenfeld).

Tethya multistella Lendenfeld [1888].
Tethya multistella Hallman [1914).

This species is represented by one small, sub-spherical specimen, greyish
white in colour, 12 mm, in diameter ; and a fragment of another, pale brown

496 PROF. A. DENDY AND MISS L. M. FREDERICK ON

mm colour and apparently about 30 mm. in diameter. The surface of the
former is only slightly tessellated, while that of the latter, as far as can be
judged, is fairly smooth but uneven. ‘The cortex of both is lacunar and not
densely charged with spicules.

The skeleton consists of radially arranged bundles of strongyloxea, which
penetrate the cortex and spread out in brushes beneath the surface ; loose
radially arranged megascleres are scattered between the bundles.

Spicules :—(1) Strongyloxea of the usual type, measuring about 1:26 by
0-016 mm.

(2) Spherasters ; about 0:06 mm. in diameter, with conical, sharp-pointed
rays, nearly as long as the diameter of the centrum.

(3) Smallasters of cortex and choanosome ; these range from tylasters, with
the ends of the rays beset with small spines, to oxyasters. The diameter of the
small aster varies from 0:01 to 0:012 mm.

This species is undoubtedly closely related to Donatia robusta (Bowerbank),
but differs from it in the size of the spherasters and in small differences
in the small asters.

Previously known Distribution, Port Jackson, N.S.W (Lendenfeld, Hall-
mann) ; Port Phillip Heads (Hentschel).

Register Nos. and Locality. VI. 17a, VII. 17, Sandy Isle.

23. CHONDRILLA AUSTRALIENSIS Carter.
- (For literature and synonymy vide Dendy [1916 B].)

There are two small specimens in the collection ; one is buff-coloured in
spirit, the other liver-brown. The external form and spiculation agree closely
with those of the type.

Previously known Distribution. Port Jackson, H. Coast of Australia
(Carter, Lendenfeld) ; Sharks Bay, S.W. Australia (Hentschel) ; coast of
Cochin China (Lindgren) ; Okhamandal, Ceylon, Indian Ocean (Dendy).

Register Nos. and Localities. VII. 4a, Turtle Bay ; VIII. 1, Sandy Isle.

24, ReNIERA AQuzDucTUS O. Schmidt. (Pl. 25. fig. 12.)

Reniera aqueductus Schmidt (1862).
Renera aqueductus Kolliker [1864].
Renera aqueductus Czerniaysky {1878}.
Reniera aqueductus Swartschewsky [1905}.
Remera aqueductus Ostroumoy [1893].
Reniera aqueductus Babié [1921, 1922).

The single specimen (Pl. 25. fig. 12), which measures 55 mm. in height,
is erect, tubular and thin-walled ; at the upper end it divides into two short
tubes, each of which terminates in a large round osculum 3 mm. in diameter.
Inhalant apertures, visible with a pocket lens, are scattered over the smooth
surface. Colour in spirit greyish brown ; texture delicate, spongy, fragile.

SPONGES FROM THE ABROLHOS ISLANDS. 497

The skeleton consists of a somewhat rectangular, unispicular reticulation ;
towards the outer surface the spicules become concentrated into plurispicular
fibres running vertically to the surface.

Spicules :—Oxea ; small, smooth, straight or slightly curved, sharply
pointed, measuring about 0:23 by 0:008 mm.

Previously known Distribution. Adriatic (Schnudt, Babid); Black Sea
(Czerniavsky, Swartschewsky).

Register No. and Locality. VII. 3, Abrolhos Islands.

25. RENIERA ORIBRICUTIS Dendy [1921 B].

There are two specimens in the collection, one (R.N. VII. 14), the colour
of which is almost white in spirit, agrees very closely in all respects with the
type-specimen, except that it is smaller, measuring 25 mm. in length and
10 mm. in diameter, and has only a single vent, 5 mm. in diameter.

The second specimen (R.N. VIII. 2) consists of two fragments, lobate in
appearance, each possessing one large, round osculum, about 4 mm. in dia-
meter, with a slightly raised margin, opening out of a wide oscular tube.
This specimen, which is light brownish yellow in colour, does not conform
so closely to the type, the subdermal cavities being smaller and the primary
lines of the skeleton less differentiated. We feel justified, however, in
making the identification.

The spicules of the Abrolhos specimens are of much the same size as those
of the type, measuring about 0:14 by 0-006 mm.

Previously known Distribution. Amirante, Indian Ocean (Dendy).

Remster No. and Locality. VII. 1g, VIII. 2, Sandy Isle.

26. RENIERA PERMOLLIS (Bowerbank).
Tsodictya permollis Bowerbank [1866, 1874, 1882).
Reniera permollis Topsent | 18883].

Reniera permollis Nendy [1916 A}.

We identify with this species seven specimens of an encrusting nature
which vary considerably in size and general appearance. The surface is
slightly hispid, and the large subdermal cavities can be seen throueh the thin
dermal membrane, which is almost aspiculous. The oscula are small, single
and scattered, measuring up to about 3 mm. in- diameter. The colour in
spirit varies from light yellowish brown to nut-brown ; texture in most cases
very soft and friable, that of R.N. VI. 17d and R.N. VI. 17e firm and
compact.

The skeleton arrangement and spiculation agree closely with those of
specimens from Okhamandal, but the tendency to form primary lines is not
so strongly marked.

Previously known Distribution. Huropean seas (Bowerbank, T opsent) ;
Okhamandal, Indian Ocean (Dendy).

LINN. JOURN.—ZOOLOGY, VOL, XXXV, 36

498 PROF. A, DENDY AND MISS L. M. FREDERICK ON

Register Nos., Localities, etc. VI. 17d, VI. 17e, Sandy Isle; Il. 73,
III. 11, IV. 6, 1V. 7, IV. 10, Wooded Isle.

RENIERA spp.
There are also in the collection a number of more or less fragmentary
specimens representing other species of this difficult genus.
Register Nos., Localities, ete. 1. 3, VII. 46, Turtle Bay ; VI. 14, VI. 15,
Sandy Isle.

27. Perrosra puRA (Nardo) Vosmaer [1887].
Reniera dura Nardo.
Reniera? dura Nardo, Schmidt [1862].
Schmidtia dura Balsamo Crivelli [1863].
Schmidtia dura Schmidt {1868}.
Petrosia dura Topsent [1897].
Petrosia dura Ferray [1914].
Petrosia dura Babi¢ [1921, 1922).

The single specimen in the collection consists of a somewhat irregularly
shaped, digitiform fragment, 40 mm. long and 10 mm. thick, which had
probably been growing erect and has evidently been cut off transversely from
a larger sponge. The surface is smooth, and the dermal membrane, which
is thin but distinct, is pierced by inhalant pores visible with a lens. ‘There
are two round oscula, 2 mm. in diameter. Colour in spirit light reddish
brown ; texture hard but brittle.

The skeleton arrangement and spiculation agree closely with those of one
of Schmidt’s specimens in the British Museum.

The spicules are oxea, which show a great tendency to become rounded oft
at one or both ends ; they measure about 0:35 by 0-015 mm.

Previously known Distribution. Adriatic (Schmidt, Babié); Mediterranean
(Balsamo Crivelli, Topsent) ; Cantabrico (ferrar).

Register No. and Locality. VI. 16a, Long Island.

28. HALICHONDRIA PHAKELLIOIDES n. sp. (PI. 26. fig: 10.)

The single specimen consists of a portion of an erect lamella, 190 mm. in —
height, 75 mm. in width, with a maximum thickness of 8 mm,, constricted
towards the base of attachment so as to form a very short, thick stalk. The
lamella, which has a narrow, very slightly sinuous margin, appears similar
on both surfaces, but one has been somewhat cracked and rubbed.

The surface is smooth and practically even, covered everywhere by a very
thin, transparent dermal membrane (rubbed off in places), which contains
numerous irregularly and loosely scattered tangential oxea. The dermal
membrane is dotted with numerous small, round pore-areas, the pores
themselves apparently being closed. The oscula are very inconspicuous ;
they may be represented by minute apertures at the apices of several small

SPONGES FROM THE ABROLHOS ISLANDS. 499

prominences which are sparsely scattered over both surfaces of the sponge.
The colour in spirit is light brownish yellow; texture tough, flexible,
resilient.

The skeleton consists of loose tracts of spicular fibre running longitudinally
and confined to the middle of the lamella; at right angles to these on
both sides still looser and less well-defined fibres run outwards towards the
surface, where they terminate in loose brushes of spicules, which help to
support the dermal membrane.

Numerous loose spicules lie scattered everywhere between the fibres.
There is a fair quantity of spongin present, but so pale in colour that it is
almost invisible in unstained preparations.

Spicules: —Oxea (PI. 26. fig. 10 a) ; usually slightly curved or angulated
and tapering somewhat abruptly at each end to a sharp point; measuring
about 0°58 by 9:017 mm. when fully grown, but frequently shorter and
especially more slender. ‘These spicules occasionally become stylote, with
one end broadly rounded off (PI. 26. fig. 10).

FHalichondria phakellioides seems to come very near to Halichondria vela-
mentosa (Hansen) Lundbeck [1902], a remarkable fact considering that
the former is a shallow-water, sub-tropical form, while the latter is found in
deep water in the Arctic regions.

Register No. and Locality. V1. 1, Sandy Isle.

29, CHALINA PALMATA (Lamarck).
Spongia palmata Lamarck {1913}.
Chalina paimata Ridley & Dendy (1887 |.
Cladochalina euplax Lendenfeld [1887].
Chalina palmata Whitelegge [1901).

2 Ceravchalina retiarmata Dendy [1905].

There are three small, incomplete specimens in the collection. The sponge
is of erect habit, and consists of irregular, digitate processes arising from a
narrow base and sometimes branching. ‘The surface is fairly even, slightly
hispid, and has a distinctly reticulate appearance. The thin, transparent
dermal membrane is pierced by numerous round inhalant pores, which vary
considerably in size. The small, round oscula measure up to 2 mm. in dia-
meter, and are for the most part arranged uniserially on the margins. Colour
in spirit light brown : texture spongy, tough and fibrous.

The skeleton arrangement agrees closely with that of Chalina palmata
Ridley and Dendy [1887], except that the dermal reticulation of spicular
fibre has a somewhat smaller mesh.

Spicules :—Oxea ; short, slightly curved, graduaily sharp-pointed, measuring
about 0:07 by 0:004 mm., but often more slender.

After re-examining the type slides of Ceraochalina retiarmata Dendy
[1905], we think this species may also be identical with Chalina palmata

(Lamarck).
36*

500 PROF. A. DENDY AND MISS L. M. FREDERICK ON

Previously known Distribution. Indian and European Seas (Lamarck) ;
Torres Strait (Challenger) ; New South Wales (Lendenfeld, Whitelegge).
Register Nos. and Locality. II. 3, HU. 14, I. 17, Wooded Isle.

CHALINA sp.

The single specimen, which measures 40 mm. in height and has an average
diameter of 10 mm., is erect, shortly digitiform, slightly compressed, with a
single round osculum, 3 mm. in diameter, at the apex, to which two long
oscular tubes converge. Surface smooth, minutely punctate ; inhalant pores
close-set, about 0°04 mm. in diameter. Texture firm, compact, resilient. The
specimen is almost black on the outside and brown inside, owing to the
presence of numerous pigment granules.

The skeleton is very regular; a transverse section shows numerous parallel
primary fibres, about 0:06 mm. in diameter, containing a core of well-developed
spicules, usually arranged multiserially and surrounded by a thick coating of
spongin. The primary fibres are connected crosswise at very frequent
intervals by secondary fibres of less than one spicule’s length and usually
with only a single spicule in the axis. A few loose spicules are scattered
between the fibres. The spicules at the ends of the primary fibres pierce
the dermal membrane, which is also supported by a close-meshed network of
unispicular fibre.

Spicules :—Oxea; fusiform, slightly curved, gradually sharp-pointed,
measuring about 0°14 by 0:007 mm.

Register No. and Locality. III. 1, Wooded Isle.

PACHYCHALINA sp. ?
There is a small, delicate fragment, light brown in colour, which evidently
belongs to this genus.

Register No. and Locality. 11. 10, Wooded Isle.

30. CERAOCHALINA MULTIFORMIS Lendenfeld var. MANAARENSIS Dendy.
(For literature and synonymy vide Dendy |1905).)

We identify with this variety four pieces which have apparently formed
part of an erect, flattened, lamellar specimen, with an irregularly undulating
and proliferating surface. The largest piece is 83 mm. high and 55 mm.
wide, and the thickness of the lamella is about 6 mm. The surface is
glabrous, and the thin dermal membrane is pierced by numerous inhalant
pores. The oscula, which measure about 2 mm. in diameter, are fairly
numerous and scattered, but confined almost entirely to one surface of the
lamella, while some are marginal in arrangement. The colour in spirit is
dirty greyish violet ; texture firm, rather tough and resilient.

The skeleton arrangement and spiculation agree very closely with those of
the type of the variety, so no description is necessary.

SPONGES FROM THE ABROLHOS ISLANDS. 501

Previously known Listribution (of variety). Gulf of Manaar and Ceylon
(Dendy).
Register No. and Locality. I. 2, Turtle Bay.

31. PHL@ODICTYON ABROLHOSENSIS n. sp. (PI. 26. fig. 11.)

The single specimen, which is about 45 mm. long, 35 mm. wide and
12mm. thick, is somewhat flattened, cake-shaped, and has probably been
attached to a rock by its slightly concave under surface. The smooth,
convex upper surface gives off numerous very thin-walled fistule of varying
sizes, the largest being about 15 mm. in length. The diameter of the
fistulee is from 3 to 5 mm. ; some of the smaller ones end blindly ; these
either bear inhalant pores or are young stages of the larger ones, which
terminate each in a wide vent. The body generally is covered with a thin
rind (dermal membrane), which easily peels off. Colour in spirit greyish
brown; texture internally very soft and spongy, distinctly fibro-reticulate.

The skeleton of the interior consists of a loose, irregular, wide-meshed
network of multispicular fibre, averaging about 0:065 mm. in diameter.
The fibres are compact, the spicules being surrounded by a good quantity of
pale-coloured spongin. Numerous loose spicules, mostly arranged in small
bundles of one spicule’s length, are scattered through the soft ground
substance, which also contains many small brown pigment granules.
Immediately beneath the surface and parallel with it is an irregular reticu-
lation of similar spicular fibre, which tends to separate from the underlying
main skeleton and come away with the dermal membrane when the latter is
peeled off. The skeleton of the dermal membrane and that of the walls of
the fistula consists of a very dense feltwork of single spicules lying
tangentially in approximately a single layer. i

Spicules:—Oxea (PI. 26. fig. 11); slightly curved, sharply and fairly
gradually pointed at each end, sometimes hastate, measuring about 0°22 by
0-006 mm.

This species seems to approach Philwodictyon seychellense Dendy [1921 B]
most closely.

Register No. and Locality. 11. 9, Wooded Isle.

PHLGODICTYON sp.
There are two small fistulse, light brown in colour, which have evidently
been broken off from the surface of a Phlwodictyon.
Register No. and Locality. V1. 12 ¢, Sandy Isle.

39. PsBUDOESPERIA CARTERI n. sp. (PI. 26. fig. 12.)

Sponge lobose, elongated (probably horizontally), slightly compressed,
with corrugated surface. Vents and pores not seen. The dermal membrane
is distinet and thick, overlying the cavernous and coarsely fibrous interior.
Colour in spirit brown ; texture soft and compressible.

502 PROF. A. DENDY AND MISS L. M. FREDERICK ON

The main skeleton consists of a well-developed, coarse reticulation of
stout, compact, multispicular fibre up to about 0:2 mm. in diameter, composed
of closely packed megascleres with little or no spongin. The meshes of the
reticulation vary much in size, 0° mm. being an average diameter. There
is a loose dermal reticulation of ill-defined spicular fibre and loose spicules,
with more or less dense, radially arranged brushes of slender tylostyles.

Spicules:—(1) Tylostyli (Pl. 26. fig. 12a); often slightly curved or
crooked, with a well-developed oval head separated from the fairly stout
shaft by a slight constriction ; fairly gradually sharp-pointed at the apex ;
size about 0°28 by 0:008 mm. :

(2) Sigmata (Pl. 26. fig. 126) ; slender, simple and contort, with short,
abruptly recurved, sharply-pointed ends. Length about 0:04 mm. from
bend to bend.

(3) Large quadridentate anisochelee (Pl. 26. fig. 12 c-c’’) ; very numerous,
mostly in rosettes, but scattered singly as well. The free end is composed
of four short, sharply-pointed teeth which extend more or less at right angles
to the shaft, which is markedly curved and fairly stout; the fixed end is
quadrilateral in side view, with the three teeth almost parallel to one another ;
the median tooth connected with the shaft by a long, narrow falx. The total
length of the spicule is about 0°04 mm. and the shaft is about 0°004 mm. in
diameter.

(4) Small, semi-bipocillate anisochele (Pl. 26. fig. 12d) ; mostly in
rosettes ; measuring about 0:01 mm. in length ; the free end of the markedly
curved shaft is expanded into a flat, saucer-shaped fimbria ; in front view
this appears as a narrow, slightly curved cross beam at the outer end of the
shaft ; the fixed end is quadrilateral and of the chelate type. This form of
microsclere is probably intermediate between an anisochela and a bipocillate.

(5) Small, palmate anisochelee of the Lophon type (Pl. 26. fig. 12 e) ; with
only a very minute spur at the small end; these occur scattered singly and
are not so abundant as the other microscleres ; length about 0°017 mm.

This sponge appears to be almost identical with Pseudoesperia enigmatica
(olim L’sperca parasitica Carter [1880]) except for the absence of sandy
fibre so characteristic of Carter’s specimens. We have examined a micro-
scopic preparation of Carter’s type-specimen (7) in the British Museum
collection, and find that the small, semi-bipocillate anisochele are certainly
present, though not mentioned in his description.

Register No, and Locality. 11. 15, Wooded Isle.

33. PsrUDOESPERIA TRICHOPHORA Nn. sp.

The single specimen consists of a small, light-brown fragment encrusting
a Hircinia sp.?

The main skeleton consists of a reticulation of stout, ill-defined spicular
fibre with many loose megascleres scattered irregularly ; radially arranged
brushes of tylostyles lie vertically to the surface.

SPONGES FROM THE ABROLHOS ISLANDS. 503

The spiculation agrees closely with that of Psewdoesperia carteri, except
that the sigmata are entirely replaced by trichodragmata.

Spicules :—-(1) Tylostyli; size about 0°34 by 0:006 mm.

(2) Large, quadridentate anisochele ; about 0°056 mm. in length.

(3) Small, semi-bipocillate anisochele, about 0-01 mm. in length.

(4) Small, palmate anisochelee, about 0°012 mm. in length.

(5) Trichodragmata ; short, compact bundles, measuring about 0°026 by
0:006 mm.

It is an interesting fact that this species differs from Pseudoesperia
carteri only in the replacement of the sigmata of the former by trichodrag-
mata. These spicules are so widely and sosporadically distributed throughout
the tetraxonid sponges that it is extremely difficult to estimate their taxonomic
value.

The peculiar form of the quadridentate anisochelee and the occurrence of
what we have called the “ semi-bipocillates ” seem to justify the retention of
Carter’s genus Pseudoesperia.

Register No. and Locality. 11. 12 a, Wooded Isle.

34. ESPBRELLA PLUMOSA (Carter).
(For literature and synonymy vide Dendy | 1916 A }.)

The material consists of a small, whitish-yellow fragment encrusting Hippo-
spongia intestinalis (Lamarck). No further description is necessary.

Previously known Distribution. Mauritius and Mergui Archipelago (Carter);
Ceylon, Okhamandal (Dendy).

Register No. and Locality. VI. 136, Sandy Isle.

35. BrEMNA TUBULATA Dendy.

Desmacella tubulata Dendy [1905, 1916 A}.
Biemna microxa Hentschel {1911}.
Toxemna tubulata Hallmann [1917].
Biemna tubulata Dendy [1921 B}.

The material consists of a couple of fragments of thin-walled tubes, the
outer surface of which appears slightly granular, while the inner bears
numerous minute openings of exhalant canals. (Colour in spirit brownish
yellow ; texture very soft and fragile.

The skeleton arrangement and spiculation agree very closely with those of
previously described specimens.

We consider Biemna microxa Hentschel [1911] to be identical with this
species.

Previously known Distribution. Gulf of Manaar, Okhamandal, Indian
Ocean (Dendy) ; S.W. Australia (Hentschel).

Register No. and Locality. VIII. 3, Sandy Isle.

504 PROF. A. DENDY AND MISS L. M. FREDERICK ON

36. EcHINODICTYUM BILAMELLATUM (Lamarck) Itidley.

Spongia bilamellata Lamarck | 1813].

. Echinodictyum bilamellatum Ridley {1881}.
Echinonema vasiplicata Carter [1882 B}.
Kalykenteron elegans Lendenfeld [1888].
Kalykenteron silex Lendenfeld [1888].
Thalassodendron typica Whitelegge [1901].
Echinodictyum bilamellatum Hentschel [1911].
Echinodietyum elegans Hallman {1912}.

There is in the collection one beautiful, cup-shaped specimen, 110 mm. in
height, with a short stalk and very slightly folded walls. The colour in
spirit of the inner surface is almost black, that of the outside grey merging
to violet.

The skeleton arrangement and spiculation agree very closely with those of
the specimens described by Ridley and Hentschel.

We have also examined two dry specimens of this species from the Dampier
Archipelago, N.W. Australia, which are in Professor Dendy’s collection at
King’s College, London. These are both cup-shaped with much folded walls.

Previously known Distribution. N.W. and W. Australia (Ridley) ; Fre-
mantle, S.W. Australia (Carter) ; S.W. Australia (Hentschel) ; E. and W.
Australia (Lendenjeld) ; HE. Australia (Whatelegge, Hallmann).

Register No. and Locality. V., Albrolhos Islands.

37. ANCHINOE FICTITIOIDES n. sp. (PI. 25. fig. 13; Pl. 26. fig. 13.)

There are two specimens in the collection, each forming an almost
continuous, vertical, fan-shaped lamella, with an irregularly indented upper
margin. The larger specimen (R.N. VI. 4, Pl. 25. fig. 13), which measures
150 mm. in greatest height, 95 mm. in greatest width, and has an average
thickness of 7 mm., has two conspicuous oval fenestree ; the other specimen
has only one fenestra. The width of both specimens diminishes gradually
below to a narrow base of attachment. The colour of the larger specimen
in spirit is pale greyish yellow, that of the smaller light brown ; texture
fibrous, tough and resilient. The surface is glabrous.

On both surfaces numerous circular, slightly raised, pustule-like, cribriform
pore-areas are closely but irregularly scattered. The larger of these pore-
areas measure 3 or 4mm. in diameter ; they vary, however, in size, number,
and conspicuousness in different parts of the same specimen.

A few small, marginal vents are seen in both specimens. A ramifying
and anastomosing system of inhalant and exhalant subdermal canals can be
seen through the thin dermal membrane.

In the deeper parts of the sponge the main skeleton consists of an irregular
network of horny fibre cored to a varying extent by smooth megascleres
(tornotoxea) and echinated by acanthostyli. Dense fibres, consisting only of
tornotoxea, ascend somewhat irregularly towards the surface, and the

radially arranged terminal spicules of these fibres form a fringe round

SPONGES FROM THE ABROLHOS ISLANDS. 505

the pore-areas. The sieve-membrane of these areas is free from spicules,
except for numerous isochelee.

Spicules :—(1) Tornotoxea (Pl. 26. fig. 13a); smooth, slender, usually
straight ; ends hastate but unequal; size about 0-4 by 0-006 mm.

(2) Acanthostyli of two kinds—(a) (PI. 26. fig. 136) with short, straight
or sometimes slightly curved shaft, which is slightly swollen at one end and
sharply pointed at the other, covered with small, slightly recurved spines ;
size about 0:14 by 0°013 mm.: (4) (Pl. 26, fig. 15¢) with long, slightly
curved shaft, the curve being most noticeable just above the slightly swollen
base ; the spines are very small and only extend for about two-thirds of the
shaft from the base, thus leaving the sharply-pointed apical end quite smooth.
These longer acanthostyles, which measure about 0°35 by 0:01 mm., are found
only in the larger specimen (R.N. VI. 4).

(3) Tridentate isochelee (chele arcuate) (Pl. 26. fig. 13d), with stout,
slightly curved shaft and short teeth ; measuring about 0-024 mm. in length.
These spicules are very abundant in the dermal membrane, especially in
the pore-areas, but occur also scattered through the choanosome.

This species agrees very closely with the Huropean Anchinoé fictitius
(Bowerbank) J. Stephens [1921] as regards its pore-areas and spiculation,
but differs widely in its mode of growth, which closely resembles that of
Yvesia (Grayella) spinulata Hentschel [1911], also a S.W. Australian
and Indian Ocean species (vide Dendy [1921 A]). Indeed, Yvesia spinulata
differs from Anchinoé fictitiodes in little if anything more than the replace-
ment of the acanthostyles by acanthoxea, and may probably be regarded as a
direct derivative of the latter species.

Register Nos. and Locality. V1. 4, VI. 11, Sandy Isle.

38. DENDORICELLA scHMiDII (Aedley).
Crella schmidti Ridley [1884].
Damiria australiensis Dendy [1896].
Damiria Schmidti Topsent [1897 ].
Damiria australiensis Lindgren {1897, 1898).
Dendoricella Schmidti Hentschel {1911, 1912 A}.

The larger of the two specimens (R.N. IV. 9) in the collection consists of
five erect, digitiform processes springing from a thin, common base; the
processes measure about 8 mm. in diameter, and four of them divide at their
ends into a varying number of slender, secondary, digitiform processes. The
second specimen (R.N. ITI. 10) consists of a portion of a single lobe, dividing
above into four slender, digitiform processes. The general surface of the
sponge is covered with narrow, meandering ridges running longitudinally ;
the dermal membrane, which is smooth and transparent, is interrupted by small
pore-sieves, visible with a lens. A number of small oval vents, sometimes
lying in slight depressions, are scattered singly on the processes. Colour in
spirit light greyish brown ; texture very soft, spongy, friable.

506 PROF. A. DENDY AND MISS L. M. FREDERICK ON

The skeleton arrangement and spiculation agree with those of previously
described specimens.

Spicules:—(1) Oxea ; measuring about 0°23 by 0:01 mm.

(2) Amphitylota; some straight, others slightly undulating, measuring
about 0°31 by 0-007 mm.

(3) C-shaped sigmata of two sizes, the larger measuring about 0:039 by
0°0028 mm., the smaller 0-02 by 0:0014 mm.

(4) Tridentate isochele (chelee arcuate) of two sizes, the larger being
about 0:034 mm. and the smaller about 0°022 mm. in length.

Previously known Distribution. Port Jackson (Ridley); Port Phillip
Heads (Dendy) ; Amboina ( Topsent) ; S.W. Australia (/Tentschel).

Register Nos. and Locality. II. 10, 1V. 9, Wooded Isle.

39. TRACHYCLADUS L&VISPIRULIFER Carter.
Trachycladus levispirulifer Carter (1879, 1885-6).
Trachycladus levispirulifer Dendy {1897}.
(For further possible synonymy see Hallmann’s species [1916].)

The external form of the single specimen resembles closely that of Trachy-
cladus digitatus var. clavatus Hallmann [1916]. The branches, which
anastomose freely, are subcylindrical, and their surface appears minutely
conulose owing to the up-pushing of the dermal membrane by the ends of
the impinging skeleton fibres. The inhalant pores are scattered singly, closely
and for the most part equidistantly over the entire surface, giving it a
minutely reticulate appearance. ‘The small oscula are scattered sparsely and
irregularly. Colour in spirit whitish brown ; texture tough and dense.

The skeleton arrangement calls for no comment.

Spicules :—(1) Oxea; smooth, curved, of nearly uniform diameter
throughout their length, bluntly pointed, measuring about 0:47 by 0:016 mm.;
very rarely stylote.

(2) Spinispiree ; minutely spined, corkscrew spicules, usually of two com-
plete turns, measuring about 00135 mm, in length ; very abundant. Simpler
forms, C- and S-shaped, are found frequently.

(3) Microstrongyla; of two kinds—(qa) slender, centrotylote, 0:019 mm.
long ; (6) stout, rarely centrotylote, 0:016 by 0°003 mm. ‘These microscleres
are rare.

We have examined preparations of this species in Mr. Carter’s own
cabinet, and feel justified in identifying with it the Abrolhos specimen,
although the oxea of the latter are larger and the microstrongyla less
abundant.

Hallmarn [1916] has endeavoured to distinguish between a number of
Australian species, some of which he describes as new. We are inclined to
think, however, that the differences between Hallmann’s specimens are not
sufficiently great to justify specific distinction being made, and that probably

e

SPONGES FROM THE ABROLHOS ISLANDS. 507

all the known Australian specimens of Trachycladus, as well as Lendenfeld’s
Spirophora digitata and Spirophora bacterium, should be included in Trachy-
cladus levispirulifer Carter.

Previously known Distribution. South-West Australia (Carter) ; Port
Phillip Heads, Australia (Dendy).

Register No. and Locality. II. 9, Wooded Isle.

40. SIGMOSCEPTRELLA FIBROSA Dendy. (Pl. 25. fig. 14.)
Spirastrella fibrosa Dendy [1897].
Sigmosceptrella fibrosa Dendy {1921 A].

The single fine specimen in the collection, which measures 70 mm. in
height, is sessile and lobodigitate. Surface sub-glabrous, almost smooth in
parts, slightly conulose or nodular in others. The close-set inhalant pore-areas
give the surface a minutely reticulate appearance. They overlie large, funuel-
shaped chones in the cortex, which is slightly fibrous and measures up to
0:4 mm. in thickness. The roof of each clone is not merely a dermal membrane,
but is moderately thick, strengthened by microscleres (sigmodiscorhabds),
and pierced by individual pores which take the form of short canals. The lower
part of the chone is constricted to a relatively small opening, which leads
into a spacious subcortical crypt ; from this, inhalant canals, provided with
diaphragms along their length, penetrate the choanosome. The canal system
is eurypylous, the flagellate chambers spherical and measuring up to 0:05 mm.
in diameter. The larger exhalant canals and the wide oscular tubes are sur-
rounded by a thick layer of gelatinous mesoglea. Vents small but
prominent, scattered singly on the upper margin of the sponge, sometimes at
the ends of small conical projections. Hach vent is much smaller than the
diameter of the oscular tube out of which it opens, so that the latter appears
to have a thin roof with the vent in the middle. Colour in spirit yellowish
white ; texture firm, compact, resilient.

The skeleton arrangement agrees closely with Dendy’s original description.
Between the chones the cortex is packed with microscleres, intermingled in
places with the ends of the radial bundles of megascleres.

Spicules:—(1) Styli; straight, evenly rounded or slightly enlarged at the
base and sharply pointed at the apex, very uniform in shape and size,
measuring about 0°34 by 0-01 mm.

(2) Isodisecorhabds (sigmodiscorhabds) ; measuring in total length about
0:05 mm.; the diameter of the smooth shaft is about 0:008 mm., that of a
whorl with the spines included 0°025 mm. The development and adult form
of these spicules have been described and figured by Dendy in “The
Tetraxonid Sponge Spicule—A Study in Evolution” [1921 A}.

Previously known Distribution. Port Phillip Heads (Dendy).

Register No. and Locality. VI. 2, Sandy Isle.

508 PROF. A. DENDY AND MISS L. M. FREDERICK ON

41. SPIRASTRELLA VAGABUNDA Firdley.
Spirastrella vagabunda Ridley [1884].
(For literature and possible synonymy vide Dendy [1905] and Vosmaer [1911].)

The larger of the two specimens, which measures about 60 by 20 by
25 mm., is elongated and tubular, with a hollow digitiform process (which
has been cut off) arising from one end. T'wo large, irregularly shaped vents
are present, and inhalant pore-sieves are visible over a small portion of the
smooth but uneven surface. The second specimen is a small tubular fragment
with thick walls. Colour in spirit light brown ; texture firm and compact.

The skeleton arrangement and spiculation agree closely with those of the
type.

The Abrolhos specimens come very close to Spirastrella vagabunda var.
tubulodigitata Dendy [1905], but the tylostyles of the former are shorter
and stouter.

Previously known Distribution. Torres Straits (Adley) ; Indian Ocean
(Dendy).

Register Nos. and Locality. VI. 10, VI. 15 a, Sandy Isle.

42. AapTos AAPTOS (Schmidt).
Ancorina aaptos O. Schmidt [1864).
Aaptos adriatica Gray [1867].

Ancorina aaptos O. Schmidt [1868].
Trachya pernucleata Carter | 1870}.
Tuberella tethyotdes Weller {1880}.
Tethyophena silifica O. Schmidt [1880].
Tuberella tethyoides Vosmaer [1887 ].
Aaptos adriatica Vosmaer [ 1887 }.
Amorphina Duchassaingi Topsent [1889].
Suberites spissus Topsent [1892].
Tuberella Duchassaing: Topsent {1894},
Tuberella tethyoides Topsent {1896}.
Suberttes aaptus Lendenfeld [1896].
Suberites aaptus Topsent [1898].
Tuberella aaptos Topsent [1898, 1900}.
Tuberella aaptos Wilson {1902}.
Tuberella aaptos Hentschel (1909, 1912).
Tuberella aaptos Dendy [1916 A}.
Tuberella aaptos Babié (1922).

The larger of the two specimens (R.N. ITI. 5) is irregularly subspherical,
measuring 45 by 25 by 30 mm. ; at one end, in a slight depression, lies a
small group of oscula, and three or four small round oscula are scattered
singly over the sub-glabrous upper surface. Colour in spirit dark brown
outside, light brown inside ; texture compact but compressible.

The other specimen (R.N. VI. 126) is a small, light-brown fragment
25 mm. in length, 15 mm. in breadth, and 8 mm. in thickness, with a flat,
sub-glabrous upper surface, in the centre of which is a single slit-like

SPONGES FROM THE ABROLHOS ISLANDS. 509

osculum raised on a small papilla. Pigment granules are present in the
large specimen, absent in the small one.

The skeleton arrangement and spiculation agree closely with the descrip-
tions given by Topsent and Wilson.

We propose to revive here Gray’s generic name Aaptos [1867], because
we can find no adequate reason why Keller's genus Tuberella [1880]
should stand in place of it, though we regard his species Tuberella tethyoides
as synonymous with Ancorina aaptos Schmidt. We have revised and
brought up to date Topsent’s synonymy list, which we think can be definitely
accepted.

Carter [1870] described a new genus and species of sponge under the
name T’rachya pernucleata ; after examining his preparations we think it
justifiable to identify this species with Aaptos aaptos (Schmidt), and it is
therefore included in the synonymy list.

Carter [1882A, 1886] described two more species of Zrachya, viz.
Trachya durissima and Trachya horrida; these are probably identical with
his original species, Trachya pernucleata.

Trachya globosa Carter [1885-86] and Trachya globosa var. rugosa Carter
[1886] are obviously lipotriwnose Tetillids, having exceedingly long,
slender oxea for mevascleres and sigmata for microscleres.

Carter [1876, 1882 A] proposed a group Polymastina in the Sub-order
Suberitida to include his genus Trachya as well as the typical Polymastia,
referring to Trachya pernucleata and its allies as “ Geodia-like forms of
Polymastia.”

Although Vosmaer, Topsent and Wilson have placed the genus Tuberella
(= Aaptos) in the Tethyidee (Donatiidz), we are inclined to agree more with
Carter’s views ; and so, because of the arrangement of the skeleton, the
absence of any fibrous cortex and the presence of pin-head spicules in some
cases, we now propose to transfer it to the Sigmatotetraxonida, Family
Clavulide, Sub-family Suberitinee.

Previously known Distribution. Mediterranean (Schmidt, Lendenfeld,
Topsent) ; Gulf of Naples (Keller); Gulf of Mexico (Topsent) ; Porto Rico
(Wilson) ; S.W. Australia (Hentschel) ; Aru Islands (Hentschel) ; Okhaman-
dal (Dendy).

Register Nos. and Locality. III. 5, Wooded Isle ; VI. 126, Sandy Isle.

43, POLYMASTIA MAMMILLARIS (O. Ff’. Miller) Bowerbank.
(For literature and synonymy vide Topsent [1900].)

There is a single small specimen, consisting of about six smooth, slender,
thin-walled, hollow, mammiform projections, 10 to 15 mm. in length, rising
from a plate-like base which has either been torn off a stone or from a larger
specimen ; the surface of the base is slightly hispid. The colour in spirit is
dull brown owing to the development of pigment granules.

510 PROF. A. DENDY AND MISS L. M. FREDERICK ON

The skeleton arrangement and spieulation agree with the descriptions
given by Bowerbank and Topsent.

Spicules:—(1) Small tylostyles ; size about 0-17 by 0-008 mm.

(2) Large tylostyles; size about 1:1 by 0:02 mm. ; the size of the head
varies greatly.

Previously known Disiribution. Cosmopolitan.

Register No. and Locality. VI. 12d, Sandy Isle.

44, MEGALOPASTAS ARENIFIBROSA n. sp. (PI. 25. fig. 15.)

There are three specimens of this sponge in the collection. The largest
(R.N. VI. 3, Pl. 25. fig. 15), which must be regarded as the type of the
species, is sessile, erect, and consists of a principal lamella giving off a
number of secondary lamelle of varing size. The specimen is 110 mm.
high and 70 mm. wide, and the thickness of the lamella is about 3 mm.
The second specimen (R.N. VI. 9), which is 65 mm. high, 15 mm. wide and
about 5mm. thick, is a fragment probably broken off the type. The third
specimen (R.N. IV. 4) is small and lamino-clathrous, becoming constricted
towards the base of attachment. The total height is about 50 mm., the
maximum breadth about 55 mm., and the average thickness of the lamella
about 4mm. ‘The surface appears granulated, the granules being really
numerous, minute, close-set conuli. The vents are small, round and single ;
some are arranged marginally, others are scattered, but almost always
confined to one surface of each lamella. Colour in spirit dark brown,
becoming lighter towards the base of the sponge; texture compressible,
resilient, fairly tough.

The main skeleton consists of primary fibres, about 0-09 mm. in diameter,
running vertically to the surface and connected by an irregular network of
secondary fibres about half the diameter of the primaries ; the outermost of
the secondary fibres form a distinct subdermal skeleton network, the fibres
averaging about 0°03 mm. in diameter. The fibres are laminated and “ pithed”
and the primary fibres are cored by numerous sand-grains and broken
spicules.

The dermal membrane, which sometimes contains a good many broken
spicules, is pierced by round inhalant pores, about 0:06 mm. in diameter,
which lead into large subdermal cavities lying in the ectosome, which
contains numerous close-packed, granular, stellate cells. The canal-system is
lacunar ; the flagellate chambers, which are large, sic-shaped, averaging about
0:05 mm. in diameter, are placed fairly close together and are eurypylous.
The mesoglcea of the choanosome is not strongly developed, except around
some of the larger canals, where it is collenchymatous, with stellate cells.
There are also numerous bands of fibrous tissue, composed of granular,
elo gated cells, running in from the ectosome in places and deeply pene-
trating the choanosome.

This species is well characterised by its external appearance and unusually

SPONGES FROM THE ABROLHOS ISLANDS. 511

compact texture, and, above all, by the coring of the primary fibres by
sand-grains, which indicates an approach to the Family Spongeliidee.

Register Nos. and Localities. VI. 3, VI. 9, Sandy Isle ; IV. 4, Wooded
Isle.

45, SPONGELIA DAKINIn. sp. (PI. 25. fig. 11.)

This most interesting specimen forms a very smooth encrustation (about
0°65mm. thick), almost white in colour, over a considerable portion of the
surface of a very fine specimen of Ancorina australiensis (Pl. 25. fig. 11).

There is a well-developed, minutely reticulate cortex, about 0°085 mm.
thick, formed of sand and broken spicules, which giyes the surface a very
uniform appearance and presumably covers the vents, as these are not visible.

The main skeleton consists of columns of broken foreign spicules of
various kinds, but without visible spongin ; the columns run vertically and
sometimes appear to branch slightly as they approach the surface.

The thin, transparent dermal membrane is pierced by numerous round
inhalant pores, which measure up to 0'05 mm. in diameter ; usually there is
only a single pore in each mesh of the cortex. Small subdermal cavities in
the sand cortex lead into large subcortical crypts or inhalant canals in the
choanosome, the mesogloea of which is but feebly developed. The flagellate
chambers are crowded together, sac-shaped, measuring about 0:06 mm. in
diameter ; they are eurypylous and each has several prosopyles. Some of
the larger canals appear to be surrounded by fibrous tissue; these are
probably exhalant.

This species is at once distinguished by its thin, encrusting habit and its
well-developed, reticulate sand cortex.

Register No. and Locality. V1. 66, Sandy Isle.

46. PSAMMOPEMMA CRASSUM (Carter) var.
Holopsamma crassa Carter [ 1885-6}.
Psammopemma crassum Lendenfeld [1889].

The single specimen, which is 35 mm. in height, 20 mm. in width, and
about 5 mm. in thickness, is erect, somewhat lamellar in shape, thickening
considerably towards the base of attachment. The surface is uneven and
sometimes rugose. Oscula are not visible. Colour in spirit light greyish
brown ; texture hard, gritty, incompressible, but friable.

There is a thin, pore-bearing dermal membrane, overlying large subdermal
cavities in the ectosome, which is feebly developed, with collenchymatous
mesoglea. The canal-system is lacunar; the chambers, which have several
prosopyles, are large, sac-shaped, measuring about 0-1 mm. in longer
diameter, and eurypylous. The gelatinous mesogloea is fairly conspicuous ; it
contains stellate cells and also bands of fibrous tissue.

The skeleton consists of a dense mass of sand-grains and broken spicules
with no recognisable spongin, interrupted by irregular patches of soft tissue
more or less free from sand,

512 PROF. A. DENDY AND MISS L. M. FREDERICK ON

The large size of the flagellate chambers, and the highly lacunar character
of the canal-system generally, indicate that this sponge is a true Spongeliid
and not a reduced Tetraxonid that has lost its proper spicules.

Previously known Distribution. Port Phillip Heads (Carter, Lendenfeld) ;
Port Jackson (Lendenfeld).

Register No. and Locality. IV. 11, Wooded Isle.

47. HipPpospoNGIaA INTESTINALIS (Lamarck).
(For literature and synonymy vide Dendy [1905].)

There are two specimens in the collection, light brown in colour, of
elongated tubular form, with their walls perforated at irregular intervals and
the surface slightly conulose. The tubes are about 12 mm. in diameter.

The main skeleton is composed of an irregular network of fairly stout,
amber-coloured, horny fibre, the fibres measuring up to 0:08 mm. in diameter.
There is a surface reticulation of more slender horny fibre. A certain amount
of foreign matter, including a few huge oxeote spicules, has been incorporated
within the sponge.

Previously known Distribution. Mediterranean (Lamarck) ; Zanzibar
(Hyatt) ; Mascarene Islands and Amirante Group (fdley) ; Ceylon Seas
(Dendy).

Register Nos. and Locality. VI. 13a, VII. 2, Sandy Isle.

48. CoscINODERMA PYRIFORME Lendenfeld var. a.
Coscinoderma pyriforme Lendenfeld [1889].

There are four specimens in the collection. The largest (R.N. III. 7),
which must be regarded as the type of the variety, is erect, massive, somewhat
compressed laterally, and has evidently been attached to a rock along its base
and a portion of one side; the total height is 50 mm., the maximum breadth
65 mm., and the average thickness 25 mm. A number of sphinctrate oscula,
about 3 mm. in diameter, are arranged along a crest on the upper surface.
The second specimen (R.N. II. 7a), which measures 45 mm. long, 35 mm.
wide, and has an average thickness of 12 mm., is oval and somewhat cushion-
shaped; a number of round sphinetrate oscula, the largest of which measures
2 mm. in diameter, lie round the edge of the convex upper surface. The under
surface is smooth and slightly concave, and has a few small oscula scattered
over it. The third specimen (R.N. VI. 20a) is erect, flabellate, with oval
outline, attached to a rock by a short, thick stem ; it is 35 mm. high, 28 mm.
wide, and about 6 mm. thick; there is a single osculum on one side. The
fourth specimen (R.N. VI. 22) is a small oval fragment which has probably
been torn off a larger one.

The surface is for the most part minutely conulose, the conuli being more
marked in some places than others, while in parts it is quite smooth; there

SPONGES FROM THE ABROLAOS ISLANDS. 513

are also narrow meandering grooves, covered in life by the thin dermal
membrane. Colour in spirit purplish brown in parts, lighter brown in others ;
texture compact and compressible, resilient, but tough.

There is a thin but well-developed cortex of sand and broken spicules ; this
cortex is usually of a minutely reticulate nature, the meshes being about
0:13 mm. in diameter and containing inhalant pore-areas. In other parts the
cortex seems continuous.

The skeleton consists of primary fibres about 0:05 mm. in diameter, running
vertically to the surface and containing much foreign matter (broken spicules);
the secondary connecting fibres, which average about 0:02 mm. in diameter,
form a close network with somewhat irregular, polygonal meshes about
0:2 mm. wide.

There is a very well developed collenchymatous ectosome, in which lie
large subdermal cavities. The canal system resembles closely that of
Euspongia. Some of the larger canals in the choanosome are surrounded by
a thick layer of collenchyma. The flagellate chambers are small, subspherical,
and about 0:03 mm. in diameter; they are either eurypylous or with very short
canaliculi, and closely packed together. The mesogloea between them is very
finely granular. Embryos are present, some in an advanced state of
development.

Hyatt [1877] described two sponges, Spongelia incerta and Spongelia
spinosa, which Lendenfeld [1889] makes synonymous with his Coscinoderma
pyriforme. We can find nothing in Hyatt’s description to justify this.

Register Nos. and Localities. 11. 7a, 111. 7, Wooded Isle; VI. 20a, VI. 22
Sandy Isle.

?

48 a. COSCINODERMA PYRIFORME Lendenfeld, var. 8.
Coscinoderma pyriforme Lendenfeld [1889].

The single specimen, which measures 25 mm. in greatest height, 55 mm.
in greatest width, and has an average thickness of 10 mm., is a flattened,
‘cup-shaped sponge with irregular but entire margin; it has probably been
attached to a rock at several points on its convex lower surface. Colour in
spirit dark brown on the upper surface, greyish brown on the lower ; texture
fine, compact, compressible.

The upper surface is very minutely conulose, subglabrous, and has a well-
marked but thin, minutely reticulate cortex made up mostly of broken spicules.
Inhalant pore-areas lie in the meshes of the cortex. There are no visible oscula
on this surface. The lower surface is smooth and has a thin, continuous
cortex formed of broken spicules, no inhalant pore-areas being visible; a
number of closed oscula are scattered over this surface, surrounded by
radiating subdermal exhalant canals.

The skeleton arrangement and the structure of the soft tissues agree closely
with those of Coscinoderma pyriforme var.«. A large number of small,

LINN. JOURN,—ZOOLOGY, VOL. XXXV, 3

514

dark

PROF, A. DENDY AND MISS L. M. FREDERICK ON

ly staining, spherical cells, about 0:006 mm. in diameter, are found

embedded in the mesogloea ; these cells are most abundant around the various
parts of the inhalant canal-system and congregated in immense numbers in
the inner portion of the ectosome ; they are probably symbiotic algee such as
are known to occur frequently in sponges.

Register No. and Locality. 1V. 5, Wooded Isle.

1889.

1891.

List or LITERATURE REFERRED TO.

Banic, K.—“Monactinellida und Tetractinellida der Adria” (Glasnik hry.
prirod. drustva. g. xxxiii. g, 1921, pp. 1-17).

—— « Motactinellcan inl Tetractinellida des Adriatischen Meeres” (Zool.
Jahrb. Bd. 46, pp. 217-802).

BatsaMo-CRIVELLI, G.—‘ Di alcuni spongiarj del golfo di Napoli” (Milano
Soe. Ital. Atti, vol. v. pp. 284-302).

Bowrrpank, J.S.—‘ A Monograph of the British Spongiade, Vol. II.” (London).

——— ‘Contributions to a General ister of the Spongiade, Part IV.”
(Proc. Zool. Soc. London, 1873, pp. 3-25).

—— “A Monograph of the British Spongiade, Vol. III.” (London).

‘‘A Monograph of the British Spongiade, Vol. IV.” (edited, with
additions, by the Rev. A. M. Norman. London).

Burton, M.—* A Revision of the Family Donatiide.” MS.

Carrer, H. J.—“On two new Species of Subspheerous Sponges, with Observations ”
(Ann. & Mag. Nat. Hist. vol. vi. pp. 176-182).

— “On two new Species of Gummine (Corticiwm abyssi, Chondrilla
australiensis), with special and general Observations” (Ann. & Mag. Nat.
Hist. vol. xii. pp. 17-30),

——— “Descriptions and Figures of Deep-Sea Sponges and their Spicules, from
the Atlantic Ocean, etc.” (Ann, & Mag. Nat. Hist. vol. xviii. pp. 888-410),
——— “Contributions to our Knowledge of the Spongida” (Ann. &. Mag. Nat.

Hist. vol. iii. pp. 284-360).

——— “Some Sponges from the West Indies and Acapulco, in the Liverpool
Free Museum, ete.” (Ann. & Mag. Nat. Hist. vol. ix. pp. 266-301, 346-368).

——— “New Sponges, Observations on old ones, and a proposed New Group”
(Ann. & Mag. Nat. Hist. vol. x. pp. 106-125).

—— “Contributions to owr Knowledge of the Spongida. Pachytragida ”
(Ann. & Mag. Nat. Hist. vol. xi. pp. 344-369).

— “Descriptions of Sponges from the Neighbourhood of Port Phillip Heads,

ete.” (Ann, & Mag. Nat. Hist. vols. xv., xvi., xvii., xviii.).

— “Supplement to the Descriptions of Mr. J. Bracebridge Wilson’s
Australian Sponges? (Ann. & Mag. Nat. Hist. vol. xviii. pp. 445-466).

—— “Report on Marine Sponges, chiefly from King Island, in the Mergui
Archipelago, ete.” (Journ. Linn, Soc., Zool. vol. xxi. pp. 61- -84),

CzZERNIAVSKY, VLADIMIR.—‘ The littoral Sponges of the Black and Caspian
Seas: Preliminary Report” (Mosc. Soc. Nat. Bull. vol. lili. pt. 2,
pp. 375-397).

Drnpy, A.—‘‘ Report on a Second Collection of Sponges from the Gulf of Manaar”
Ann. & Mag. Nat. Hist. vol. iii. pp. 73-99).

——— ‘Monograph of the Victorian Sponges: Part I. The Organisation and
Classification of the Calearea Homoccela, with Descriptions of the Victorian
Species” (Trans. R. Soc. Vict. vol. iii. pp. 1-82).

1892.
1895 A.
1895 B.

1896.
1897.
1905.

1913.

1916 A.

19165,

1914.
1867.
1872.
1912:
1914.

1916.

1917,

SPONGES FROM THE ABROLHOS ISLANDS. 515

Drnpy, A.‘ Synopsis of the Australian Calcarea Heteroccela, with a proposed
Classification of the Group, and Descriptions of some new Genera and Species”
(Proc. Roy. Soc. Vict. (new series), v. pp. 69-116).

—— “Studies on the Comparative Anatomy of Sponges: V. Observations
on the Structure and Classification of the Calcarea Heteroccela” (Q.J. M.S.
(n.s.), No. 138, pp. 159-257).

—— “Studies on the Comparative Anatomy of Sponges: VI. On the Anatomy
and Relationships of Lelapia australis, a living Representative of the fossil
Pharetrones” (Q.J.M.S. (n.s.), No. 142, pp. 127-142).

—— “Catalogue of Non-Caleareous Sponges collected by J. Bracebridge
Wilson, ete., Part 2” (Proc. Roy. Soc. Victoria, vol. viii. pp. 14-51).

—— “Catalogue of Non-Calcareous Sponges collected by J. Bracebridge
Wilson, etc., Part 3” (Proc. Roy. Soc. Victoria, vol. ix. pp. 280-259).

——— “Report on the Sponges collected by Prof. Herdman at Ceylon in
1902” (Report Pearl Oyster Fisheries, Roy. Soc. London, Part 3, pp.57-246).

——— “Report on the Calcareous Sponges collected by H.M.S. ‘Sealark’
in the Indian Ocean” (Trans. Linn. Soc. (ser. 2), Zool. vol. xvi. 1918,
pp. 1-29).

—— “Report on the Non-Calcareous Sponges collected by Mr. James Hornell
at Okhamandal, etc.” (Report to the Government of Baroda on the Marine
Zoology of Okhamandal in Kattiawar, Part 2, pp. 93-146. London: Williams
& Norgate). ;

— “Report on the Homosclerophora and Astrotetraxonida collected by
HLM.S. ‘Sealark’ in the Indian Ocean” (Trans. Linn. Soc. (ser. 2), Zool.
vol. xvii. pp. 225-271), '

—— “The Tetraxonid Sponge Spicule: A Study in Evolution” (Acta
Zoologica, vol. ii. pp. 95-152).

—— “Report on the Sigmatotetraxonida collected by H.M.S. ‘Sealark’ in
the Indian Ocean” (Trans. Linn. Soe. (ser. 2), Zool. vol. xviii. pp. 1-164).
Drnvy, Artuur, and R. W. Harotp Row.— The Classification and Phylogeny
of the Calcareous Sponges, with a Reference List of all the described Species,

systematically arranged” (Proc. Zool. Soc. London, 1913, pp. 704-818).

Ferrer Hernanprz, I'.—‘Hspoujas del Cantabrico. Parte segunda: III.
Myxospongida, IV. Tetraxonida, V. Triaxonida’ (Trab. Mus. Nac. Cience.
nat., Madrid, Ser. Zool. No. xvii.).

Gray, J. .—“ Notes on the Arrangement of Sponges, with the Descriptions of
some new Genera” (Proc. Zool. Soc. 1867, pp. 492-558).

Harcren, KH.“ Die Kalkschwiimne: eine Monographie” (Berlin).

Hartmann, J. F.—“ Report on the Sponges obtained by the F.I.S. ‘Mndeavour’
on the Coasts of New South Wales, Victoria, South Australia, Queensland
and Tasmania. Part I.” (Commonwealth of Australia Fisheries. Zool.
Results of the Fishing Experiments carried out by the F.1.S, ‘ Endeavour,’
1909-1910, Part IL. pp. 115-300).

—— “A Revision of the Monaxonid Species described as new in Lendenfeld’s
Catalogue of the Sponges in the Australian Museum. Part I.” (Proc. Linn.
Soc. N.S.W., Sydney, vol. xxxix. pp. 263-315).

‘CX Revision of the Genera with Microscleres included, or provisionally
included, in the Family Axinellidee; with Descriptions of some Australian
Species. Parts 1 and 2” (Proc. Linn. Soc. N.S.W. vol. xli. pp. 453-491 &
495-582).

—-— “Op. cit. Part 3” (Proc. Linn, Soc. N.S.W. vol. xli. pp. 684-675).

37

1898.

1902:
1900.
1893.

1883.
1881.

PROF. A. DENDY AND MISS L. M. FREDERICK ON

Hara, J.—“On a new species of Calcareous Sponge, Lelapia nipponica”. (Ann.
Mag. Zool. Soc. Tokyo (Tokyo Dobutsugaku Zashi), vol. vi. pp. 3859-370, pl.viii.).

HenrscHert, K.—“Tetraxonida, Th. 1” (Die Fauna Sitdwest-Australiens,
Bd. ii. pp. 847-402. Jena: G. Fischer).

—— “Tetraxonida. Th. 2” (Die Fauna Siidwest-Australiens. Bd. iii,
pp. 279-393. Jena: G. Fischer).

—— “Kiesel- und Hornschwimme der Aru- und Kei-Inseln” (Abb.
Senckenberg nat. Ges. Bd. xxxiv. pp. 291-448).

Hozawa, Sans1.—‘ On a new Genus of Calcareous Sponge” (Annotationes
Zoologicze Japonenses, vol. x. article 18).

Hyarr, A.—“ Revision of the North American Poriferee, Part II.” (Boston
Soc. Nat. Hist. Mem. vol. ii. pp. 481-554).

Krier, C.—‘‘ Neue Coelenteraten aus dem Golf von Neapel” (Archiv. Mikrosk.
Anat. vol. xviii. 1880, pp. 271-280).

Kirkpatrick, R.—‘On the Sponges of Christmas Island” (Proc. Zool. Soe.
London, 1900, pp. 127-141).

KO6.iircEr, A. von.—“ Icones Histiologicee oder Atlas der vergleichenden Gewe-
belehre. I. Der feinere Bau der Protozoen”’ (Leipzig).

Lamarck, J. B. P. A. de M. de.—Sur les Polypiers empatés: Eponges” (Ann.
Mus. Hist. Nat. Paris, vol. xx. pp. 294-312, 370-386, 432-458).

LENDENFELD, R. von.—‘‘ A Monograph of the Australian Sponges. Part IIL:
The Calcispongize” (Proce. Linn, Soc. N.S.W. vol. ix. 1885, pp. 1088-1150).

—— “A Monograph of the Australian Sponges. Part IV.: The Myxo-
spongie” (Proc. Linn. Soc. N.S.W. vol. x. pp. 3-22).

——— “Die Chalineen des australischen Gebietes” (Zool. Jahrb. vol. ii.
pp: 723-828).

—-— “Catalogue of Sponges in the Australian Museum” (Sydney, N.S.W.).

-——— ‘(A Monograph of the Horny Sponges” (Published for the Roy. Soe.
by Triibner & Co.).

—— “Die Clavulina der Adria” (Nova Acta Acad. Leopoldino-Carolinee,
T. Ixix. pp. 1-251).

——— “Spongien von Sansibar” (Wiss. Ergebn. Madagaskar Ostafr, Abh.
Senckenb. nat. Ges. Bd. xxi. pp. 93-1388).

——— “Tetraxonia” (Das Thierreich, Lief. xix.).

—— “Die Tetraxonia” (Wiss. Ergebn. deutsch. Tiefsee-Exp. Bd. xi. Lief. ii.).

LinperEn, N.G. “Beitrag zur Kenntnis der Spongienfauna des Malayischen
Archipels und der chinesischen Meere” (Zool. Anzeiger, Bd. xx. pp. 480-
487).

—— “Beitrag zur Kenntnis der Spongienfauna des Malayischen Archipels
und der chinesischen Meere” (Zool. Jahrbuch, Abth. Syst. Bd. xi. pp. 283-
378).

Lunpsreck, W.—‘ Porifera, Part I.” (Danish Ingolf-Expedition, vol. vi.
Copenhagen).

Mincur, A. E.—“ Porifera” (‘A Treatise on Zoology,’ edited by E. Ray Lankester).

Ostroumov.— Liste des Spongiaires de la Mer Noire tirée des travaux de
Mr, Czerniavsky” (Cong. Zool. 2° sess. Moscou (1892), part 2, pp. 159-
160).

PortsaErr, N. pE—“ Calearea” (‘ Challenger’ Reports, Zoology, vol. viii.).

RIDLEY, 8. O.—‘ On the Genus Plocamia Schmidt (Dirrhopalum), and some other
Sponges of the Order Echinonemata” (Journ, Linn. Soc., Zool. vol. xy.
pp- 476-487, 493-497).

1884.

SPONGES FROM THE ABROLHOS ISLANDS. 517

Ripiey, 8. O.—‘ Spongiida” (Report on the Zoological Collection made in the
Indo-Paeifie Ocean during the yoyage of H.M.S. ‘Alert’ in 1881-2, pp. 366—
482, 582-630).

Rivrey, 8. O., and Drenpy, A.—“ Report on the Monaxonida” (‘Challenger’
Reports, Zoology, vol. xx.).

Row, R. W. H.—“ Report on the Sponges collected by Mr, Cyril Crossland in
1904-5” (Reports on the Marine Biology of the Sudanese Red Sea. Journ.
Linn. Soc., Zool. vol. xxxi. pp. 287-400).

Scumipt, O.—* Die Spongien des Adriatischen Meeres ” (Leipzig).

— ‘Supplement der Spongien des Adriatischen Meeres” (Leipzig).

——— “Die Spongien der Kiiste von Algier, etc.” (Leipzig).

——— “Zusatz zu “Neue Ceelenteraten aus dem Golf yon Neapel” von C. Keller’
(Arch. Mikr. Anat. vol. xviii. 1880, pp. 280-282).

Sorzas, W. J.— Report on the Tetractinellida” (‘Challenger’ Reports, Zoology,
vol. xxv.).

SwarrscuEwsky, B.—“ Beitrag zur Kenntnis des Schwamm-Fauna desSchwarzen
Meeres” (Mém. Soc. Nat. Kiew, T. 20).

Tete, J.—“ Mieselschwimms von Ternate, I.” (Abh. Senckenb. Nat. Ges.
Bd. xxv. pp. 17-80).

Torsent, E.—“ Contribution & Etude des Clionides” (Arch. Zool. Exp. et
Gén. y. Suppl. vi. pp. 407-496).

—— ‘Quelques Spongiaires du Banc de Campéche et de la Point-a-Pitre”
(Mém. Soe. Zool. France, II. i. pp. 30-52).

——— ‘Diagnoses d’ ponges nouvelle de la Mediterranée et plus particuliére-
ment de Banyuls” (Arch. Zool. Exp. et Gén. (2) x. pp. 26-32, 65-71).

— “Application de la taxonomie actuelle & une collection de Spongiaires
du bane de Campéche et de la Guadeloupe décrite précédement” (Mém. Soc.
Zool. France, vii. (1894) pp. 27-87).

—-— “Matériaux pour servir 4 l’Etude de la Faune des Spongiaires de France”
(Mém. Soe. Zool. France, ix. 1, pp. 115-133). :

—— “Spongiaires de la Baie d’Amboine” (Rey. Suisse Zool. vol. iv. pp. 421-
487).

——— “Sur les Hadromerina de l’Adriatique” (Bull. Soc. Scient. Méd. Ouest,
T. 7, pp. 117-180).

——— “Etude monographique des Spongiaires de France. III. Monaxonida
(Hadromerina)” (Arch. Zool. Exp. et Gén. (8) viii. pp. 1-331).

——— “Eponges recueillies par M. Ch. Gravier dans la Mer Rouge” (Bull.
Mus. Hist. Nat. Paris, 1906, pp. 557-570).

——— ‘“ponges de San Thomé. Essai sur les genres Spirastrella, Donatia et
Chondrilla” (Archives de Zoologie Expérimentale et Générale, Tome 57
Fascicule 6, pp. 5385-618).

VosmarrR, G. C. J.—“ Porifera”” (Bronn’s ‘ Klassen und Ordnungen des Thier-
reichs,’ vol. ii.).

“ The Porifera of the ‘Siboga’ Expedition. II. The Genus Spzrastrella ”
(‘ Siboga’ Expedition Monographs),

WuirrLEces, T.—‘‘ Report on Sponges from the Coastal Beaches of New South
Wales” (Records of the Australian Museum, vol. iv. pp. 55-118).

Witson, H. V.—*‘ The Sponges collected in Porto Rico in 1899 by the U.S. Fish
Commission Steamer ‘ Fish Hawk’” (Bull. U.S. Fish Comm. 1900, vol. ii.
pp. 375-411).

518

eS

OID op ce to

ra
(=)

11.

Lo

“N

x
. Grantiopsis cylindrica Dendy. R.N. Vil. 1a. x
. Grantiopsis cylindrica Dendy. R.N. VII.16. x
. Grantiopsis cylindrica Dendy. R.N. VI. 17¢. x 2.

. Grantiopsis cylindrica Dendy. R.N. 1.1. x 2.

. Grantiopsis cylindrica Dendy var. fruticosa nov. R.N.IUI4. x 2.
. Lelapia antiquan. sp. R.N. VII. le. x 23.

PROF. A. DENDY AND MISS L. M. FREDERICK ON

EXPLANATION OF THE PLATES.

PuaTE 25,
(All the figures in this plate are from photographs.)

. Leucosolenia grisea n. sp. R.N. VI. 21. x 13.
. Leucosolenia protogenes Heckel. R.N. VIII. 4. x 2.
. Ascoleucetta compressa n. sp. R.N. 11.12. x 2.

Vosmaeropsis mackinnoni n. sp. R.N. IV. 1. 13

tole

3
3.

Ancorina australiensis Carter. R.N. VI. 6a; encrusted by Spongeha dakini n. sp.,
IRIN[G Wl @O, 52 il,

. Reniera aqueduetus Schmidt. R.N. VIL. 3. x 1.
. Anchinoé fictitioides n. sp. R.N. VI. 4. x 3.
. Sigmosceptrella fibrosa Dendy. R.N. VI. 2. x 3.

. Megalopastas arenifibrosa n. sp, R.N. VI. 3. Xx 3.

PLATE 26.

. Leucosolenia grisea n. sp. R.N. VI. 21.

a. Triradiates, x 150; 6. Quadrivadiates, x 150.

, Ascoleucetta compressa n. sp. R.N. II. 12.

a. Large trivadiates, x 60; 6. Small triradiates, x 150; c-e. Quadriradiates,
x 160

. Ascoleucetta compressan.sp. R.N. 111.12. Diagrammatic vertical section through

margin, showing canal-system. X 20,

i.a., inhalant apertures; d.cor., dermal cortex; 2.c., inhalant canal; /2.ch.,
flagellate chamber; e2.0., exhalant opening of chamber; e2.c., exhalant canal;
ose., osculum ; mes., mesogloea.

. Ascoleucetta compressa n. sp. KR.N. III. 12. Portion of similar section. x 60.

e.c., collared cells ; pr., prosopyle; 77., large triradiates of dermal cortex (d.c.) ;
a.r., apical rays of quadrivadiates projecting into flagellate chamber and exhalant
canal. Other lettering as before.

. Ascoleucetta compressa n. sp. R.N. IU. 12. Dermal surface, showing skeleton

arrangement round inhalant aperture, x 100.
tr., trichoxea.

. Vosmaeropsis mackinnont n. sp. R.N. VI. 1.

a, Large oxea, X 40; 6. Subgastral or tubar sagittal triradiates, x 40;
ce. Subdermal pseudo-sagittal triradiates, x 40; d. Trivadiates of the dermal
cortex, x 40; e. Trivadiates of the gastral cortex, x 40.

. Grantiopsis cylindrica Dendy. R.N. I. 1.

a. Trivadiates of dermal cortex, x 60; b. Dermal microxea, x 150; c. Sub-
castral quadriradiates, X 150; d. Reduced tubar trivadiate, x 150; e. Head of
tubar triradiate, showing the two vestigial rays, x 950; / Gastral quadriradiates,
x 150; a7. apical ray; g. Oxeote from oscular fringe, X 150; h. Oscular tri-
radiate, x 150.

Dendy & Frederick.

SHPO wW EisS 11)

JOURN. LINN. Soc. ZooL. VOL. XXXV, PL.25

Huth coll.

ROLHOS ISLANDS.

Dendy & Frederick

Journ. Linn. Soc. Zoot. VoL RXV PES

SPONGES FROM ABROLHOS ISLANDS Huth coll.

ne

Dendy & Frederick.

Sa OLN GRE Sisk

JouRN. LINN. SoG. ZoO0L. VOL. XXXV, P]. 26.

ABROLHOS ISLANDS.

hte.

ae
AE
:

us

JouRN. LINN. Soc. ZOOL. VoL. XXXV P). 26

Dendy & Frederick.

Fig. 8
Fig. 9
Fig. 10.
Fig. 11.
Fig. 12
Fig. 13

SPONGES FROM THE ABROLHOS ISLANDS. 519

. Lelapia antigua n. sp. R.N. VII. Le.

a. Large oxea, x 40; b. Slender oxea from dermal brushes, x 40; c. Normal
sagittal triradiates of dermal cortex, x 60; c’. Subgastral sagittal triradiate,
x 60; d. Alate triradiates of gastral cortex, x 60; e. Alate quadriradiates of
peristome, x 60; 7. “ Tuniny-fork ” spicules, x 60,

. Ancorina brevidens n. sp. R.N. VI. 160.

a. Orthotrizene, x 100; 6. Anatriewne, x 100; c. Large oxeote, x 100;
c’. Small ectosomal oxeote, x 100; d. Microrhabds, x 360; e. Tylasters, x 360.
Halichondria phakellioides n. sp. R.N. VI.1.
a. Oxea, X 100; b. Stylote, x 100.
Phieodictyon abrothosensis n. sp. R.N. II. 9.
Oxea, x 190.

. Pseudoesperia carteri n. sp. R.N. IL. 15.

a. Tylostyli, x 270; 6. Sigmata, x 550; c. Quadridentate anisochela, side
view, X 590; c’. Quadridentate anisochela, face view seen from the back,
x 550; c'’. Young quadridentate anisochela, side view, x 550; d. Small semi-
bipocillate anisochele, x 550; e. Small palmate anisochele, x 550.

. Anchinoé fictitioides n. sp. R.N. VI. 4.

a. Tornotoxea, X 190; 6. Small acanthostyli, x 190; c. Long acanthostyli,
x 190; d. Tridentate anisochele (chelee arcuate), x 650.

ban

re Ne
SAY

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS.

Or
ho
e

Report on Opisthobranchiata from the Abrolhos islands, Western Australia,
with Description of a new parasitic Copepod. By Cuas. H. O’Donocuurn,
D.Sc., F.R.S.C., Professor of Zoology, University of Manitoba, Canada.
(Communicated by Prof. W. J. Daxty, D.Sc., F.L.S.)

[Percy Sladen Trust Expedition to the Abrolhos Islands
under the leadership of Prof. W. J. Dakin. ]

(PLATES 27-30.)

[Read 21st June, 1923.]

From a very cursory examination of the distribution of the Opisthobranchs,
it is evident that the Indian Ocean is very rich in members of this group and
is characterized particularly by certain genera. For this reason we may
look forward to the coasts of Australia furnishing a large number of
interesting forms. Up to the present, however, they have not been studied
in this area with any degree of thoroughness, save in the neighbourhood of
Sydney by Angas (7) and less extensively in South Australia by Basedow
and Hedley (8). In Western Australia it may be said that no systematic
collecting has been done, so that our knowledge of the Opisthobranch fauna
of this coast is very limited.

The present collection, while small in numbers, is, nevertheless, interesting
and, it is to be hoped, useful. Certain of the species themselves appear to be
new, and some of the others, although previously recorded, have only been
incompletely described, so that it has been possible in all cases to give here
further essential information regarding their structure or distribution and to
clear up certain doubtful points. In these ways therefore it will extend our
hitherto meagre knowledge of Western Australian forms and so help to fill
a noticeable gap.

The student of the Opisthobranchiata, and especially the Nudibranchiata,
is generally faced with one of two very serious difficulties. On the one
hand, if he collects living material, he does not as a rule have the oppor-
tunity to dissect it and, more particularly, to examine the radule. On the
other hand, if he is working out a collection in a museum or laboratory,
while he is able to make such dissections and preparations of the radule as
the material allows, he is usually not in a position to recognize the original
colour, or perhaps even shape, of the living creature. I doubt if any other
group of animals is so disappointing when preserved ; anyone used to seeing
them alive, with their beautiful coloration and often graceful shape, would
hardly recognize them in the shrivelled discoloured lumps that they become
after preservation. Hence it follows that unless the two aspects are dealt

522 PROF. ©. H. O'DONOGHUE: REPORT ON

with simultaneously, there is always the possibility of one species receiving
two different names, one from the collector who knows the living animal
and the other from the laboratory worker who has handled only preserved
material—indeed, it would not be difficult to cite instances of this. That this
may occur in Australian material is made possible in the case of the paper
by G. F. Angas on the Nudibranchs of New South Wales (7). This work is
illustrated by a number of excellent coloured figures, from which I should
judge a collector would have practically no difficulty in recognizing the
living forms at once. As no details or figures of the radulz or other internal
organs are given, however, it would be impossible to recognize a number of
the species from a collection of preserved material without colour-notes.

This difficulty may be overcome if the same person works at the forms
from the two aspects, or by the co-operation of two individuals. In the
present case I have been fortunate enough to look through the collection
with Professor Dakin, who gave me colour-notes, which, while brief, should
be sufficient to allow of the easy recognition of the living animal. In two
instances he supplied me with coloured drawings.

It has already been noted that the Opisthobranchs of Australia are but
little known, and they would well repay further study, but it is of the utmost
importance that when collections are being made adequate colour-notes of
the living animal should be taken.

The list of workers who have treated of West Australian forms is very
limited. he earliest records are those of Cuvier, 1804 (31), who described
two species of Nudibranchs brought back by Péron from that coast. Quoy
and Gaimard in 1833 (68) described one further species. Sowerby in 1869
(71) described the shell of a Teetibranch from the Cuming collection in the
British Museum. In 1876 Abraham (1) described another Nudibranch from
the British Museum material, and Bergh (12) one from the Cuming collection
in the British Museum. A species brought back by Dr. Studer in the
‘Gazelle’? was described by Bergh in 1880 (22). Kent (54) in 1897
recorded a single species that actually came from the Abrolhos Islands.
Lastly, in 1917 Odhner (59) described three further Nudibranchs which were
brought back by Dr. Mjoberg’s Swedish scientific expeditions to Australia in
1910-13.

The species recorded therefore are :—

TECTIBRANCHIATA.
Tethys { Aplysia gigantea] (Sowerby), recorded from Swan River.
NUDIBRANCHIATA.
CLADOHEPATICA,
Seyllea pelagica Linn., recorded by Cuvier from Terre d’Edels.

Phyllirhoé lichtensteinii Eschscholtz, recorded by Quoy and Gamard from
Terre d’Hdels.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 523

Armina (Pleurophyllidia) cygnea (Bergh), recorded from Swan River.
Madrella ferruginosa Ald. & Hane., recorded by Odhner from Cape Jaubert.

HOLOHEPATICA.
Hexabranchus imperialis Kent, recorded from the Abrolhos.
Kentrodoris maculosa (Cuvier), recorded from Baye des Chiens marins.
Ceratosoma brevicaudatum Abraham, recorded from West Australia.
Ceratosoma corallinum Odhner, recorded from Cape Jaubert.
Phyllidia varicosa Bergh, recorded from Dampier’s Archipelago.
Trevelyana marginata Odhner, recorded from Cape Jaubert.

This is a small list of only 11 species and the present collection contains
16 species, only three of which have been recorded previously from West
Australia, and so the total now known from the coast is 24——a very small
number when one considers the length of the coast-line and the fact that
there is every indication of its possessing a rich fauna.

The following species are here represented :—

TECTIBRANCHIATA.
Tethys | Aplysia | gigantea (Sowerby).
Tethys | Aplysia] denisoni (Smith).
Dolabrifera pelsartensis, sp. nov.
Berthella plumula (Montagu).

ASCOGLOSSA.

Placobranchus expansa, sp. nov.

NUDIBRANCHIATA.

HOLOHEPATICA.
Spherostoma | Tritonia| dakini, sp. nov.
Hexvabranchus imperialis Kent.
Alloiodoris hedleyi, sp. nov.
Asteronotus fuscus, sp. nov.
Glossodoris [ Chromodoris| westraliensis, sp. nov.
Aphelodoris agfinis Hliot.
Ceratosoma brevicaudatum Abraham.
Dendrodoris {Doriopsis| nigra (Stimpson).
Dendrodoris | Doriopsis| mammosa (Abraham).
Nembrotha purpureolineata, sp. nov.
Notodoris gardineri Hliot.

With such a small list it is hardly profitable to spend much time in the
discussion of the distribution and relationships of its members, for as soon as
any systematic collecting is done the list will be enormously increased.

Professor Dakin has called attention to one or two points in his general
description of the expedition (32) that need brief consideration here. In

524 PROF. CG. H. O DONOGHUE: REPORT ON

the first place, the Abrolhos Islands are probably the most southern coral-
islands in the world. A tropical current from the north and north-east
passes down and bathes their shores, and the available evidence goes to show
that the temperature of the sea round the islands is usually some degrees
higher than that near the adjacent coast—indeed, even in the winter, it rarely
falls below 20° C. He further suggests that if a preponderance of tropical
species is found it will probably be due to this warm current, although the
fishes apparently are southern forms.

Of the Tectibranchs, one (i.e., 7. gigantea) has been recorded previously
from Queensland. The two genera Dolabrifera and Placobranchus are,
on the whole, distinetive of tropical or sub-tropical seas, while Berthella
is perhaps more widely distributed.

It may be stated of the Nudibranchs that, as a general rule, the Clado-
hepatica are characteristic of cool or cold waters, while the Holohepatica
characterise warmer regions. It is therefore probably not without signifi-
cance that all the forms in the present Abrolhos material belong to the
Holohepatica. In passing, too, it may be worth while to notice that, of the
Cladohepatic genera recorded from the coast of the mainland of Western
Australia, i.e., Armina (Pleurophyllidia), Scyllea, and Madrella, the two
former are of world-wide distribution and the last is a more or less tropical
group. All other West Australian forms so recorded are Holohepatic.
Turning again to the present material, the Spheerostomids are universally
distributed, but practically all the other genera are distinctly those of tropical
and sub-tropical seas.

Comparing this list with that of the British coast or of the northern part
of the Pacific Coast of North America, the difference is most striking, and it
is impossible not to recognize that it is far more sub-tropical in character.
Cold-water groups appear to be entirely unrepresented up to the present, and
it is hardly possible that, even if found, they will ever outnumber the tropical
and sub-tropical genera.

We may conclude, therefore, that, as far as the Opisthobranchs at present
known are concerned, the fauna of the Abrolhos Islands is distinctly sub- .
tropical or tropical in its constitution.

The remarkable way in which the ordinary usages of systematic taxonomy
have not merely been ignored but flagrantly violated by many workers on
Nudibranchs has created such a series of pitfalls that even with the utmost
care it is almost impossible to avoid them all. Among the most obvious of
these are examples where species have been given new names, apparently
because the author did not like the previous one, and this although the older
name has many years’ priority, and the identity of the two forms is empha-
sized. Again, several genera have been combined and a new generic name
invented, or the reverse of this may have happened and an old genus split
up to make way for two or more new ones, while the original name has been

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 525

discarded. It is obvious that, if this sort of procedure goes on unchecked, the
ultimate result will be a chaotic muddle, out of which it will be very difficult
to find a way. ‘The present author, in conjunction with Iredale (51a), has
spent a great deal of time in an attempt to straighten out the tangle of the
synonymy of the British Nudibranchiata, and, as a result, certain fairly
widely used generic names have heen replaced by less known ones. The very
obvious eriticism will be made that such changes will cause a certain emount
of inconvenience, and it must be admitted at the outset that this is so. On
the other hand, a few hours’ work in the taxonomy of any group will
soon show that this inconvenience is nothing to the deadlocks that arise
from the slipshod and incorrect use of names. The common remark that
“everyone knows what so-and-so means” is not merely untrue, for the
original describer of the species would not know, but it is frequently a flimsy
excuse for avoiding the work necessary to justify the employment of a
correct terminology. Furthermore, it becomes impossible to discuss such
topics as the comparative anatomy, systematic relationship, and distribution
of the members of a large group of animals unless one can be more or less
certain of the species involved. Wherever changes from the common
usage have been made, the reasons for so doing have been briefly given and a
synonymy provided.

My friend Professor Dakin handed the collection over to me for exami-
nation, and I wish to offer him my sincere thanks not only for so doing but
also for giving the colour-notes mentioned above. TI also desire to thank the
authorities of the British Museum—in particular, Mr. G. C. Robson—for the
privilege of working at the Museum, where ready reference to books and
specimens greatly facilitated my work ; and, lastly, also Professor Dendy,
F.R.S., of King’s College, London, in whose laboratories the dissections,
preparations of the radulee, and the drawings were made.

All the drawings of teeth were made with the camera lucida from my own
preparations. Type-specimens, examples of all the species, and, when
present, the radulee have been deposited in the British Museum.

Order OPISTHOBRANCHIATA H. Milne-Edwards, 1848.

The Gastropoda belonging to the class Euthyneura are all hermaphrodite
and characterised by the detorsion of their visceral mass and nerve-
commissure ; in addition, they generally have two pairs of tentacles and
exhibit a tendency, sometimes marked, to concentration of the nervous
system. They fall fairly naturally into two orders—the Pulmonata, which
are adapted for aerial respiration and life on land, and the Opisthobranchiata,
which are adapted for aquatic respiration and a marine life. The
Opisthobranchs thus come to form a well-defined group ; on the other hand,
however, the division of this order is not quite so simple.

526 PROF, C. H. O'DONOGHUE: REPORT ON

Certain authorities—e.g., Bergh (14 & 17), von Jhering (52 & 53)—recognise
three sub-orders: namely, the Tectibranchiata, the Nudibranchiata, and the
Ascoglossa. The last-named sub-order is characterised inter alia by the
possession of a uniseriate radula with teeth of a special form which, when
worn out, are not ejected, but retained in a sac lying at the base of the buccal
apparatus [hence Saccoglossa von Jhering (52)]. The difficulty in accepting
this classification lies in the fact that the Ascoglossa undoubtedly contain
two separate groups—(1) the Lophocercide: (Lobiger and Lophocercus), which
show certain marked affinities with the Tectibranchs, so that by some
authorities they are included with them, and (2) the Hermeeidee, Phyllo-
branchide, Placobranchide, LHlysiidee, and Limapontiidee, which are
undoubtedly more nearly related to the Nudibranchiata. Eliot (40), to
overcome this, places the Lophocercide with the Tectibranchs and the
remaining families with the Nudibranchs. While this avoids one difficulty
it creates another, for it does not sufficiently mark off these Ascoglossan
forms, which deserve some recognition. Vayssiére (73) includes the
Lophocercidee (Oxyneeide) with the Tectibranchs and the remaining
families he groups as a separate sub-order—the Ascoglossa, allied to the
Nudibranchs. This restricts the application of the term Ascoglossa to the
forms allied to the Nudibranchs, and excludes from it the Lophocercid:e
which are included under the older and original usage of the term and which
are, of course, Ascoglossan. Pelseneer (65) has but two orders in the
Opisthobranchs—the Tectibranchia and Nudibranchia. The latter he divides into
four Tribes, of which the fourth (the Elysiomorpha) is identical with the
Ascoglossa of Vayssitre. While agreeing that the Elysiomorpha constitute
a group of forms related to the Nudibranchs, I cannot accept Pelseneer’s
view that they are highly specialised Holidomorpha. Indeed, for a number
of reasons that need not be entered into here, I am inclined to think that if
there is any relationship between the Hlysiomorpha and Holidomorpha it is
the reverse of this, and the former are to be regarded as the more primitive
and not the derived group.

The difficulty remains, and further work on the doubtful and transitional
forms will be necessary before it can be satisfactorily settled. For the
purpose of the present paper it is intended to adopt the classification set
forth by Vayssi¢re in 1885 (78) and to divide the order Opisthobranchiata
into three sub-orders—namely, the Tectibranchiata, the Ascoglossa, and the
Nudibranchiata, using the term Ascoglossa in the restricted sense as identical
with the Hlysiomorpha of Pelseneer.

Sub-order TECTIBRANCHIATA Cuvier, 1817.

The Tectibranchs are characterised generally by the possession of a lateral
or dorso-lateral gill (ctenidium) on the right side, protected by a fold of the
mantle ; usually a shell, internal or external ; a radula, whose used teeth are

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 527

discarded. ‘The asymmetrical position of the ctenidium involves also a
certain asymmetry of the viscera that is not characteristic of the Nudi-
branchiata.

The main sub-diyisions employed here are those suggested by Fischer in
1884 (41) and adopted by Vayssiére (74) and Pilsbry (67) among others, and
the sub-order is split into three divisions : (1) Cephalaspidea, (2) Anaspidea, and
(3) Notaspidea. Three of the species in the collection belong to the second
group (the Anaspidea) and fall within the one family (the Tethyide). The
other belongs to the last group, the Notaspidea.

(2) TECTIBRANCHIATA ANASPIDEA Fischer, 1884.

Tectibranchs that lack a fleshy head-shield ; the head possesses two or
four folded or split tentacles ; the shell is spiral or plate-like, usually hidden
by mantle, and with a posterior terminal nucleus, rarely absent; the penis
is near right anterior tentacle, the female aperture and vas deferens open
near gill.

Family TETHYIDAl (Apiysirp 2).

The animal is elongate; the shell is in the form of an uncoiled concave
plate, almost or entirely hidden, or absent; the neck and head are narrower
than the body, the mouth is a vertical fissure ; the epipodia or pleuropodia
are recurved over back, forming two lateral or dorsal lobes enclosing mantle
and gill. The genital orifice lies within dorsal slit, communicating by a long
furrow with the invertible penis, which lies near the anterior right tentacle ;
the mouth is armed with horny jaws; the multiserial radula possesses
similar teeth ; the stomach is armed with cartilaginous nodules; the anus is
situated behind the gill.

The name generally used for the family is Aplysiidee from the generic
name Aplysia, but Linneeus in his tenth edition, 1758, uses the term Tethys
for an animal which he briefly defines and refers to a figure in Rondelet, and
this is beyond doubt the ordinary sea-hare usually referred to now as Aplysia.
Later, in the twelfth edition, 1767, he applies the term Aplysia or Laplysia
to the very same creature, and uses the name Tethys for the Nudibranch
commonly known as Tethys fimbria. Pilsbry has dealt fully with this
matter (66), and points out, quite correctly, that the term Tethys is the
proper one for this genus and not applicable to Nudibranchs. He adopts
this nomenclature in his monograph (67, p. 59), but strangely enough retains
the old family-name Aplysiidee, whereas the family should take its name
from the oldest or typical genus of the family The correct family-name,
however, is used by Hedley (50, p. 107), 1918. There is no doubt that Pilsbry
was right in making this change, and the genus is Tethys with Aplysia as a
synonym. It is to be regretted that earlier writers did not go back to the
original literature, but continued to use the word Aplysia and misapply
the term Tethys.

528 ' PROF. C. H. O'DONOGHUE: REPORT ON

Genus Trerays Linn. 1758.

Type by designation, 7. leporina Linn. ; T. limacina Linn.
being unidentifiable.
Synonymy: Apnysra Linn., 1767, and subsequent authors.
Type, Laplysia depilans Linn.

The animal is swollen behind, narrower in front, with rather a long neck
and head, bearing folded tentacles and slit rhinophores as is usual in the
family ; the latter lie about midway between tentacles and dorsal slit. The
pleuropedia arise in front of the middle of the animal’s length, are ample,
freely mobile, free throughout their length or united for a distance behind.
They are functional as swimming-lobes ; their anterior ends are separated.
The mantle nearly covers the gill, having a median tube, foramen, or orifice
communicating with shell-cavity, and produced behind in a more or less
developed lobe or lobes, folded to form an excurrent siphon. The genital
orifice lies under the front edge of the mantle, in front of the gill; an
opaline gland is present a short distance behind the genital opening. The
foot is well developed. The shell is very thin and membranous with a thin
calcareous inner layer; it is nearly as large as the mantle, concave, with
pointed small apex, bears a recurved lamina, and hasa concave posterior sinus.

IT am aware that it is an extremely difficult matter in certain cases to
identify members of the genus Tethys by their shells, which are on the whole
very similar, but the two following species have been placed under the same
names as previously recorded, but poorly described, forms. This has been
done because I have had the opportunity to examine the type-material and in
neither case have I been able to detect significant differences between the
present and the previously described species. Further, the localities are in
each instance sufficiently near to make it probable that they fall within the
limits of the distribution of the respective species. Literature, moreover, is
strewn with nomina nuda due to the multiplication of names founded upon
unsatisfactory descriptions or without sufficient notice of previous work.

Species TeTHYS GIGANTEA (Sowerby), 1869, Conch. Icon. vol. xvii., Aplysia,
pl. 1, sp.1,fig.la&o6. (Pl. 27. fig. 1. Pl. 29. figs. 20-22.)

Body. The animal is very large, quite justifying its name of gigantea, and
its body is plump, high, and somewhat egg-shaped. The visceral hump is
placed relatively far back in the largest part of the body, which narrows and
gets lower as it passes forwards to the cephalic region. The very well-
developed pleuropodial lobes arise a short distance behind the rhinophores,
are freely and widely separable, and unite posteriorly to form a transverse
fold across the hinder end of the body above the short but well-marked
caudal prolongation of the foot. The moderately thick mantle is perforated
by a small aperture over the centre of the shell. It covers the ctenidium,
save for a narrow strip postero-laterally, but more completely than in the

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 529

next species. The anal funnel is strongly marked and protrudes between the
hinder margins of the pleuropodia on the right side of the middle line.
The integument is soft and smooth, but thrown into a series of raised areas
of irregular sizes and shapes, separated from one another by deep narrow
grooves. The whole animal, more particularly in lateral aspect, presents a
very characteristic appearance, recalling to some extent the appearance of
erocodile-hide, but, as just noted, it is quite soft.

The opaline gland occupies a well-defined region below the right anterior
part of the mantle. It opens to the mantle-cavity by a very distinct,
circular or slightly crescentic, single aperture and not bya series of apertures
asin 7. punctata.

Colour. The colour of the preserved specimens is a very deep brownish
grey with black lines running in many of the cracks, but also forming areas
of interlacing tracery under the pleuropodia and in the caudal region.
Professor Dakin informs me that in life the animal was of a deep opaque
purple-green,

Dimensions. The larger specimen measured, in the preserved condition,
15 em. long by 6°5 em. wide and 6 em. high, but of course would have been
much larger in life, as all these animals shrink considerably when preserved.
Judging from the type-shells named by Sowerby, one of the original speci-
mens must have been at least a half or two-thirds as large again. If we
take the estimate made by Hales (33, p. 12) that when preserved 7. punc-
tata rarely measures “more than a third of its length when alive,” and
assume it to apply to 7. gigantea, we see that it is possible that the present
species may reach a length of about 60 cm. (say, 2 ft.).

Head. The head and neck-region of this form are quite well developed,
although noticeably smaller than the body of the animal. The vertical slit-
like mouth has a large auriculate tentacle on each side. These cephalic
tentacles do not meet in the mid-dorsal line above the mouth, but an extension
of each forms a sort of lateral flange passing on each side of the mouth.
The postero-dorsal border of each tentacle is rolled, so as to form a very con-
spicuous groove facing backwards and outwards. Below and completely
hidden by the right oral tentacle is the penis.

Foot. The foot is well developed, broad, and muscular. It is not sharply
marked off from the pleuropodial lobes laterally, and in the larger specimen
its surface presents the same “‘ crocodile skin” pattern as the sides of the
body. At the anterior end it is separated from the head by a well-marked
groove, and it is expanded into large semicircular lobes which meet across
the anterior end. This is present in both specimens, but much more marked
in the large one, where this frontal expansion of the foot projects laterally
almost the width of the head beyond it on each side. The hinder end of the
foot has a similar, but much smaller, expanded lobe borne on each side of the
caudal prolongation.

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 38

530 PROF. C. H. 0 DONOGHUE: REPORT ON

Rhinophores. The rhinophores, although considerably smaller than the oral
tentacles, are very obvious cylindrico-conical structures lying close together
in the dorsal region about halfway between the oral tentacles and the
beginnings of the pleuropodia. Their terminal portion takes the form of a
fairly thin flap folded upon itself so as to produce a deep groove facing
laterally.

Gill or Ctenidium. The large gill is crescentic in shape and forms a lobate
plume. The anterior portion is attached by a wide base along the pallial
cavity and the posterior end is free. Just the edge of the hinder portion
protrudes from beneath the mantle.

Shell. The shell is transparent, very thin and soft, and of a pale horn-
yellow colour. The concentric lines of growth are clearly visible. That of
the smaller specimen measures 62 mm. by 41 mm. The shell of the larger
specimen was considerably larger, but left intact. The largest of the Sowerby
shells in the British Museum was 96 mm. by 79 mm., but itis very dry and
shrivelled.

Labial Armature. The wall at the hinder end of the buccal cavity is lined
by a thin semi-transparent cartilage-like membrane; this takes the form of a
compressed tube, whose diameter is 9 mm. high and 4 mm. at its widest point
and about 5 mm. long. This is strengthened by two flat cushion-shaped
bands, 3°25 mm. wide, of a deep brown colour. ‘hey almost touch dorsally,
but are slightly more widely separated at their ventral ends. Under the
microscope it will be seen that these bands are composed of an enormous
number of tiny cylindrical rods with rounded ends, closely packed together to
form a sort of “ pile’ as in velvet. In some works these bands are spoken
of as “jaws,” but this is not a good term, as they cannot be considered as
either homologous in structure or analogous in function with the jaws of
Nudibranchs.

Radula. The radula is very well developed, and in the smaller of the two
specimens measured 15°5 mm. long by 14°5 mm. wide. The total number of
rows in the radula was from 83-85. The number of teeth in the various
rows was as follows: in the first row 5 (?.¢., 2.1.2); in the tenth row 15
(i.e., 7.1.7); and in the seventeenth row about 119 (2.e., 59.1.59). The
rachidial tooth is large and consists of a basal plate in the form of a trapezium
with a deep notch in the middle of the anterior edge and a leaf-shaped blade.
The blade has a broad median denticle with 3 or 4 lateral spines increasing
in size towards the base and the two lateral smaller denticles each flanked by
one or two spines. ‘The bases of the inner pleurals are roughly rhomboidal
with curved sides. The blade contains two main denticles, of which the
inner is larger and has 5-6 lateral spines. The outer has 3-4 laterals, the
basal one being larger than the others. The pleurals increase in size towards
the outside, reaching their maximum more than halfway out. Here they
are much longer and the two denticles are very large and almost smooth—

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 531

they have a very thin crenulated edge along their basal portions. The teeth
decrease in size rapidly at the margin, and the last one or two consist of a
single elongated basal plate with no spine.

The conspicuous genital aperture is to be found just in front of the anterior
edge of the mantle to the right of the middle line. The seminal groove is
well marked and passes forward around the outside of the right rhinophore
to the penis underneath the right cephalic tentacle.

The anus lies on a fold of the mantle to the right of the middle line behind
the visceral hump. In this region the mantle is somewhat thickened and
produced to form.an anal funnel which projects beyond the pleuropodia.

Notes. Two specimens were obtained from near Fremantle (some 250
miles south of the Abrolhos region), and as Sowerby’s specimens were
also from “‘Swan River” the present ones can probably be regarded as

topotypes.

Species Terays DENISONI (Smith), Rep. Zool. Coll. ‘Alert,’ p. 89 (1884).
(Pl. 27. fig. 2; Pl. 29. figs. 23-25.)

Body. The body is plump, high, and somewhat egg-shaped, being wider at
the hinder end than at the front end, and it passes forwards into a narrower
head or cephalic region. The visceral hump appears to be thrown rather far
back as compared with 7. (Aplysia) punctata. The pleuropodial lobes arise
laterally about as far behind the rhinophores as those are behind the oral
tentacles ; they are moderately large and freely separable. Posteriorly they
unite and form a sort of transverse flange across the hinder end of the body
above the caudal prolongation of the foot. The mantle covers the shell, but
possesses a small central perforation, and it also largely covers the ctenidium,
which is free for a short distance at its postero-lateral end. The anal funnel
is well developed. The integument covering the whole animal is very
smooth, but thrown into wrinkles.

The opaline gland is a well-defined body lying under the right anterior
portion of the mantle and opening to the mantle by a single fairly large pore
and not by a series of apertures as in the Huropean 7. punctata.

Colow. When preserved the animals are of a pale greyish white and the
wrinkles are often black, very similar to those in 7. gigantea. Professor
Dakin informs me that in life they are of an olive-green colour with some
darker mottlings.

Dimensions. The largest specimen when preserved measured 6 cm. long
by 3 em. wide and 3:2 em. high.

Head. This may be regarded as including the head and neck. The mouth
is terminal and takes the shape of a vertical slit; on each side is a fairly
large auriculate tentacle. These pass down on each side of the mouth, and
at their upper border are curled round to form a groove directed posteriorly

and outwards, They are separated in the mid-dorsal line. and show no
30"

§32 PROF. CG. H. O'DONOGHUE: REPORT ON

tendency to fuse and forma sort of veil as in 7. punctata. Below and in
front of the right oral tentacle lies the penis.

Foot. Vhe foot is large and muscular and fairly broad. It is sharply
marked off from the head at the anterior end by a groove, and posteriorly it
is continued as a sort of tail. The pleuropodia arise as dorsally-directed
flaps from the median and postero-lateral region of the foot.

Rhinophores. The rhinophores are two sub-conical projections abovt 12 mm.
high, situated near the mid-dorsal line about halfway between the oral
tentacles and the front end of the pleuropodia. They are deeply grooved
along the terminal portion of their postero-lateral borders.

Gill or Ctentdiwm. The gill is well developed and is a lobate plume of
erescentic shape. It is attached by a wide base along the antero-median
margin of the pallial cavity, and posteriorly it passes out freely and the tip
is exposed to a relatively greater extent than in the preceding species.

Shell. The shell is pale yellow, transparent, with concentric lines of growth
clearly showing, and it measures 28 mm. by 25 mm. (Smith’s specimen
measured 30 by 27 mm.—a fairly close approximation). The postero-dorsal
umbo lies far back on the visceral mass. On the longer, more arched, left
side of the shell lie a ridge and a groove which run parallel with the edge.

Labial Armature. The inner parts of the buccal cavity are lined by a thin
hyaline layer of cartilaginous consistency. This takes the form of a tube
flattened laterally ; its internal dimensions are 5-75 mm. high by 1°5 mm. at
its widest point and it is about 4 mm. long. It is strengthened by two
slightly raised bands 2°5 mm. wide ; these almost touch dorsally, but are a
little more separated ventrally. As in the preceding species they are com-
posed of an enormous number of tightly packed circular rods.

Radula. The radula is well developed and measured 12°5 mm. long by
10°5 mm. wide when spread out on a slide. It is of a deep horn-vellow
colour. The number of rows in the radula is about 48 and the number of
teeth in the first row 3 (7. e., 1.1.1) ; inthe tenth row 45 (2. e., 22. 1. 22) ; and
in the thirty-fifth row about 87 (7. ¢.,43.1.43). The base of the rachidial
tooth is roughly trapezoidal, with a deep rounded notch in the middle of the
anterior border. The blade is leaf-shaped and relatively larger than in
T. gigantea. It has a median spine bearing 3-4 lateral denticles and two
lateral spines. The first of these has two lateral denticles and the other is
smooth. The base of the first pleural tooth is approximately rhomboidal
with curved sides ; its blade consists of two well-marked spines, the larger
on the inside, and each bears a series of from 2-4 flanking denticles. The
pleural teeth get larger as they pass outwards, reaching their maximum well
over halfway out. The blade becomes elongated and the spines much
longer, but they retain their lateral denticles, about 3-5 in number.
Finally, near the lateral margin, first the smaller spine disappears and then

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 533

the larger, so that the last two or three teeth consist simply of a narrow
oval basal plate. Ail the types of teeth differ from the corresponding ones
in T. gigantea.

The genital aperture lies a little to the right of the middle line, just under
or very slightly in front of the anterior edge of the mantle, and from it the
quite distinct seminal groove passes forwards to the penis, which, as noted
above, lies just in front of and below the oral tentacle.

The anus is situated at the hinder end of the visceral hump, practically in
the mid-dorsal line and a short distance away from the hump, upon a fold of
the mantle. In the neighbourhood of the anus the mantle is thickened and
projects outwards in the form of « flap, which is coiled round in such a manner
as to form a funnel, the anal funnel, which in this species is relatively larger
than in 7. punctata. {

Notes. In all, five specimens of this species were obtained: two large ones
about the same size from Wooded Island, two smaller ones with no exact
locality given, and one quite small one from Pelsart Island. Professor Dakin
in his paper on the Abrolhos (82, p. 170), when dealing with Wooded Island,
makes the following reference, presumably to this species : “ Nudibranchs and
Tectibranchs (Aplysia sp.) were extremely common. We could have
obtained hundreds of specimens of Aplysia by merely picking them up as we
waded in the shallow water.”

Smith (70, p. 89) described a form Aplysia denisoni collected by H.M.S.
‘Alert’ in the following manner :—

“Body (in spirit) high, exhibiting a distinct pedal dise, produced pos-
teriorly into a caudal termination. The entire surface dirty whitish, black-
veined in the wrinkles (? stains only). Mantles-lobes moderately large,
commencing in front some distance behind the posterior tentacles and
terminating a little in advance of the cauda. Anterior tentacles large,
cylindrical, with the apical slit not extending halfway down the outer side,
placed a little nearer the oral tentacles than the beginning of the mantle-
lobes. Hyes minute, situated near the outer anterior base of the tentacles.
Shell very thin, straw-colour, 30 mm. long and 27 broad. Animal about
three inches in length in its contracted state.”

This description, while not very exact, applies closely to the present speci-
mens. It was, however, of an animal obtained from Port Denison, (ueens-
land, and I should have hesitated to regard it as identical, had it not been
possible to examine the original type-specimen in the British Museum.
Smith’s specimen, much discoloured by age and not well preserved, and the
shell, dried and somewhat shrivelled, are still available. Taking into account
the difference in condition of the specimens, I can see no way of distin-
guishing between them, and so I have placed the present forms under Smith’s
name, although they come from fairly widely-separated areas.

534 PROF. C. H. O'DONOGHUE: REPORT ON

Genus Donaprirera Gray, Proce. Zool. Soc. London, p. 162 (1847).
Type by tautonomy and designation :
D. dolabrifera Rang, Hist. Nat. Aplys. p. 51, pl. 4. figs. 1-6, ex Cuvier,
Rég. Anim. (edit. i.) ii. p. 398, “1817” for 1816 (name only).
Synonymy: DoxiaBrirer Fischer, Man. Conchyl. 1888, p. 568.

The general form is ovate-oblong or sack-like, tapering towards the tail.
Tentacles and rhinophores are slit and expanded distally, the latter lie nearer
to the front margin than to the dorsal slit. The eyes are as in Tethys.
The pleuropodial lobes arise tar behind the middle of the length, are
contiguous, scarcely mobile, united behind, and enclose a large gill-cavity ;
the dorsal slit is short. The mantle is small, not perforated over the shell,
nor covering much of the gill. The foot is broad, often expanded at the
edges. The genital pore lies in front of the gill, under the mantle-edge.
The shell is small, not spiral, solid and calcareous, and sub-triangular,
trapezoidal or irregularly oblong; the apex is projecting and calloused,
with no spiral tendency. The radula bears large sub-triangular rachidial
teeth, with several denticles on the cusp, and the lateral teeth possess long,
coarsely denticulate cusps.

Species DoLABRIFERA PELSARTENSIS, sp. nov. (Pl. 2%. fig. 3; Pl. 29.
figs. 26-28.)

Body. The body is oblong-ovate, reaching its maximum width behind the
middle and passing backwards to a bluntly pointed tail. The back is arched
and the whole integument is wrinkled, and bears a series of wart-like pointed
projections. The pleuropodial lobes take their origin towards the middle
region of the body ; they are not easily movable, and are separated by a slit
lying slightly to the right of the mid-dorsal line and mainly in the posterior
half of the animal’s length. At the anterior end the lobes are practically
contiguous, but separated by the narrow genital groove. Within the gill-
cavity, the mantle entirely covers the shell and is not perforated; it only
covers a small part of the gill.

Colour. The colour of the preserved specimens is of a uniform dirty
brownish grey, and Professor Dakin informs me that the living animals were
much the same colour, but of a deeper warmer brown.

Dimensions. The larger specimen measured 32 mm. long by 9mm. wide,
and 8°5 mm. high.

Head. The head is relatively small and the front of it is occupied by the
vertical slit-like mouth. It is separated from the front end of the foot by a
groove. Antero-laterally it bears two split auriculate oral tentacles. The
tiny eyes lie just in front of and outside the rhinophores. In the larger of
the two specimens the head is practically withdrawn into the body.

Foot. The muscular foot is well developed and broad. At the anterior end
it is bluntly rounded and, as noted, separated from the head by a transverse
groove. Posteriorly it narrows off and terminates in a blunt point.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 535

Rhinophores. The two rhinophores lie fairly close together, dorsally, just
behind the oral tentacles and nearer to them than to the slit between the
pleuropodia. Their extremities are grooved postero-laterally, but they are
not so dilated as in some members of the genus.

Gill or Ctenidium. The gill is composed of a single lamellated plume with
the lamellee lying longitudinally, and it runs more or less transversely across
the animal.

Shell. The shell was very small, calcareous, and apparently trapezoidal, but
was too crushed to allow of accurate description.

Labial Armature. The labial armature, if present at all, is only very feebly
developed.

Radula. The radula is not large, but the teeth are well developed. The
total number of rows is 31. he first row consisted of 2 (t.e., 1.1.0), the
fifth row of 27 (7.e.,13.1.13), and the twenty-fourth of 57 (¢.e., 28. 1.28).
The rachidial tooth has a broad trapezoidal basal plate, which is incurved on
both its anterior and posterior edges. The blade consists of a median spine
bearing two tiny laterals, and four smaller but almost equally developed spines
lie on each side of this. The basal plate of the inner pleurals is rhomboidal
with incurving long sides, and the blade consists of four large spines of which
that nearest the rachis is usually the largest. Further out the spines become
extremely large, and the one nearest the rachis much the largest, so that the
others are borne on its outer side. The outermost pleurals still have a large
inner spine, but the next one to it is almost as well developed, and so they
appear bifid.

In general, the teeth are somewhat similar in type to those of Tethys, but
they differ sufficiently to be recognised as of a different genus.

The genital aperture lies under the mantle-edge well in front of the .gill.
The seminal groove passes forwards to the penis, which is situated just below
the right cephalic tentacle.

The anus opens on the right side of the middle line behind the gill.

Notes. The two specimens in the collection were both collected on Pelsart
Island.

They do not appear to be referable to any previously described forms, and
so have been recorded here as a new species with the name Dolabrifera
pelsartensis from the island on which they were taken.

(3) TECTIBRANCHIATA NOTASPIDEA Fischer, 1884.

The mantle is well developed and covers the whole dorsal surface. It
contains a somewhat small concave lamellar shell within a large pallial cavity.
The shell may be absent or may itself be hidden by an external shell. The
oral veil is marked and carries two dorsal tentacles (rhinophores). The gills
are pinnate and situated laterally on the right side of the body.

536 PROF. C. H. O'DONOGHUE: REPORT ON

Family PLEHUROBRANCHID.

The Pleurobranchidee have not the large head-sheath of the Bullidee nor
the large parapodia of the Tethyide. They have a well-developed oral veil
on a head marked off from the body. The oral tentacles are only moderately
developed and cleft; the rhinophores are usually considerably larger and
are deeply cleft longitudinally. The back is covered with a thick or a thin
mantle and this generally projects freely over tlie head anteriorly and over
the tail posteriorly. Within the mantle is a more or less large shell (absent,
however, in Plewrobranchea and Oscaniopsis). A single (in Oscanius double)
genital aperture lies at the front end of the right side of the body. Above
or below this, but still anterior to the gill, is 1 prebranchial opening or papilla
leading into a sac. The gill is frequently tripinnate, and below it lies the
urinary aperture. At the hinder end of the strand of tissue attaching the
gill (the gill mesentery) lies the anus (in Pleurobranchwa it is above
the strand). The foot is moderately large and overhung by the mantle
(save in Pleurobranchwa, where it passes over into the side and back of the
body). The sole of the tail-region of the foot often bears a gland.

Genus BurrHeLua Blainville, Man. de Mal. et Conch. 1825, p. 469.
Type by monotypy : B. porosa Blainville, zbed.=(Bulla) plumula
Montagu, Testacea Brit. i. 1803, p. 214, pl. 15. fig. 9.

Type, therefore, 5. plumula (Mont.).

Synonymy: Crurantuus Leach, Synop. Moll. Gt. Brit. 1852, p. 28.
{ PLEUROBRANCHUS (pars) Bergh, Malak. Unters. 1898, p. 117.

The shell is haliotidiform, fairly tough, and shows lines of growth and also
longitudinal strize which, while faint, are nevertheless clearly visible. It is
of a translucent amber-yellow colour.

The animal is plump and oblong ; the mantle covers the whole body, its
margin is entire, its front and hinder ends rounded, and it is separated from
the foot by a deep groove. The gill is bipinnate and of moderate size. ‘The
two dorsal tentacles or rhinophores are inserted side by side between the
mantle and the oral veil, and are auriform and deeply cleft. The foot is
truncated anteriorly and bluntly rounded posteriorly where it projects
slightly beyond the mantle. The labial armature is composed of two plates,
each built up of a large number of irregularly arranged close-fitting spine-
like pieces of chitin. The radula is well developed, without a rachidial
tooth and the numerous pleural teeth have the form of knife-like blades
bearing denticles at their distal extremities.

Species BERTHELLA PLUMULA (Montagu), Test. Brit.1. 1803, p. 214. (PI. 29.
figs. 29 & 30.)

Body. The body is an elongated oval and slightly flattened. The mantle
is well developed and overhangs the body all round, save for a narrow strip at
the hinder end. The cral veil also projects in front of the mantle.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 537

Colour. The colour of the preserved specimen is a translucent yellow-grey,
and Professor Dakin informs me that, in life, it was of a uniform dull yellow
colour.

Dimensions. The preserved example measured 22 mm. long by 14 wide
and 9mm. high.

Head. Vhe head is flattened and covered by the forward extension of the
mantle. It bears a strongly developed oral veil, and this is continued out
laterally into two short triangular expansions, which represent the oral
tentacles and are deeply cleft along their lower outer edge. The mouth is
a round opening under the head, and between it and the anterior extension
of the foot.

Foot. The foot is well developed, and, although fairly broad, it is overhung
along the sides of the mantle, but probably the posterior end is free during
locomotion. The anterior end is abruptly rounded and carried on beyond
the mouth as a shallow flange, and the posterior end is bluntly pointed.

Rhinophores.. The rhinophores are well-developed cylindrico-conical
structures lying under the front end of the mantle at the hinder dorsal region
of the head. They arise practically touching one another in the middle line,
but diverge slightly as they pass forwards and they are deeply sulcate on
their outer margins. Just behind the base of each a well-marked, dull black
eye-spot shows through the skin of the neck.

Gill. The gill is situated well forward on the right side of the body under
the mantle. It is attached at the front or basal end and passes backwards as
an elongated pyramidal structure, bearing’ about 24 lamellee on each side of
a smooth rachis.

Shell. The shell was, unfortunately, too damaged to admit of accurate
description, but it appears to be very similar to that of Berthella | Pleuro-
branchus| plumula as described and figured by various authors.

Lalnal Armature. The labial armature consists of two oblong plates
measuring 3mm. long by 15mm. wide. They are of a horn-brown colour
and are composed of tooth-like elements. Hach consists of a lanceolate
flattened blade coming to a single, sharp, non-denticulate point, and bears,
some way down on each side, a blunt triangular process, but these two
projections are not opposite to one another. ‘he elements agree with those
figured by Bergh (27) and Vayssiére (75).

Radula. When flattened on a slide the pale yellow radula measured 4mm.
by 2mm. It has no rachidial teeth and the pleural teeth are arranged in
78-82 rows; a typical row contains 280-290 teeth and the formula is
(140-145) .0.(140-145). The teeth increase in size from the rachis out-
wards, reaching their maximum well beyond the middle line; they then
decrease slightly, but the outermost teeth, although slender, are still longer
than the innermost. All the-teeth are of very similar shape and stand
practically upright. They have a small base, from which comes off a long

538 PROF, C. H. 0’ DONOGHUE: REPORT ON

thin curved spine like the blade of a knife. At its distal extremity it bears
a set of saw-like denticulations on one side, over about one-third or one-half
its length. These vary in number from tooth to tooth from 7-14 and are of
different sizes on the same tooth.

The genital and prebranchial apertures lie immediately in front of and
below the insertion of the gill. ‘The anus is situated just at the hinder end
of the gill-mesentery.

Notes. This species is represented in the collection by a single specimen
from Sandy Island.

Cheesman (30) has described a form, Pleurobranchus (perhaps Berthella)
ornatus, from New Zealand, but it differs from the animal here described
in having brownish mottlings on the back and dark brown rhinophores.
Unfortunately, details of the radula and labial armature are not given.

According to Bergh (27) the radula of B. | Pleurobranchus| plumula has
100 rows of teeth 150 in a row, or, again (26), the radula is stated to contain
120 rows with about 220 in the row, but it is not clear whether the 150 and
220 in these descriptions refer to the number of pleural teeth on one side of
the radula or on both. Vayssiére (75), however, while he does not state the
number of rows in the radula, gives the formula as 140-155 . 155-140, which
agrees quite closely with that of the present specimen. ‘The form of the
teeth also falls within the limits of the variations figured by different authors
for this species. The radula, therefore, appears to be somewhat variable in
constitution.

The labial armature, as a whole and in its individual elements, and also the
shell, as noted above, agree with previous descriptions. It would appear,
then, that the present specimen belongs to the species B. plwnula, although,
if the agreement were not so close, I should hesitate to refer it to that species
on account of its locality. B. plumula, however, had previously been
recorded from Norway, Great Britain, France, Portugal, the Azores, the
Mediterranean, and Lower California, so that it is of wide distribution.

Sub-order ASCOGLOSSA Bergh, 1876.
(Saccoglossa von Jhering, 1876; Hlysiomorpha Pelseneer.)

Branchiz or gills are absent or poorly developed. No shell is present.
The pharynx is mainly adapted for sucking and does not possess jaws. The
radula is short, uniseriate, and has teeth of a characteristic shape, which, when
no longer required, are passed down into a sac or ascus lying at the base of
the pharynx. The central nervous system is composed of 6 or 7 ganglia
massed around the oesophagus. The otocysts contain only a single otolith.
Two seminal vesicles are usually present. The penis is often armed with a

spine.

On
Ss
Ne)

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS.

Family PLACOBRANCHID A.

The head is flattened, and passes over laterally into short leaf- or ear-shaped
tentacles, which have their lateral margins rolled. The two closely set eyes
show through the skin of the neck. The body is much flaitened and its sides
or mantle-folds expanded into wing-like flaps (these are not epipodial
developments as in Tethys), which are usually held up over the body. Behind
the neck is a noticeable swelling, the pericardial swelling containing the
pericardial cavity and the kidney, and at the right side of this lies the anus.
The dorsal surface of both body and jateral expansions generally bears a series
of low longitudinal folds or lamella. The foot is not sharply marked off
from the body, its edge projects very slightly, and often its anterior end is
bilabiate.

There are no jaws; the radula is short, uniseriate, and has large teeth.
These, when worn, are shed into the inferior radula-sac. The stomach is
short and sac-shaped. The hermaphrodite gland is not discrete, but in the
form of isolated follicles. The penis is protrusible and armed with a curved
spine.

Genus PLacopraNncuts van Hasselt, Ale. Konst en Letter-Bode,
1824, I. Deel, no. 3, p. 34.
Type by monotypy, P. ocellatus v. Hass.

The head is flattened, with a broad front, which bears on each side a short
leaf-like or auriform tentacle whose side-margin is rolled. Two eyes, set
close together, lie on the neck. The body is much depressed and bears two
wing-like lateral expansions ; these are mantle-lobes belonging to the body
and not epipodial appendages as in Tethys ; they ure usually turned up over
the body. Behind the neck is a fairly large pericardial swelling, on the
right side of which lies the anal aperture. The upper surface of the body
and the mantle-lobes bear numerous, longitudinal, parallel, low folds. The
double genital aperture lies behind the right tentacle. The foot is not
distinctly marked off from the body, anteriorly it is divided by a transverse
groove into two equi-sized lips.

The bulbus pharyngeus is much as in Phyllobranchia. The radula is
short and uniseriate, the teeth are large and persistent and passed on into
an inferior sac. The penisis armed with curved spines.

Species PLACOBRANCHUS EXPANSA, sp.nov. (Pl. 27. fig.4 ; Pl. 29. figs. 31-34.)

Body. The body of this form is extremely flat and somewhat elongated.
It passes out without a sharp line of demarcation into the enormously developed
mantle-lobes which project, as leaf-like expansions, many times the width of
the body and are continuous behind with the posterior end of the body. The
whole animal presents the appearance of a flat leaf slightly thickened in the

540 PROF. C. H. O'DONOGHUE: REPORT ON

middle anterior region. It is usual in this genus for the mantle-folds to be
folded over the dorsum, but there was no suggestion of this in the present
species. Indeed, these lobes are much more developed than in other members
of the genus and if folded would far more than overlap. Also their antero-
lateral corner projects well in front of the head, so giving the animal a
characteristic appearance not found in other members of the genus.

Colour. The colour of the preserved specimen is a light brownish fawn
with a large number of tiny black spots scattered irregularly all over it, and
the mantle-margin is bordered all round with a black line about 1 mm. wide.
Professor Dakin informs me that in life the colour was more of a greenish
fawn with the black markings as in the preserved specimen.

Dimensions. The length along the middle line was 23 mm., but the antero-
median corners of the mantle projected about 1:5 mm. in front of this.
Width across the widest point was 21°5 mm.

flead. The head is flattened and abruptly rounded at the anterior end.
On each side is a flattened auriculate tentacle rolled at the lateral margin.
In the specimen the head was somewhat retracted.

Foot. The foot is practically indistinguishable, being represented simply
by the thickening in the antero-median line on the ventral side. It is so
reduced that it suggests that the animal was incapable of creeping.

Branchie.: None.

Radula. The radula is short and uniseriate ; it contains 12-14 teeth. The
tooth has an oblong base with rounded corners, and the upper portion is
cacried on as a long, stout, slightly curved, non-denticulate, almost conical
spine. The bases of the teeth are closely approximated, so that each spine
overlaps the whole of the base of the tooth immediately in front of it, and
about half the next also. ‘he spines lie down upon the succeeding tooth,
which is slightly hollowed to receive them, save in the front tooth, which is
almost upright and appears to be the only one in use.

The dental sac contained twenty or more discarded teeth. Some of these
are quite small, not a quarter of the length of that in use, so that it would
appear that they are carried for a relatively long period of time.

Notes. But one specimen is represented in the collection, obtained from
the shore of Wooded Island.

While it is here referred to the genus Placobranchus, it is, as noted above,
different from the previously described members of the genus in the following
points :—

(1) It has non-denticulate teeth. (2) It has a relatively much greater
extension of the mantle-lobes. (3) It has forward extensions of these lobes.
Rather than make a new genus, however, it is included here, since in other
points it agrees closely with the previously known members.

The large expansions of the mantle and almost total lack of a foot rather
suggest that it may be a swimming form. The name P. expansa is here

5

suggested to call attention to the large mantle-lobes.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 541

Sub-order NUDIBRANCHIATA Cuvier, 1817.

Nudibranchs are marine hermaphrodite Opisthobranchiata without
ctenidium and osphradium, and in the adult state without a shell. The larva,
however, has a shell and operculum. The visceral mass is not marked off
from the foot; the body shows complete or approximate external symmetry
and usually bears plumes or other appendages which assist respiration. The
neryous system is concentrated in a collar behind the buccal bulb, and the
chief ganglia are placed on the dorsal surface of the cesophagus, being often
partially united and sometimes fused into a single mass. The vas deferens
is always an internal tube, never an external groove. The teeth, when used,
are discharged from the body and not retained in a sac. Among the common,
but not universal, characteristics of the Nudibranchiata are also the
following :—(1) The dorsal tentacles or rhinophores are often laminated and
retractile, features not recorded in any other group. (2) The kidney is
rarely compact, but usually a system of ramified tubes. (3) The genitalia are
often extremely complicated, both in their essential plan and also owing to the
presence of accessory glands and armatures. Besides this, the various sub-
divisions show remarkable peculiarities of their own, such as the ramification
of the digestive organs, the reduction of the teeth to a single row, and the
presence of nematocysts.

The classification of the Nudibranchiata here adopted is essentially that of
Bliot (40) with the following four main differences :—(1) The Ascoglossa have
been removed and treated separately for reasons noted in the introduction,
Wliot’s family-names Doridide Cryptobranchiate and Doridide Phanero-
branchiatz, being binomial, cannot strictly be used, as the family must have a
single name derived from that of its oldest and typical genus. Moreover, both
these terms include a number of groups of forms, which, for the present at any
rate, would appear to be more conveniently handled separately. It must be
confessed that the classification of both these assemblages stands in need of
further revision, a task outside the scope of the present work. (2) For the
reasons just given the group Doridide Cryptobranchiate has been sub-divided
into smaller families, whose limits correspond with the similarly named sub-
families as given by Bergh in his ‘System der Nudibranchiaten Gastero-
poden* in 1892. In certain cases the laws of priority have necessitated a
change of name. (3) Hliot’s four divisions of the Doridide Phanerobranchiate
have been regarded as independent families. (4) Investigation of the synonyms
of various species has revealed the fact that certain names are incorrectly
applied, as was pointed out previously. These have been rectified by the
alteration of the names in the cases that appear to admit of no doubt, and such
a change sometimes involves both the names of the genusand the family, even
though the limits of these groups have not been altered.

The sub-order is divided into two Tribes (A. Holohepatica and B. Clado-
hepatica), a separation dependent in the main, as the terms imply, upon

542 PROF. C. H. O'DONOGHUE: REPORT ON

whether the liver remains in one compact mass or is split up and branched.
The two groups defined in this way, however, are separated by other
associated anatomical differences, that indicate clearly a wider gap between
them than is perhaps suggested by the mere condition of the liver.

Tribe A. Holohepatica.

Nudibranchs in which the liver forms a compact mass, neither branched
nor divided, and usually accompanied hy the following characters. There is
complete external symmetry ; the vent usually hes in the mid-dorsal line and
is surrounded by a circle of branchial plumes; the radula is of moderate or
considerable width, save in the family Dendrodoridee (Doriopsidee) ; mandibles
are rare ; the hermaphrodite gland is usually spread over the liver as a layer,
but sometimes forms a discrete mass; as a rule, the genital ducts are
triaulic and there are two receptacula seminis.

Family DUVAUCELIID (Trironimps).
The branchiz consist of tufts set along the mantle-region or occasionally
they may be absent altogether.

Genus SpoHmrostoma Macgill. Hist. Moll. Anim. Aberdeen, ete. (1843).
Type by monotypy : S. jameson Macgill. = 7. hombergii Cuvier, 1803.
Type: S. hombergit (Cuvier).
Synonymy: Trrironta Cuvier, Tabl. Elém. Hist. Nat. 1798, p. 887, auct.

Necromantss Gistel, Natur. des Tierr. f. Schulen, 1848, p. xi.
Liri0pek Gistel, op 90 ii 1845, p. 171.

The body is limaciform, but somewhat rectangular in outline, except at the
end, where it tapers to a short tail. The foot is broad. Over the mouth is an
oral veil bearing two grooved tentacles at the ends, and in the middle
tubercles or processes. ‘The rhinophores are retractile into raised sheaths ;
they are not perfoliate, but surrounded by a few plumes. The dorsal margin is
slightly prominent, and bears a single row of branchial tufts which are more
or less arborescent. Anal and renal openings lie on the right side. The jaws
are large, with several rows of denticles or prominences near the edge. The
radula varies greatly in size in the different species, but always consists of a
broadish central tooth with a moderate or large number of laterals which are
usually simply hamate. The first lateral is usually larger than the others,
and somewhat clumsily shaped. The liver is not divided and sends off no
branches to the gills. There is no armature either in the stomach or on the
genitalia. The hermaphredite gland forms a layer over the liver (Zliot).

It will be noted that the name of the family is changed from Tritoniidee
Cuvier, as this name is preoccupied, and Duvauceliide substituted. Vritonia
Cuvier being preoccupied has to be replaced by Spherostoma Macgillivray,

2

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 543

43. “‘Dunanueelia Risso, 1826, has to replace Candiella Gray, 1853, as the

valid name for this genus, and thus the oldest generic name in the family
becomes Duvaucelia, from which the family-name is derived. The matter has
been fully dealt with by Iredale and O’ Donoghue (51a, p. 229), and is again
referred to under the Family Euphuride (Polyceride).

Species SPHHROSTOMA DAKINI, sp. noy. (Pl. 27. fig. 5 ; Pl. 29. figs. 35-40.)

Body. The body is oblong, limaciform, and like a typical Spherostoma. It
is moderately high, the back flat, and there is a slight pallial extension along
the edges of the back in the line of the branchial plumes. The dorsum passes
forward and out as a well-marked, strongly bilobed, oral veil, which bears
about six or seven dendriform processes on each side of the middle line. The
short tentacle at each lateral margin of the veil is grooved throughout its
length upon the ventral side.

Colour. In the preserved condition some of the specimens were of a dull
erey colour, and some of the same general colour but plentifully marbled
with dark brown. Professor Dakin informs me that the living animal is of
a dirty pinkish colour with brown markings.

Dimensions. The specimens yaried somewhat in size, and the largest
measured 88 mm. long by 25 mm. wide and 25 mm. high. The lateral
margins of the oral veil project about 15 mm.

Head. The head is sub-globose, distinct, but not projecting, and the mouth
is in the form of a longitudinal cleft. It is completely overhung by the
bilobed oral veil.

Foot. The foot is of moderate width and of the same length as the body.
At the front end it is bilabiate, and posteriorly it passes back to a blunt point.

Rhinophores. The rhinophores are retractile within short, stout, cylindrical
sheaths. This sheath is provided with a funnel-shaped collar with a wavy
edge and a deep cleft on its antero-lateral side. The clavus bears a ring of
six simply pinnate plumes, the most anterior is quite small, the most
posterior is much larger than the others and is continued upwards asa bluntly
rounded projecting cylinder.

Branchiw. The branchiz are in the form of a series of dendritic tufts along
the side of the dcrsum. They are paired and decrease in size as they pass
backwards; the anterior plumes are attached by a wide long stalk and quite
distinct from one another, but at the posterior end the stalks are shorter and
closer together, so that they form a more or less continuous group. About
ten large distinct plumes are present on each side, and a terminal group
composed of from 3-5 smaller branchize. They are of the same colour as the
body of the animal.

Radula. As in all the Duvauceliide the pharyngeal complex is very large
and the raduia very well developed. The total number of rows in the
radula is 65-67 and the number of teeth in a row—in the first row 9 (2. e.,

544 PROF. C. H. O'DONOGHUE: REPORT ON

3.1.1.1.3); in the tenth row 81 (@. ¢.,39.1.1.1.39), and in the sixteenth
row about 273 (7. e., cirea 135.1.1.1.135). When removed and flattened on
the slide the radula measured 11 mm. along the middle line by 12 mm. at
its widest point. The basal plate of the rachidial tooth is approximately
rectangular, and the blade consists of three large spines, of which the
median is the largest, the most pointed, and it projects well in front of the
anterior margin of the basal plate. The lateral spines are blunt and much
shorter. The first pleural tooth has a roughly rectangular base, but it is
very irregular and it bears a single blunt and irregular spine. The remaining
pleural teeth are all of a simple hamate form: they increase in size as they
pass outwards, reaching their maximum between the 90th—-120th, and then
they decrease rapidly to the margin.

Jaw. The jaw, as is usual in the family, is very large and strong. Its
most prominent feature is the series of serrations that mark its cutting-edge.
They are quite small at the upper anterior end near the erista connectiva,
where they first appear as the continuation of a wavy raised ridge, but
they increase noticeably in size as they pass backwards. On the processus
masticatorius they are quite large, nearly 1mm. high and easily discernible
with the naked eye. The denticles are deep brown in colour, and appear to
be inserted into a lighter-coloured ridge. The well-developed jaws measure
18 mm. long by 45 mm. at their widest point, and are of a deep horn-brown
passing off almost to black near the cutting-edge. They are somewhat
narrower than in other species of the genus, and perhaps more widely
divergent at their posterior ends.

The genital aperture is situated about halfway up on the right side near
the level of the second branchial plume. The penis is short, conical, and
unarmed.

The anal papilla lies just behind the third branchial plume, high up on
the right side of the body.

Notes. There are five specimens of this species in the collection: three
with brown mottlings and two quite grey—none with closer locality given
than Abrolhos.

However, it is almost certainly the animal referred to by Professor Dakin
in his report (82, p. 170) as coming from Wooded Island*, since his remarks
can apply to no other form in the collection, and the long branchiz do give
it a spurious resemblance to Dendronotus. He says, “‘ Another animal which
was extraordinarily abundant on this little stretch of Lagoon flat (only on
the margin) was a large and beautiful nudibranch, almost certainly new
and allied probably to Dendronotus. The singularity of its occurrence is
accentuated by the fact that, notwithstanding its abundance here, not a
specimen was captured anywhere else at the Abrolhos, and on our second
visit in 1915 it was just as common at this place as two years before.”

* [This observation is quite correct,—W. J. D.]

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 545

It does not appear to agree with any previously recorded species, and so
is described here as new with the name Sphwrostoma dakeni, after my friend
Professor W. J. Dakin, who collected it.

Family HEXABRANCHID 2.

This family contains only one genus, and its characters are those of the

genus.
Genus Hrxasrancuus Ehrenberg, Symb. Phys. (1828).
Type by monotypy : /Z. pretextus Khrenb., pl. 1. figs. A-C.
N.B.—The plates in this work were published in 1828 and the text
in 1831. This form appears on the plates.

The body is large and of an elongate-oval shape: the back is smooth
and broad. It passes out laterally to a folded margin, which is continued
out as the enormous sheet-like much folded mantle, which may be wider
than the body. The branchial plumes, generally 6-8, are dendritic and
distinct, and arranged ina cirele around the anus. They are not retractile,
but contractile, and their bases are surrounded by a shallow wide-open
depression. The rhinophores have a cylindrical stalk and a perfoliate clavus
set at an angle to it. The oral tentacles are large and auriculate with a
folded margin. The foot is relatively well developed. The labial armature
is well developed and composed of tiny hooked spines. The radula is well
developed, with no rachidial tooth, and the pleural teeth are numerous,
hamate, and have no lateral denticles. The penis is long and unarmed.
Only the posterior blood-gland is present. Bergh classes these as a
sub-family of the Dorididse Cryptobranchiate. This name cannot stand,
and the forms are not even Cryptobranchiate. They are here raised to a
separate family.

Species HEXABRANCHUS IMFERIALIS Kent, Nat. in Austr. 1897, p. 150.
(Pl. 28. fig. 11; Pl. 29. figs. 41-44.)

Body. The body is moderately stout, of an elongated oval shape, and the
skin is smooth. The most striking feature, however, is the mantle, which is
extraordinarily well developed and takes the form of a thin wide flange,
wider than the body, passing all round save at the anterior end, where it is
only narrow. The rhinophores with their sheaths stand up prominently at
the anterior end, and the circle of gills is conspicuous at the hinder end.

Colour. The colour of the preserved specimens is a dull greenish grey
without any indications of markings. Professor Dakin says it is identical
with Saville Kent’s drawing, and that ata distance it looked not unlike a
great piece of fresh lung torn from a vertebrate. Kent figures it as of
a flaring red colour all over with no markings.

Dimensions. The body of the largest specimen measured 125 mm. long
by 50 mm. wide and44 mm. high. Beyond this the mantle extended 80 mm.
along each side, 45mm. across the posterior end, znd 6mm. across the
anterior end as a sort of oral veil.

LINN. JOURN.—ZOOLOGY, VOL. XXXY. 39

546 PROF. CG. H. O'DONOGHUE: REPORT ON

Head. The head is fairly large and sub-globose; the mouth is large and
conspicuous, having the form of a longitudinal sht. On each side of it is an
oral tentacle in the form of a large somewhat ear-shaped flap with a wavy
margin. This measured 27 by 24 mm.

Foot. The foot is well developed, strong, and muscular. Its front end is
abruptly rounded and bilabiate, and the hinder end passes back to a blunt
point. The sides of the foot are thrown out into a wide, folded, undulating
flange which, in the large preserved specimen, projected 16-17 mm.

Rhinophores. The rhinophores are well developed, and the sub-conical
perfoliate clavus with 55-60 leaves is set at an angle of about 45° to the
cylindrical stalk. The stalk is surrounded by a tubular sheath with a
smooth margin, and in some specimens the clavus is retractile within this.

Branchiv, The six branchial plumes are dendritic, very conspicuous, and
arranged in a wide circle. Hach of the anterior two pairs gives off a small
branch from the base, but the posterior pair each arise by two equally
developed main stems coming from the same point.

Radula. The radula is well developed and strong, and when removed
and flattened measured 10°5 mm. along the middle line and 10 mm. at
its widest point. The total number of rows in the radula is 47-49, and
the number of teeth is in the first row 40 (2.e., 20. 0.20), in the tenth
row 130 (i.e., 65.0.65), and in the forty-fifth row 152 (i.e. 76.0. 76).
There is no rachidial tooth. The inner pleurals consist of a roughly
rhomboidal basal plate with a smail curved spine at the antero-median
corner. As they pass outwards they increase in size, the base gets longer,
and the spine increases very markedly. They reach their maximum just
beyond the middle of the row, and then decrease in size mainly in the
basal plate. Thus it comes that, while the inner pleurals are mostly basal
plate with a small spine, the outermost teeth are mainly spine with a small
basal plate.

Labial Armature. The lips are guarded by a thin tough membrane which
is beset with a large number of tiny hooked spines.

The anus is not conspicuous and lies within the circle of gills towards
the hinder end.

The genital aperture is well marked, and situated up on the right side of
the body behind the level of the front end of the foot. The penis is long
and unarmed.

Notes. Five specimens were in the collection :—

A large one and a moderately large one with no locality-label.

Two smaller ones from Pelsart Island.

One of fair size from Wallaby Island.

These specimens were obtained from the same place as Kent obtained his
HT, imperialis and are thus topotypic with it. While he gave a coloured
drawing of the living form, with which the present specimen agrees closely,

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 547
he gave no anatomical details, so that the above is the first description of the
radula ete. of the species.

Family ARCHIDORIDID/.

The body is not hard and slightly depressed ; the noteeum is tuberculate
or granulate, the pallial margin is fairly broad ; the tentacles are small ; the
branchial plumes are nearly always tri- or quadripinnate ; the foot is fairly
broad. There is no labial armature. The radula has no rachidial tooth ; the
pleurals are numerous and hamate. ‘The penis is generally unarmed.

Genus ALLoroporis Bergh, 1904.

Type by monotypy: A. marmorata Bergh, Malak. Unters. vi.,
; Semp. Reis. 1904, p. 42.

The body is depressed, the notzeeum minutely granulose and fairly broad.
The branchiz are not numerous and generally tri- or quadripinnate.
Labial armature is feeble or absent. The foot is strong and as wide as
the body. The mantle is fleshy and well developed. There is no rachidial
tooth. ‘The lateral teeth are numerous and hamate. The last part of the
vas deferens in the penis bears a series of hooks. The hermaphrodite gland
forms a discrete mass quite distinct from the liver.

This genus was founded by Bergh on some specimens from Tasmania, and
is remarkable in that the hermaphrodite gland forms a diserete mass and is ©
not spread out as a layer over the liver. This is a condition encountered
also in Bathydoris and Trevelyana, forms that for other characters are quite
widely removed from the present genus. Hliot suggests (38, p. 333) that
this perhaps represents an older condition than that in which the gland is
spread over the liver, “and it would seem that very different families of the
Dorididae sporadically preserve or revert to the older arrangement.”

Up to the present only two species have been referred to the genus,
viz., A. marmorata Bergh and A. lanuginata Abr.

Species ALLOIODORIS HEDLEYI, sp. nov. (Pl. 2%. figs. 6 & 7; Pl. 80.
figs. 45-47.)
Synonymy: A. marmorata Basedow & Hedley, Trans. Roy. Soc. South Austr. 1905, p. 152,
non Bergh,

Body. The general shape of the body is elliptical, slightly broader at the
posterior end, and only moderately convex. The entire dorsal surface is
covered with minute spiculate papille, making it somewhat rough to the
touch. The mantle is very well developed, being much wider than the foot,
and its fairly thin edge is wavy and undulating.

Colour. The colour of the preserved specimen is a dark muddy-fawn with
a tinge of green. Irreeularly scattered over it are a series of dark brown,
almost black, roughly circular, ring-shaped marks from 2-3 mm. wide,

a)

548 PROF. C. H. 0’ DONOGHUE : REPORT ON

A similar series of markings is present on the under surface of the mantle.
The foot is slightly lighter in colour and speckled with dark brownish-black
spots.

Dimensions. The largest specimen measured 52 mm. long by 34 mm. wide
and 23:5 mm. high. The foot measured 42 mm. long by 16°5 mm. wide.
Thus it is larger ‘hewn Bergh’s specimen of A. marmorata (45 by 25 by 13 mm.),
and considerably larger than the measurements" given by Basedow and Hedley
(22:5 by 10 mm.).

Head. The inconspicuous head is very small and tucked in between the
front margin of the foot and the large overhanging mantle. The mouth
appears as a tiny longitudinal slit, and the oral tentacles are sub-cylindrical
and when contracted measure 1:75 mm.

Foot. The foot is elongated and narrow ; the front end deeply bilabiate
and rounded ; the hinder end bluntly pointed and in the specimen not quite
reaching to the edge of the mantle.

Rhinophores. The rhinophores are club-shaped, and the perfoliate clavus
occupies almost three-quarters of their length. They are completely
retractile within cavities with a circular, slightly raised aperture.

Branchic. The branchiz were six in number. Basedow and Hedley (8)
state that they were seven or eight, but show only six in their figure. They
are retractile within a cavity which opens by a raised, transversely oval
aperture with a wavy margin.

The anal papilla lies within the circle of gills ; it is lowand bears the renal
aperture on its right anterior base.

Radula. The vals is well developed, and when removed and flattened
measured 6 mm. along the middle line and 5-5 mm. at its widest point. The
total number of rows in the radula was 383-35 and the rows contained :—the
first row 6.0. 6 (2. e., 12) teeth ; the tenth row 55.0. 55 (7. e., 110) teeth ; and
the thirtieth row 57. 0.57 (2. e., 114) teeth. The innermost pleural teeth are
small, and consist of a curved spine on an elongated basal plate. They
increase very markedly in size to beyond the middle of the row, where they
are falciform, and then they decrease again towards the outside. No sign
of denticles was found on the lateral teeth.

Labial Armature. None was noticed.

The glans penis is armed with a series of rows of hooks.

The hermaphrodite gland is discrete and does not form a layer over the
liver.

Notes. The collection included three specimens. One small one from
Pelsart Island and the other two larger ones with no definite locality-label.

The specimens described by Basedow and Hedley (8, pp. 139 and 152) are
from St. Vineent’s Gulf and Port River, South Australia, and from Wdith-
burgh, York Peninsula. It has been noted that the specimens here measured
are considerably larger than those measured by Basedow and Hedley.

OPISTHOBRANGHIATA FROM THE ABROLHOS ISLANDS. 549

Mr. E. Ashby, who is familiar with this form on the coast of South Australia,
informs me that they are usually of the smaller size. It has not been
recorded previously from West Australia.

Some confusion exists in regard to the nomenclature of the members of
this genus, which deserves comment.

The following forms have been recorded :—

A. (Doris) lanuginata Abraham, Proc. Zool. Soc. Lond. p. 255,
pl. xxix. figs. 15-17 (1877).

A. marmorata Bergh, Malak. Unters. vi., Semper’s Reisen, 1904, p. 42.

A. marmorata Bergh (sic), Basedow & Hedley, Trans. Roy. Soc. South
Australia, vol. xxix. p. 152 (1905).

A. lanuginata Abr., Eliot, Proc. Malac. Soc. Lond. p. 333 (1907).

A. hedleyi mihi, described above.

The original specimen described by Abraham is in the British Museum ;
it is not well enough preserved for detailed examination and the pharyngeal
complex is removed. Hliot (98, p. 333) states that, on comparing this
specimen with that sent by Suter from New Zealand, he has no doubt that
they are identical. He gives the genus as containing two species (A.
lanuginata Abr. and A. marmorata Bergh), but lower down on the same page
he gives A. marmorata as a synonym of A. lanuginata, and on p. 335 he states
that Bergh’s A. marmorata is perhaps a distinct species. Further, he
suggests that it is not clear that the animal figured by Basedow and Hedley is ~
either of the two species. On comparing the descriptions we find :—

A, LANUGINATA (Abr.).—Colour preserved ‘ greenish-grey, blackish
mottlings” (lot). In the specimens in the British Museum the
mottlings are uniformly dark areas (O’/).). Colour in life “ dirty red,
numerous white pustules” (Su¢er), but the applicability of this to the
specimen is doubtful.

A weak labial armature. Tentacles long, flat, furrowed on upper
surface, and almost auriculate. Rhinophores much closer together
in large specimens of A. hedleyi than in the far smaller original
specimen of A. lanuginata. Size 50 mm. long, 33 mm. broad,
16mm.high. Radula 26 rows, (40-45) .0. (40-45). Ist and 2nd teeth

characteristic, unlike A. marmorata or hedleyt.

A. MARMORATA Bergh.—Colour preserved “von hell gelblichweisser Farbe,
die von zahlreichen kleineren und (bis 3 mm. diam.) grosseren hell
aschegrau, verschwimminden, unregmassigen Flecken unterbrocken
war” (no mention of spots being ring-shaped, which would have
hardly escaped Bergh’s acute eye).

Labial armature not mentioned. Tentacles 1:5 mm long, “ finger-
formig.” Size 45 mm. long, 25 mm. broad, 13 mm. high. Radula
34-35 rows, (40-42) .0. (40-42). 1st and 2nd teeth like remainder,
outer teeth denticulate and not quite same shape as 4. hedleyi.

550 PROF. C. H. O’DONOGHUE: REPORT ON

ALLOIODORIS HEDLEY! mihi.—Colour preserved; dark muddy-fawn with a
tinge of green with irregularly scattered dark brown, almost black,
roughly circular, ring-shaped marks (O’).). Colour alive: yellowish
white to greyish brown, covered with minute spiculose elevations on
the dorsal surface, which impart to it a brownish tint; also less
numerous, larger elevations, surrounded by irregular circles of deep
brown. The latter occasionally have a centre of opaque white
surrounded by a ring of reddish brown.

No labial armature. Tentacles small, sub-cylindrical, 1°25 mm. long.
Size 02 mm. long, 34 mm. wide, 23°5 mm. high. Radula 33-35 rows
(55-57) .0. (55-57). Innermost pleurals small, but much like the
remainder, not like those of A. lanuginata ; outer teeth not denticulate.

I think it will be seen from the above that three distinct forms appear
to be represented, of which A. marmorata Bergh and A. hedleyi mihi,
while resembling one another more closely than either of them does A. lanu-
ginata, are, nevertheless, distinct species. The form described by Basedow
and Hedley is not the same as that recorded by Bergh, and hence it is in
need of anew name. I propose to call it A. hedleyi after Dr. Charles Hedley,
who has added so much to our knowledge of the marine fauna of Australia.

Family PLATYDORIDID A.

The body is somewhat flattened, of leathery consistency, sometimes tough
and sometimes brittle ; it has an oval or sometimes almost circular outline ;
the notzeum is fairly smooth or minutely granulate, the dorsum often marked
by tubercles or ridges ; the pallial margin is ample ; the branchial aperture
is generally stellate: the tentacles are digitate ; the anterior margin of the
foot is bilabiate, and the upper lip indented or split in the middle line.

There is no labial armature; the radula lacks rachidial teeth ; the pleural
teeth are numerous and hamate.

The prostate gland is large.

Genus AstERonotus Ehrenberg, 1831.
Type by monotypy: A. hemprichii Ehrenb. Symb. Phys. (1831).

The body-form is oval and somewhat flattened ; the animal is of a peculiar
leathery or india-rubber-like consistency, not hard or brittle. The skin is
smooth to the touch, but covered with raised protuberances in the form of
ridges, of which one well-marked one runs in the mid-dorsal line from
between the rhinophores back to the branchial aperture. The margin of
this aperture is produced into lobes (usually six). The foot is narrow,
bilabiate anteriorly, and the upper lip indented.

There is no labial armature; the radula has no median tooth and the
pleural teeth are simply hamate.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 551

The penis is unarmed, the prostate large, and, according to Bergh, a
glandula and hasta amatoria are present.

Species ASTERONOTUS FUSCcUS, sp. nov. (Pl. 28. figs. 12,13; Pl. 80.
fis. 48-50.)

Body. The body is shaped like a typical Dorid—oval and with a moderately
arched dorsum. The manile is well developed, and projects considerably
beyond the foot all round. The whole animal is like a piece of india-rubber
to feel. The dorsum bears a series of raised elongated tubercles, the largest
of which forms an irregular ridge stretching from between the rhinophores
back to the branchial aperture. From this median projection a number of
unequal and irregular short side-ridges are given off, getting lower as they
pass from the middle line, and all around the margin is a number of small
oval or roundish tubercles.

Colour. The colour of the preserved specimen is a muddy-brown, and
a note included with it states that it was ‘“‘dirty brown, rough surface.”
Professor Dakin informs me that it was of a dirty brown with some lighter
brown spots, but no conspicuous colour-pattern.

Dimensions. The specimen measured 27°5 mm. long by 15 mm. wide and
9 mm. high, and the foot 24 mm. by 6°5 mm.

Head. The head is quite small with a small mouth in the form of a short
transverse slit, and on each side is 1 small tentacle.

Foot. The foot is only moderately broad, with an undulating margin.
The anterior end is bluntly rounded, strongly bilabiate, the upper lip is
indented in the middle line, and the posterior end is pointed.

Rhinophores. The rhinophore has a bluntly conical, perfoliate clavus with
a fair number of thin leaves and a short cylindrical stalk. They are com-
pletely retractile within cavities, whose margins are slightly raised, so that
they look somewhat like a tubercle. In the preserved specimen they are of
a yellow colour.

Branchie. The branchie are five in number, each consisting of a large,
branched, plume-like gill, arranged in a circle. They are completely
retractile within a cavity with a raised margin, so that it stands up like one
of the larger tubercles. The edge of this is furnished with six small lobes,
which, when the aperture is tightly closed, are tucked inside.

Radula. The pale yellow radula when flattened on a slide measured 3 by
2:75 mm. and contained 38-40 rows of teeth. A median tooth is absent, and
the number of pleurals in each row was :—in the first 6.0.6 (2. e., 12) and in
other rows from the third on 65.0.65 (i.e., 130). The teeth themselves
are simply hamate and increase in size as they pass outward to beyond the
middle of the line. ‘They decrease in size rapidly at the outer margin,
and the last one or two are reduced.

The anus lies on a well-marked papilla in the middle of the branchiz, and
it bears at its base the small oval excretory pore.

IU

Ot
to

PROF. C. H. 0’DONOGHUE: REPORT ON

The genital aperture, as usual, lies on the right side of the body near the
anterior end.

Labial Armature. No labial armature was found.

Notes. One specimen only is in the collection, and this was accom anes
by the locality-label “ Shore of Wooded Isle.”

Eliot (36, pp. 305-356) calls attention to the difficulties encountered when
trying to determine the position of a Cryptobranchiate Dorid, and in dealing
with the present form they have come before me frequently. Bergh’s family
Dorididee Cryptobranchiatee is divided into five sub-families and about 30
genera, practically all of which were created by Bergh himself, and certain
of them contain only a single species. The differences between some of the
genera are so slight that, unless Bergh’s own species are dealt with, it is
difficult to determine where to place a new form. There seems no doubt
that the whole family stands in need of considerable revision before it can be
regarded as satisfactory. The preseni specimen agrees closely with the
genus Asteronotus, and in general form, and perhaps even colour, somewhat
resembles the A. hempricht Ehr. as figured by Eliot (36, pl. xxxiv.), but it
differs in certain points. The radula formula is different, the individual
teeth are not quite same, the branchise number five, etc. For these reasons
I have described it as a new species under the name of Asteronotus
fuscus.

Family GLOSSODORIDID (Curomoporipin» Bergh).

Tt has been pointed out above that Bergh’s binomial family-name Doridide
Cryptobranchiate cannot possibly be aillonsad tostand. It is proposed, there-
fore, to split this group into several families, of which oue is the same as the
old Chromodoridine of Bergh. But, as the name Chromodoris has to be set
uside for the older name (rlossodoris, the family-name must be changed as
above.

The body is elongated, compressed, and soft; the colour is striking, often
magnificent, and generally with lines or spots ; the noteeum is nearly always
smooth, the pallial margin is fairly broad at the anterior and more particu-
larly at the posterior end, but usually quite narrow in between ; the tentacles
are small, conical, and generally partly eversible; the branchial plumes are
usually simply pinnate,

The labial armature is strong and composed of minute rods. The rachis
of the radula is very narrow and often furnished with minute compressed
psceudo-teeth ; there are numerous pleural teeth of hamate form and generally
with a denticulate margin, the first teeth are usually denticulate on both
sides.

There is no proper ventricle.

The penis is unarmed.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 553

Genus Guossoporis Hhrenberg, 1831, Symb. Phys. [unpaged,
but on p. 92 |.
Type by subsequent designation: G. wantholeuca Hhrenb. Red Sea.
Gray, Proe. Zool. Soc. Lond. 1847, p. 164.
But G. xantholeuca Hhrenb.== Doris pallida Riipp. et Leuck., 1828.
The type is therefore here designated (. pallida (Riipp. et Leuck.), 1828.
Synonymy: Actrnoports Ehrenberg, 1881.
Type by monotypy: A. sponsa, Khrenb. Symb. Phys. [p. 93]; wide also
Gray, Proc. Zool. Soc. Lond. 1847, p. 164.
Prrroporis Khrenberg, 1831.
Type by subsequent designation: P. picta Ehrenb., Gray, Proc. Zool, Soc.
Lond. 1847, p. 164.
Curomoporis Alder & Hancock, 1855.
Type by original designation and monotypy : D. magnifica Quoy et Gaim.,
Alder & Hancock, Mon. Brit. Nud. Moll. pt. vil. 1855, p. xvii.
GONIOBRANCHUS Pease, 1866.
Type by designation: G. vibrata Pease, Pease, Amer. Jour. Conch. ii.
1866, p. 204.

Chromodoris, the generally accepted name for this genus, was introduced
by Alder and Hancock (wide supra). These authors in a subsequent paper
(6, p. 115) say :—“ The Goniodoris of Forbes has hitherto been considered a
northern form—the southern species which some authors have referred to it
belonging almost without exception to the allied genus Chromodoris, which, -
on the other hand, has not been found further north than the Mediterranean.”
The name thus introduced was used by Bergh (11, p. 72), who gives as
synonyms Doriprismatica d’Orbigny and Goniobranchus Pease, and again
in 18 (p. 1), but this time with the synonyms Glossodoris, Actinodoris, and
Pterodoris, all of Hhrenberg, 1831, twenty-four years earlier than Alder and
Hancock’s name. Bergh points out there and again in 1884 (28, p. 65) the
identity of Ehrenberg’s genera with that of Alder and Hancock, and in this
he is right.

Thus, in spite of the common usage of the generic designation Chromodoris,
there is no doubt that Ehrenberg’s names have considerable priority. The
question as to which name should be employed is easily settled, for, while
they were all published at the same time, Glossodoris comes first in order,
and the first species is given as G. xantholeuca, which Gray (44, p. 164)
designated as the type-species. Bergh, in a paper where he re-examines
Khrenberg’s types, states, in our opinion rightly, that G. vantholeuca is the
D. pallida (Ritpp. et Leuck.), and that all species of Glossodoris are con-
generic. The genus then stands as Glossodoris with the type-species
G. pallida (Riipp. et Leuck.).

In various places Bergh includes Voriprismatica d’Orbigny, 1837 (11), as
a synonym; but, while part of d’Orbigny’s genus belongs to the present
group, the type, D. atromarginata Cuv., 1804, is in reality a member of a

554 PROF. C. H. 0 DONOGHUE: REPORT ON

genus of its own, often but incorrectly termed Cassella H. & A. Adams,
1858 (8).

The Gontodoris of Forbes, 1840 (42), still remains as a valid genus, and so
does not enter the synonymy, although certain forms originally included in
it really belong to Glossodoris.

The body is elongated and rectangular or almost square in transverse
section. The smooth dorsum is an elongated oval, rounded in front and
behind, and with its long sides approximately parallel. The mantle projects
slightly all round and usually forms a frontal and caudal veil. The foot is
large and continued backwards as a fairly long pointed tail. The sides of
the body are vertical. The oral tentacles are small and conical; the
retractile rhinophores have a perfoliated clavus. The retractile branchice
are composed of simply pinnate leaves. The armature of the labial disc is
strong and composed of a number of densely-set small hooks bifid at the tip.
The radula contains no rachidial teeth, but there are frequently thickenings
which take their place. ‘The lateral teeth are numerous and hook-shaped ;
the first lateral tooth is denticulate on both sides, the rest denticulate only
upon the external margin ; the outward teeth are smaller, and denticulate
at the extremity. The penis is unarmed.

Glossodoris resembles fairly closely in general appearance and shape
Groniodoris, but the latter is phanerobranchiate. It is also much like Aphelo-
doris, but differs in the presence of labial armature, of denticulate teeth, and
simply pinnate branchie.

Species GLOSSODORIS WESTRALIENSIS, sp. nov. (PI. 27. figs. 8,9; Pi. 30.
figs. 51-53.) :

Body. A coloured sketch by Professor Dakin from the living animal
shows that its general shape was that of a typical Glossodorid, but the
preserved specimens have shrunken considerably, more particularly in the
tail-region. The noteeum is a fairly narrow, flat oval, equally rounded at each
end and the sides almost parallel. In the preserved specimens it contracts
considerably, becoming less rectangular and more arched. The sides of the
body pass inwards to a moderately narrow foot and the mantle only projects
a very short distance.

Colour. From Professor Dakin’s sketch the general body-colour appears
to bea bright blue. The notzeum is bordered by a narrow band, yellow on the
inside and orange-red right at the margin. The rhinophores are orange-red
and the branchiee red. On the noteeum inside the marginal line is a narrow
band of blue and then a slightly wider band of deep purple, almost black.
This deep band passes round in front and lateral to the rhinophores, and may
or may not enlarge slightly to include the margin of their sheaths. At the
posterior end it passes round more or less parallel with the margin and
behind the branchize. In the middle of the noteeum the dark band enlarges,

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 5955

so that on the outside it obliterates the blue band and on the inside it passes
across to the other side as a broad band, thus isolating, as it were, two islands
of the light blue ground-colour on the noteum. ‘wo similar bands of dark
colour pass back on the dorsal surface of the tail.

In the preserved specimens all trace of the orange, red, and bright blue
colours has disappeared and the whole ground-colour is of a dirty brownish
fawn ; the dark lines, on the otlier hand, are quite clearly visible. In addition
to those already noted on the drawing, a narrow dark band passes around
the sides of the body at the line of junction of the mantle-fold, and near the
anterior end it loops downwards and backwards. It then passes back
parallel with its former course, but lower down the side of the body, as a
fairly broad streak running out on the dorsal surface of the tail as the two
lines already noted. Under the posterior end of the mantle all these dark
lines may unite and pass back right to the end of the tail as a single broad
band. A narrow dark band runs right around the upper margin of the foot
and out to the tip of the tail, where it may or may not unite with the lines
already described.

The striking and characteristic distribution of these dark lines is fairly
constant in all the specimens and shows well after preservation.

Dimensions. The specimens were all about the same size, and measured
when preserved :—noteum 20 mm. long by 10 mm. wide; foot 22 mm.
long ; body 17°5 mm. wide. ;

Head. The head is quite small and overhung by the forward extension of
the mantle. The mouth is a small longitudinal slit, and on each side there
is a small cylindrical tentacle, much contracted in all specimens.

Foot. The foot is linear and its sides almost parallel. It is rounded and
bilabiate in front and passes off to a pointed tail posteriorly. The sides of
the foot are marked off from the body by a marginal flange. Judging from
the sketch of the living animal, the tail may project beyond the noteeum
about half the length of the latter, but in the preserved specimens it is
usually curled up and hardly projects at all.

Rhinophores. The rhinophores have a short stalk and a cylindrico-conical
perfoliate clavus. They are completely retractile within cavities which have
a slightly raised margin.

Branchie. The branchiz are fairly simple and arranged in an almost
complete circle. The anterior ones are simply pinnate thread-like plumes,
sometimes tending to be bifid at the distal extremity; these number nine,
although there is a tendency to fusion at their bases. At the posterior end
on each side is a group of four similar, but smaller, plumes more closely
united at the base, and these when extended may appear as four branches
from one stalk.

The anus lies in the middle of the branchial circle. All can be completely
retracted within a pocket with a circular aperture surrounded by a slightly
raised margin.

or
or
or

PROF. C. H. O?7 DONOGHUE: REPORT ON

Radula. When removed and flattened the radula measured 5 mm. by
3°25 mm. and was large and well developed for the size of the animal. The
total number of rows in the radula was 83-85, and the number of teeth in
each row :—in the first row 20.0. 20 (i.e., 40) ; in the tenth row 72.0.72
(z.e., 144); and in the eightieth row 76.0. 76 (7.e., 152). There is no rachidial
tooth. The innermost teeth are small but stout, and consist of an elongated
base with a marked hook, on the side of which area series of from 4-6 small
triangular denticles. ‘lhe teeth increase in size as they pass outwards and
the hook becomes larger, bearing on its inner curved edge a series of 6-8
denticles and terminating in a rounded point. The outermost teeth are
slightly smaller than those nearer the middle of the row. ‘They have their
bases much smaller, and the denticles are reduced to one or two blunt pro-
jections near the end.

The genital aperture is placed high up on the anterior end of the right
side of the body.

Notes. In all there were six specimens in the collection: three were from
the Abrolhos Islands with no further details; two were from the shore of
Wooded Isle, one of which had the pharynx exserted ; one was from the
shore of Long Island, and this was very dark in colour.

It does not appear to correspond with any previously described form.

Genus APHELODORIS Bergh, Malakozool. Blatt. 1879, p. 107.

Type by monotypy: A. antillensis Bergh, ibid. p. 108.

In outward appearance these forms very closely resemble the Glossodorids,
and have a well-developed noteeum beyond which the tail projects. The
mantle-edge only projects a very short way. The foot is fairly small, rounded
in front, and not very sharply marked off fr.m the body. The rhinophores as
in Glossodoris. The oral tentacles are short and stunted, and have a cleft on
their underside. The retractile gills are composed of a few (5) tripinnate
plumes. The lips are only covered with a moderately thick cuticula. There
is no rachidial tooth, and the pleural teeth are numerous and hamate. Only
a posterior blood-gland is present. The prostate is large and the penis
unarmed.

Species APHELODORIS AFFINIS Eliot, Proc, Malac. Soc. Lond. 1907, p. 343.
(Pl. 27. fig. 10; PJ. 80. figs. 54-56.)

Body. The shape of the body is typical of the Glossodorids. The noteeum
is a moderately long oval with its long sides approximately parallel and the
ends equally rounded. It is carried over all round into a narrow paliial
ridge. The body is fairly stout and its sides almost upright, running down
to a moderately wide foot. The lower part of the body extends out beyond
the noteeum as a tail.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS, 557

Colour. The body-colour in the preserved state is of a pale yellowish grey,
in two specimens with a deeper yellowish-brown narrow line around the edge
of the notzeum and the foot. In places this line bears still darker dots. In
a third specimen the anterior end of the sides of the body, the noteeum in
front of the rhinophores and behind the branchial aperture, and the dorsal
region of the tail are of an umber-brown coyered with dark spots. The
largest specimen has this coloration still more strongly developed, and the
whole of the notzeum, body, and foot is of the same umber-brown covered with
dark irregular spots, and the anterior region of the notzeum is of a very dark
brown, almost black. Professor Dakin informs me that the living animals
were similarly coloured, but the brown somewhat warmer.

Dimensions The largest (and darkest) specimen measured 24 mm. long by
13 mm. wide and 12 mm. high, but this issomewhat distorted. Three others
were more or less of a size, and measured 20 mm. long by 9°5 mm. wide and
9 mm. high.

Head. The head is small and ineenspicuous ; it bears a short sub-cylindrical
tentacle on each side, between which the mouth appears as a small circular
aperture. Hach tentacle bears a longitudinal groove on its under surface.

Foot. The foot is moderately broad and its sides fairly parallel. The
front end is abruptly rounded and the hinder end passes off to a point.

Rhinophores. The rhinophores have a short cylindrical stalk and sub-conical
perfoliate clavus. They are completely retractile within cavities, whose rims
are raised into short dark-coloured tubes. ;

Branchiw. The five small, dark-coloured, tripinnate, branchial plumes are
arranged in almost a complete and very narrow circle. They are retractile
within a cavity with a circular non-raised aperture.

Radula, The radula when removed and flattened measured 3°5 by 2°5 mm.
It is well developed and has no rachidial tooth. The number of rows is from
27-28, and the number of teeth in a row is :—in the first row 6.0.6 (¢.e., 12);
in the tenth row 42.0.42 (¢.¢., 84) ; and in the twentieth row 46.0.46
(¢.e., 92). The inner teeth are small simple hooks, somewhat angular and
without denticles.. They increase markedly in size and become simple
eurved hooks. Towards the outside they decrease noticeably, and they turn
into short hocks only slightly curved.

The anus lies within the cirele of gills at its hinder side.

The genital aperture lies about halfway up on the side near the anterior
end of the body.

Notes. This species was represented by four specimens collected on
“¢ Wooded Island.”

Kliot (88, p. 343) described a form from New Zealand of which he says,
“ Colour dirty white with irregular mottlings of dark reddish brown. The
epidermis peels off very readily, and it is possible that the brown mottlings

may have been much more extensive, or eyen that the dorsal surface may

558 PROF. C. H. O’DONOGHUE: REPORT ON

have been wholly brown.” This agrees closely with my own observations,
and the peeling of the epidermis is also similar. The radula formula is
given as 60.0. 60, and the total number of rows 23 ; this is not quite the
same, but the teeth are apparently identical. He says, further, “oral
tentacles white ; large, flat, and distinctly grooved.” This is not at all in
agreement with the present specimens, and, indeed, is not found in other
members of the genus. Hliot’s form from its teeth and other features
undoubtedly is referable to this genus, and I am inclined to think that this
description of the tentacles is not quite accurate.

In spite of this one serious discrepancy, the present forms agree so closely
with the description of A. affinis given by Hliot, that I think it safer to refer
them to that species than give them a new specific name.

Genus Crrarosoma Adams & Reeve, Voyage of ‘ Samarang,’ 1848, p. 67.
Type by monotypy : C. cornigerium, Adams & Reeve, idem, p. 68.

Species CERATOSOMA BREVICAUDATUM Abraham, Ann. & Mag. Nat. Hist. (4)
xvill, p. 142, pl. 8. fig. 6 (1876). (Pl. 28: fie. 14; Pl. 30:
figs. 57-59.)

Synonym: C, oblongum Abraham, J. c. p. 148, pl. 7. figs. 7, 7a, 70.

Bergh (25) suggests that C. caledonicum Fischer, C. tenue Abraham,
and (. oblongum Abraham are all synonyms of C. brevicaudatum Abraham.
Basedow and Hedley (8, p. 154) call attention to the following :—in the case
of C. caledonicum, Fischer’s description indicates an animal with the lobes of
the noteeum more developed, and the colour-scheme in the two forms is
entirely different. This being so, it seems undesirable to merge the two into
one species without further evidence. The same two authors suggest that the
differences between C. brevicaudatum and C. oblongum are due to the amount
of contraction undergone upon or before preservation, and state that they
have obtained examples of this species exhibiting similar differences from the
same dredging. An examination of the external characters of the original
specimens described by Abraham, and preserved in the British Museum, leads
me to think that they are probably right, and that C. oblongum is to be
regarded as synonym of C. brevicaudatum. I have also examined the original
example of C. tenue described by Abraham, and, while it is possible that it
may also be a synonym, it is somewhat different in appearance, and it is
probably better—for the present, at any rate—to keep it separate, as has been
done by Basedow and Hedley.

The specimens here described agree closely with the original description
given by Abraham and that furnished by Basedow and Hedley, and are
undoubtedly referable to C. brevicaudatum.

Body. The body is large, elongated, slightly wider in the middle than at
the two ends, and it increases in height from before backwards to the middle
or just beyond. ‘The sides are high, almost vertical, and practically parallel.
The back is flattened and continued out into a narrow undulating mantle-

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 559

fold, which is rounded at the anterior end, much wider posteriorly, and
terminates in a median tongue-shaped projection. The dorsum and sides of
the body are quite smooth.

Colour. The colour of the preserved specimen is a uniform dull yellowish
brown. Basedow and Hedley (8, p. 143) give a full account of its coloration
and some excellent figures (8, pl. i.), and Professor Dakin informs me that
the present specimens agree with them. The general colour is shaded pinkish
buff and pinkish brown ; the darker colour is more prevalent towards the
margin. Along the margin of the dorsum are a series of light areas, each of
which has a violet-purple spot surrounded by a ring of reddish purple. The
mid-dorsal region is strewn with scattered circular spots of a light violet-
purple colour, in some cases surrounded by a rim of light yellow, and some
of these lie in front of the rhinophores. The postero-median projection of the
dorsum is covered with a network of brown lines. The sides of the body are
similarly coloured to the dorsum, and have a number of small spots approxi-
mately arranged in three bands :—the upper and lower are of a rich purple
and the median spots of a light violet-purple. The rhinophores and the
branchize are of a rich reddish-orange colour.

Dimensions. The larger specimen, preserved in spirit, measured: 92 mm.
along the foot from the tip of the tail to the front end; breadth of the body at
the widest part 22 mm.; height of the body 26 mm. ; breadth of the dorsum
and mantle at the anterior end 22 mm.; at the posterior end 36 mm.;
length of tail on dorsal side from the end of dorsum to tip 38 mm. Basedow
and Hedley’s measurements are “length 111, breadth 25, height 31 mm.,”
so that, allowing for shrinkage, the present specimen was almost the same
size.

Head. The head is small and inconspicuous, and is completely overhung. by
the anterior end of the mantle. The mouth is small and in the form of a
longitudinal slit running vertically. On each side of the mouth is a short
sub-conical oral tentacle, which can be retracted into a sort of shallow pit.

Foot. The foot is linear, rounded at the anterior end, passes off to a blunt
point posteriorly, and its margin is wavy.

Rhinophores. The club-shaped rhinophores are perfoliate, and the clavus,
which is a little more than half the total length, bears a number of closely
packed leaves. Hach is completely retractile within a cavity, which has the
margin of its aperture slightly elevated.

Branchie. The branchial plumes are twelve in number and incompletely
surround the anal cone. The bases of the plumes show a marked tendency
to fusion, and the posterior five on each side almost appear to arise from one
ridge. Hach plume is somewhat dendritic in arrangement.

They are all retractile together with the anus within a cavity that has the
margin produced into a short tube.

Radula. The radula is strong and when removed and flattened measured
12 mm. by 95mm. The number of rows of teeth is 74-76 and the number

560 PROF. C. H. 0’DONOGHUE: REPORT ON

of teeth in a row:—in the first row 25.0.25 (7.e., 50); in the tenth
row (117-120). 0. (117-120), i.e. (284-240) ; and in the seventeenth row
(130-135). 0. (130-135), 7. e. (260-270). There is no rachidial tooth. The
inner teeth are in the form of an oblong base bearing a short curved spine.
They increase noticeably in size to beyond the middle of the row, where they
are hamate with a bluntly pointed corner in the middle of the back. At the
outer edge they become somewhat more triangular in side-view, and the last
5-6 have a small rounded denticle near the apex of the spine.

The prominent genital aperture lies on the right side about a quarter of
the length of the body from the anterior end and high up beneath the narrow
mantle.

Notes. his species is represented by two specimens. The larger one taken
from near Fremantle and the smaller from the Abrolhos Islands. The
latter is interesting, because it bears the parasitic copepods described later.

The type-specimen in the British Museum described by Abraham (1) has
for its locality simply Australia, it was recorded by the same author from
West Australia as C. oblongum. Basedow and Hedley report it from
Sydney Harbour, New South Wales; St. Vincent Gulf, South Australia,
20 fms.; Antechamber Bay, Kangaroo Islands, 20 fms. (?); Port Noarlunga,
low water, and Salt Creek Bay, Yorke Peninsula *.

Family DENDRODORIDID& (Dortorsip).

The body is nearly always soft and the general shape much as in
Archidorididee. The mouth is in the form of a small pore ; the tentacles are
small and partly adherent ; the rhinophores and branchize are very similar to
Archidoridide. The notzeum is smooth or tuberculate; the pallial margin is
well developed and often with an undulating margin. The foot is broad and
much as in Archidoridide.

The buccal tube is simple and non-glandular. The Bulbus pharyngeus is
suctorial, elongated, cylindrical, and destitute of mandibles or radula. The
end of the liver is deeply notched.

The penis is armed with a series of hooks.

Genus Denproporis Ehrenberg, Symb. Phys. 1831
[unpaged, but on p. 94].
Type by subsequent designation: ). lugubris Ehrenb.; vide Gray,
Proc. Zool. Soe. Lond. 1847, p. 164.
Synonymy: Dorropsis Pease, Proc. Zool. Soc. Lond. 1860, p. 32.
Type by monotypy, D. granulosa Pease.

Racuoports Mérch, Jour. de Conch. 3rd ser. iii. 1863, p. 34.
Type by original designation, D. Jaciniata “ Cuv.,” auct.

* [It was found later by me at Albany, King George Sound, on the South Coast of Western
Australia. It has thus a wide southern range with northern extension.—W. J. Dakin. ]

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 561

Dorwoprsts Alder & Hancock, Trans. Zool. Soc. Lond. vol. v. pt. 3, 1864, p. 125.
Type by original designation, D. gemmacea Ald. & Hancock. ‘

Havstrtnoporis Pease, Amer. Jour. Conch. vol. vi. 1871, p. 299, for the
Doridiopsis of Alder & Hancock.

Doriopsis, the generally accepted name for this genus, was introduced by
Pease, 1860 (62), who failed to give a satisfactory definition of the genus. Of
it Bergh (11, p. 83) says: ‘ Das Geschlecht Doriopsis wurde von Pease in der
gewobnlichen Art des Heeres von Art- und Geschlechte-Pabrikanten mit
ganz flaschen Characteren schon in 1860 aufgestellt.”

Four years later Alder and Hancock (6) more accurately described the same
genus, but gave it the name Doridopsis, without, however, referring to Pease’s
genus or species. Again, the following year Hancock (46) gave a detailed
description of the genus, also without reference to the work of Pease. The
latter author in 1871 re-affirmed his genus, claimed that the genus of Alder
and Hancock was not identical, and suggested for their genus the name
Haustellodoris. There can be no doubt, however, that all the species of both
authors belong to the same genus.

However, Ehrenberg in 1831 had founded the genus Dendrodoris for two
species, one of which (D. lugubris) was subsequently designated the type by
Gray (44, p. 164). Bergh (18, p. 21) points out that Hhrenberg’s definition
of the genus is incomplete and unsatisfactory, although both the species
included within it are recognisable and without doubt belong to the genus
subsequently termed Doriopsis. He further says (/. ¢. p. 22), “ Hs hegt daher
um so mehr wohl kein Grund vor, die Benennung Dendrodoris fiir die durch
Hancock und durch mich jetzt so gut bekannten Doriopsen zu restituiren.”

This is a strange line of reasoning to follow, for, in the first place, Hancock
uses the term Doridopsis, not Doriopsis, and was not the author of either
name. Secondly, while admitting that both Hhrenberg’s original species
belong to the genus under discussion, Bergh prefers to adopt a much later
term, not simply incompletely defined, but actually with “ganz flaschen
Characteren.”

There seems little doubt that the genus should stand Dendrodoris Hhrenb.
with D. lugubris Hhrenb. as the type by subsequent designation, and so the
name of the family also needs to be altered to Dendrodoridide.

It is interesting to note that the same species is described under the same
name, Doris lugubris, by Grayenhorst (43a) in the same year, 1831. I have
been unable to ascertain whether Gravenhorst’s or Ehrenberg’s name has
priority, but, in any case, it does not alter the generic name.

Species DmnpRoporis nigra (Stimpson), Proc. Philad. Acad. Nat. Sci.
vol. vii. 1855, p. 380.
Synonymy: Doris nigra Stimpson, 1855.
Dorvopsis nigra auct.
Doridopsis nigra auct.
Body. The body is of moderate size, elliptical, and somewhat depressed.
The dorsal surface is finely granulate, while the mantle is more smooth. The
LINN. JOURN.—ZOOLOGY, VOL, XXXY. 40

562 PROF. C. H. O'DONOGHUE: REPORT ON

mantle is well developed, fairly thin, and extends beyond the body all round,
save that the tip of the tail probably projects slightly during locomotion.

Colour, In the preserved specimens the colour is black, shading off toa
dark grey near the mantle-margin, but showing no sign of the red margin
described by Stimpson in the living animal. It is covered irregularly with
tiny white spots, which grow more abundant and larger towards the margin
of the mantle. The branchize are of the same dark colour as the body. The
under surface of the foot is also of a dark grey shade. In life the white
spots are more striking and the sole of the foot of lighter grey.

Dimensions. The specimens varied in size, and the largest measured
23 mm. long by 15°5 mm. wide and 9°2 mm. high. The smallest was
only 12°5 mm. long.

Head. The head is very small and inconspicuous, and bears a much reduced
tentacle at each side. The mouth is a small circular pore lying between the
lips of a deep cleft in the anterior margin of the foot.

Foot. The foot is moderately large; its sides are continued out a short
distance as shallow flanges and its posterior is bluntly pointed. The front
end is deeply cleft, and between its margins, as noted above, the mouth is
situated.

Rhinophores. The rhinophores are fairly small, completely retractile within
sheaths with smooth circular margins, and beara perfoliate clavus on a short
cylindrical stalk.

Branchie. The tripinnate branchial plumes are eight in number and
arranged in the form of a circle, incomplete behind, but the gap is filled in
by the low anal papilla. They are of the same general colour as the dorsal
surface.

Radula. No trace of a radula or pharyngeal complex is present. The
cesophagus takes the form of a straight narrow tube.

Notes. In all seven specimens were represented in the collection: three
large specimens from Sandy Isle; three smaller specimens from Wooded
Isle ; one deformed and swollen specimen from Pelsart Island.

This species of Dendrodoris seems to be widely spread in the Indian and
Pacific Oceans, and the type-specimen was described from Loo Choo and
Kikaisima.

Species DENDRODORIS MAMMOSA (Abraham), Proc. Zool. Soc. 1877, p. 266,
pl. Ixix. figs. 20-21. (P28. fig. 15.)

Synonymy : Doris mammosa Abrahaw.
Doriopsis mammosa 2uct.
Doridopsis mammosa auct.

Body. The body is fairly large and of an elongated oval shape. The
mantle is strongly developed, and projects all round as a thin fold about half
the width of the body, and has a wavy margin. The very soft dorsum and
in part the mantle also are covered with very large irregular warty

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 56:

protuberances, giving them a very uneven appearance. The large projections
also have smaller ones near their bases, and similar small ones are scattered
over the general dorsal surface. Three noticeably large projections lie on
each side between the rhinophore and the gills, and two more are situated
between the rhinophores.

Colour. The general body-colour of the preserved specimen is a dirty
yellowish grey, becoming darker towards the periphery. - Down the middle
of the back is a series of three large roughly rhomboidal black marks, which
have other verydark, almost black, markings within them. Five or six similar
but slightly smaller marked areas lie lateral to the large projections on each
side, the first pair of lateral marks being in front of the rhinophores and the
last behind the branchiz. The clavus of the rhinophore is light-coloured
and the tips of the branchiee very dark ; Professor Dakin informs me that in
life the pattern was similar, but the colours brighter and the contrast more
marked.

Dimensions. The preserved specimen measured 72 mm. long by 40 mm.
wide (mantle extended) by 17 mm. high. The foot measured 65 mm. long
by 15 mm. wide, so that the mantle, when extended, projected 12°5 mm.
beyond the foot.

Flead. The head is extremely small and inconspicuous, and the mouth is a
tiny aperture lying between the two edges of the front end of the foot. On
each side of the head is a tiny flap-like tentacle about 1°5 mm. long, partly
retracted within a shallow groove-like cavity.

Foot. The foot is well developed, long, fairly narrow, and with an expanded
flange-like margin. At the anterior end the foot bears a deep longitudinal
cleft, between the edges of which lies the mouth.

Rhinophores. The rhinophores are fairly small clavate structures with a
perfoliate clavus, and are entirely retractile within cavities provided with an
elevated tubular margin. They lie fairly close together near the anterior
end.

Branclie. The five branchial plumes are tripinnate and arranged in a
fairly close circle, with a tendency to fusion at their bases. Hach is dendri-
form. ‘They are retractile withina wide shallow cavity with a slightly raised |
margin.

Radula. No trace of a radula or pharyngeal complex is present, and the
first part of the cesophagus is in the form of a straight narrow tube.

The anus lies between the bases of the two hindermost gill-plumes upon a
fairly well-developed sub-conical papilla.

The genita aperture is, as usual, high up on the right side of ihe body
about one-quarter of the way from the anterior end.

Notes. Only one specimen is represented in the collection with a label
“ Nudibranch from Fremantle, W.A.” It is not therefore an Abrolhos

type, so far as we know.

40%

564 PROF. C. H. O’DONOGHUE: REPORT ON

The type-specimen described by Abraham (2) measured 56 mm. long by
44 mm. broad and 16 mm. high, so that it was not so large as the present
one. No locality is given, but it is stated to be “obtained during the
Antarctic Expedition.” It was possibly obtained at some port of call, how-
ever, for the Dendrodorids as a genus are characteristic of the warmer seas.

Family EUPHURID (Potycrripm, Potycrrap#).

The body is more or less elongated and limaciform. The dorsum is not
marked off from the sides or separated by a prominent margin. The frontal
veil is more or less prominent, simple, or furnished with simple or compound
appendages, and the dorsum generally has a solitary dorsal appendage
(branchialis) or several dorsal and lateral. The branchize are generally
paucifoliate, but may be composite ; they are not retractile within a cavity.
The tentacles are small, lobiform, folded, or auriform. The foot is not broad,
and generally rounded anteriorly. The bulbus pharyngeus is simple. The
buccal cavity is often armed with laminz (often composed of minute rods).
The radula nearly always has no rachidial teeth ; the majority of the pleural
teeth are uncinate, and the outermost teeth simple, not hamate. The glans
penis is armed ; two spermathecas are present.

In 1798 Cuvier instituted the name Tritonia for a genus of Nudibranchs,
but while describing the genus he mentioned no particular species, and later,
in 1803, he enlarged this description and dealt with a definite and new
species, 7. hombergu. Lamarck in 1801 took Cuvier’s name and gave as e
species of the genus 1. clavigera. Rafinesque in 1815 proposed the name
Euphurus for the Tritonia of Lamarck—i. e., with T. clavigera asa type. At
a later date Johnston (1838) dismissed Lamarck’s species as being outside
the limits of Cuvier’s genus, and proposed for it the name Triopa clavigera,
under which it has since been dealt with. To turn to other animals than
Nudibranchs, however, we find that in 1774 Tritonium was employed as a
generic name by Miller, and in 1800 Meigen used Vritonia as the name
of a genus.

Thus it will be seen that the term Zritonia as applied to Nudibranchs
refers to two entirely different and unrelated forms, and in any case the
name is pre-ocecupied. As noted previously, Tritonca Cuvier (i.e., with
T. hombergu as type) has to be replaced by Spherostoma Maceillivray, and
as will be seen above Jritonta Lamarck (7. e., with 7. clavigera as type)
becomes Huphurus Rafinesque. and this intercepts the later name Triopa
Johnston. In this way the oldest generic name in the present family
becomes Huphurus, from which the family-name is derived. This matter
is also referred to by Iredale (51) and Iredale and O’Donoghue (51a).

Genus Noroporis Bergh, Journ. Mus. Godeffroy, viii. 1875, p. 64.
Type by monotypy: NV. cttrina Bergh, 1. c.
The body is limaciform with the sides not marked off from the dorsum ;

3
it is hard and rough, often marked with prominent ridges. he frontal veil

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 565

is large. The branchiz and sometimes the rhinophores are protected by
valves. The rhinophores are not perfoliate. There is no labial armature of
rods or spines, but a chitinous ring may be present. No rachidial tooth is
present ; the pleurals are similar and hamate with indications of an accessory
denticle.

Species Noroporis GARDINERI Hliot, Fauna & Geog. Maldive and Laccadive,
vol. ii. pt. 1, 1903, p. 548. (PI. 28. figs. 18, 19 ; Pl. 30. figs. 62-64.)

Body. The body is limaciform and covered with small tubercles, which are
more numerous and larger at the anterior end. Nearly halfway back the
body reaches its highest point, and here arise the branchice covered by their
three characteristic protecting valves. A row of larger tubercles runs from
the sides of the oral veil to the branchial valves. Beyond this the body
falls to a lower, narrower, tail-like region. A well-marked valve-like
tubercle lies on the outside of each rhinophoral aperture, and between these
openings is a group of two or three large tubercles. Lateral to the rhino-
phores the noteeum passes out and forwards as a very well-marked oral veil
with a tuberculated margin. The very tough skin contains a number of very
hard, semi-transparent spheres, which appear to be composed of a chitinous
material. They are irregularly scattered, and may be sparse or so close
together that they form hard masses.

Colour. According to the coloured sketch made by Professor Dakin, this
species is of a deep chrome-yellow all over; the oral veil and branchial _
valves are of the same colour. There was apparently no trace of the black
spots deseribed by Eliot. 3

Dimensions. The specimens were all about the same size and measured
when preserved 43 mm. long by 9 mm. wide by 10 mm. high ; length of the
tail-region 18 mm. Hliot’s specimen was 33°5 mm. by 9 mm. by 12 mm.

Head. The head is small and in the specimens completely hidden between
the front end of the foot and the overhanging oral veil. This veil is very
well developed, andin all the specimens is tightly curled down over not only
the head but also the anterior end of the foot.

Foot. The foot is fairly broad and lanceolate. ‘The front end is rounded
and the hinder end passes off to a point.

Ehinophores. The rhinophores are completely retractile within deep
cavities. The aperture is in the form of an oval slit placed at an angle of
about 45° to the main axis of the animal. At the posterior outer corner of
the aperture liesa very large tubercle whose flap-like apex overhangs it.

Branchiw. The branchiz are fairly numerous, very small, and completely
hidden by the three large valves. ‘The three valves rise from a transverse
ridge stretching across the middle of the back. The middle one passes back
and curls downwards until it practically touches the dorsum. The lateral
valves pass backwards, down and round partly forwards again, forming the
commencement of a spiral. They also practically touch the dorsum. All

PROF. C. H. O’DONOGHUR: REPORT ON

Ot
oS.
for}

the valves, particularly the lateral ones, possess on their margins curious
spatulate processes, which were described and figured by Hliot. The valves
are very hard and bend over almost to touch the dorsum, so that the tiny
gill-plumes, apparently arranged in three groups, cannot be seen without
cutting off the valves.

Labial Armature. According to Eliot there is no labial armature, but in the
specimens here described a circular ring of chitinous cuticle was present.
Tt is not large and quite thin, but, nevertheless, it resisted boiling in potash
solution and separated off from the lips as a distinct ring.

Radula. The radula contains no rachidial teeth. It is only moderately
developed, and contains from 56-58 rows. ‘The first row contained 18.0.18
(2. e., 36 teeth), and the middle rows (34-42) .0. (84-42), 7. e. (68-84). The
teeth are erect, moderately long, and slender, and so closely packed at the
outside that they are difficult to count accurately. They are a little larger
on the outside, but otherwise very similar throughout. Hach consists of a
more or less rectangular blade-like plate, one corner of which is carried on
as a thin, slightly curved rod, bearing at its tip a small rounded secondary
denticle. They are similar in general appearance, although differing in
detail from those deseribed by Bergh for JV. citrina (11).

Notes. Three specimens were represented in the collection, and bore the
label “ Ist Island. Shore Collecting. Outer side.”

Professor Dakin, in his general account of the Abrolhos, says of this
species :—“ A straggling brilliant lemon-yellow sponge was one very evident
specimen. After turning several stones over and noticing what appeared to
be pieces of this sponge falling off, it was discovered that the falling pieces
were Nudibranchs resembling the sponge both in colour and general appear-
ance. ‘The species belongs to the genus Notodoris, this being the first record
for the genus on the Australian coast. It was instituted by Bergh for a
single specimen of JV. citrina from Rarotonga. Two other species have been
made by Hliot for specimens from Zanzibar and the Maldive Islands. All
three are yellow in colour. It is stated in Khot’s report that nothing is
known of the habits of these animals, and further :—‘ With Mr. Gardiner’s
specimen is a piece of hard yellow sponge. There is no note, but as the
colour and consistency of the sponge closely resemble those of the Nudibranch
it is highly probable that the latter frequent it.’

“It is interesting to find that this is actually the case and in an entirely
different region. Our specimens were always found associated with this
sponge. The Nudibranch moves about slowly, so far as could be observed,
and apparently lives in the dark under coral blocks.”

This association between Nudibranch and sponge is not unknown, as
witness the entirely different brilliant red Nudibranch Rostanga pulchra living
on a similarly coloured sponge on the Pacific coast of North America (vide
O’ Donoghue, 60, p. 152).

OPISIHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 567

As pointed out in the passage quoted above, only three species of this
remarkable genus have been recorded : namely, NV. citrina Bergh from Raro-
tonga by a single specimen, WV. minor Hliot from Chuaka on the east coast of
Zanzibar by a single specimen, and JV. gardineri Eliot from the Maldive
Islands also by a single specimen.

In spite of certain minor differences, 7. e. the presence of black spots, the
presence of a Y-shaped arrangement of small tubercles on the dorsum, the
absenee of labial armature inter alia in Eliot’s specimen, I have no
hesitation in referring the present form to his species, WV. gardineri. The
size is about the same, the teeth similar, the valved rhinophoral aperture
similar, and also the branchial valves with their elaborate spatulate
processes.

As will be seen the genus is very little known, and it is to be hoped that
sufficient material for a detailed investigation of its structure will be forth-
coming. It apparently contains three distinct species :—

N. crrrina Bergh.—Pale yellow ; somewhat broader; 7 quadripinnate
gills ; gill-valve with 8 points more than one-third of the way for-
ward; valved rhinophoral aperture; radula 56 rows typically
14.0.4, tooth with indication of third blunt denticle.

N. Garpryert Hliot.—Chrome-yellow with or without black spots,
narrower ; numerous tiny gill-plumes ; gill-valves marked, three in
number, furnished with spatulate processes, about halfway back ;
valved rhinophoral aperture ; radula 56-58 rows typically (84-42) .
0. (84-42), tooth with only second blunt denticle. i

N. wiyor Eliot.—Lemon-yellow sharply marked with black lines,
narrower ; 27 small tufts of gills in three areas ; gill-valves three in
number, inconspicuous, simple ; radula 33 rows typically 25.0. 25,
tooth not nearly so distinctly denticulate as in foregoing species.

These differences appear to warrant the retention of the three forms as
valid species.

Genus NemBrorua Bergh, Malakol. Unters. Hoft xi. p. 450 (1877).

Type here designated, NV. nigerrima Bergh.

The body is limaciform, nearly smooth ; dorsum not differentiated from
sides ; the rhinophores are retractile with a perfoliate clavus ; the branchize
are paucifoliate and bi- or tri-pinnate; the tentacles are short and lobate ;
the foot is quite narrow. The labial armature is inconspicuous or absent.
The radula fairly narrow ; the rachidial tooth is depressed, subquadrate, or an
armed curve; the first pleural tooth is large and falciform, and the several
external teeth are depressed.

568 PROM. C. H. 0 DONOGHUE : REPORT ON

The hermaphrodite gland is connate with the liver ; no discrete prostate is
present ; the glans penis is armed,

There is a possiblity that the generic name Nembrotha Bergh, 1877, will
have to be replaced by Angastella Angas & Cross, 1864: the form
described by the latter being very much like those in Bergh’s genus—
indeed, Bergh himself includes it as a doubtful member of the genus. The
matter can hardly be definitely settled unless the collection of further
specimens renders it possible to examine it more closely.

Species NEMBROTHA PURPUREOLINEATA, sp. nov. (PI. 28. figs. 16, 17;
Pl. 30. figs. 60, 61.)

Body. The body is limaciform, rising from the front end to the middle,
where it bears the branchise, thence it gets lower again and terminates in a
short tail. The smooth dorsum passes over into the side of the body without
line of demarcation or pallial edge. The anterior end bears a narrow
veil-like expansion.

Colour. The general body-colour of the preserved animal is a dirty
yellowish grey marked with pale brown lines: A broad dark band com-
mences just behind the rhinophores and passes back in the mid-dorsal line to
the branchie. A thinner dark line passes round the front, anterior to the
rhinophores, and back on each side of the dorsum. Behind the branchie
these two lines converge and meet, about halfway between the gills and the
posterior end, to form one line in the middle which runs back to the end of
the tail. Another line starts at the front end about halfway up the body on
each side. It runs back roughly parallel with the edge of the foot to join
the dorsal line near the tail. Professor Dakin informs me that in life the
body-colour of the animal was a translucent slate-grey and the dark bands
were purple-brown.

Dimensions. The form of the specimen was not well preserved, but it
measured 35 mm. long by 10mm. wide and 11 mm. high.

Flead. The head is small and not conspicuous. At each side it bears a
tiny tentacle, somewhat knob-shaped in the retracted condition, and between
them is the circular mouth.

Foot. The foot is linear and not strongly developed. The hinder end
passes off into the pointed tail, and the anterior end is cleft and las rounded
corners.

Rhinophores. The rhinophores are small, have a perfoliate clavus, and are
completely retractile within cavities with smooth round apertures.

Branchie. The non-retractile branchize consist of five bipinnate plumes,
joined at their bases. The anterior one is in the middle line and larger than
the others.

Radula, The radula is relatively small and when flattened measured 4 mm.
by 2mm. It bears 32 rows of teeth and the number in the oldest row is

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 569

7.1.1.1.7 (¢.¢.,17) and in the newer portions 8.1.1.1.8(@e.,19). The
rachidial tooth is in the form of a broad irregular octagon with its posterior
side uncurved and thicker in its anterior half. Its front edge is straight,
but furnished with four curious papilliform denticles directed backwards—
one of the inner of these bifid. In the radula of the specimen examined
it was always the inner one on the same side. ‘The first lateral has an
irregular elongated base lying at an angle of about 40° to the middle line
of the radula. It bears two very strong, backwardly directed, conical
spines—one at its anterior end and the other about two-thirds of the way
back. This is followed by a series of teeth, seven or eight in number. They
take the form of approximately square plates in the inner teeth, but become
oblong and less strong towards the outside.

Labial Armature. The lips are furnished witha lining of thin chitin, which
has an oval opening, whose long axis is vertical.

The anus lies on a short rounded papilla in the middle line bekind the
Junction of the bases of the posterior branchial plumes.

The genital aperture lies near the anterior end in the light space between
the two lateral dark bands and about halfway between the head and the
branchize.

Notes. The species is represented by one specimen, which was taken on the
Abrolhos, but with no details of the exact locality.

In spite of the presence of a labial cuticle rather more strongly developed
than in other forms, there is no doubt that this belongs to the genus
Nembrotha. It does not appear to agree with any previously described.

A Coprpop PARASITIC ON C'nRATOSOMA BREVICAUDATUM.

The presence of Copepoda parasitic upon Nudibranchs has been reported
by several authorities. The first record is apparently that of Leydig (56), in
1853, who constituted the genus Doridicola for Copepods found on Dorids.
Similar forms are also reported by Alder and Hancock (5) from Arechidoris
tuberculata and Antiopa cristata, and they were referred to the genus
Ergasilus, although they were probably congeneric, if not even conspecific,
with Leydig’s forms, as was pointed out later by Hancock and Norman (47).
They are not highly modified animals and apparently more in the nature
of epizoitic forms, and their parasitism has not been proved.

In addition to these, however, a group of highly specialised forms,
undoubtedly parasitic, have also been described. The first, reported by
Hancock and Norman (47), are :—Splanchnotrophus gracilis, parasitic on
Acanthodoris pilosa and Idalia aspersa from the Hnglish coast, and S. brevipes,
parasitic on Doto coronata and Coryphella rugibranchialis also from the English
coast. The only other authority that has dealt with these parasites is
R. Bergh, who has recorded the following :—

570 PROF. C. H. 0’DONOGHUE: REPORT ON

| Parasite. Host. heen Oeand
— dea steel

Splanchnotrophus brevipes. | Galvina viridula. North Sea (21, p. 568).

| Splanchnotrophus sp. Coryphella rufibranchialis. Norway (21, p. 568).
Ismalia monstrosa. Phidiana lynceus. W. Coast, S. America (9).

ae os re inca. W. Coast, 8. America (26).

sy aA Archidoris incerta. W. Coast, 8. America (26).
Ismailia sp. | Lomanotus gener. Mediterranean (21, p. 558)
Briarella microcephala. Ceratosoma trilobata. Red Sea (13, p. 409).

| Briareila sp. probably. | Glossodoris elisabethina. Philippines (16, p. 472),

| B. microcephala. | Asteronotus bertrana. Philippines (19, p. 641).

The total number of species is very small, only four named for certain and
a possibility of seven at the most, and, from the infrequency with which they
have been recorded, it would appear that they are rare. Indeed, in many
hundreds of specimens I have handled from British Columbia, I have not
yet encountered one. In spite of this, however, it will be seen from the
above list that they are widely distributed over the world. The following
form is of interest, not only on its own account, as a new species of a rare
group, but also because it shows that the group also extends to the Australian
seas.

Family CHONDRACANTHID.
Genus SpLaNcaNorropuus Hane. & Norm. Trans. Linn. Soe. Lond.
xxiv. 1864, p. 51.

Type here designated: S. gracilis Hanc. & Norm. J. c. p. 51.
Coast of England.

Female. Head and thorax either blended into a single segment, the
thoracic portion of which is furnished on each side with unarticulated arm-like
appendages or lobes, or the first part only of the thorax is united with the
head and the last part forms a second, but comparatively minute, segment.
In this case, however, all the thoracic appendages are attached to the first
segment. Hirst antennee minute and few-jointed; second larger, in the
form of prehensile hooks. Labrum large, overhanging the mandibles, which
organs, together with the maxilla and two pairs of foot-jaws, are minute and
crowded round the mouth. Thoracic feet two pairs, minute, simple or two-
branched, terminating in hooks. Abdomen two-jointed, the last joint ending
in two caudal appendages, which are furnished with one or two simple sete.
Ovigerous sacs elliptical.

Male minute. Cephalothorax with lateral appendages and divided into
fonr segments, the first of which bears two pairs of thoracic feet.

OPISTHOBRANCHIATA FROM THE ABROLEOS ISLANDS. 571

Species SPLANCHNOTROPHUS SACCULATUS, Sp. nov. (PI. 80. figs. 65, 66.)

Female. The body is elongated and roughly cylindrical, but slightly
depressed dorso-ventrally, and it is encased in a thin, not very strong,
layer of semi-transparent chitin. It is composed of a cephalothorax, of
which the anterior portion is the head, but it is not distinctly marked off
from the thorax, and a much smaller abdomen. On the dorsal surface of the
cephalothorax are three sac-like outbulgings of the body-wall. The first of
these is in the form of a transverse collar-like ridge right across the body,
and if is somewhat firmer than the other two. The second is much larger,
more inflated, and more irregular. The anterior border and the lateral lobes
of it are sharply marked off from the body, but in the mid-dorsal region the
hinder parts of the two lateral lobes merge with the general surface of the
body. The third sac is, if anything, larger than the second, and its lateral
lobes show a slight marginal indentation.

The last portion of the thorax is narrower than that in front, but it swells
out towards its hinder edge and then rapidly narrows again still further to
the point where it joins the abdomen.

From the ventro-lateral aspeets of the cephalothorax arise three pairs of
long, tapering, cylindrical outgrowths. The first pair arise at a level between
the first and second dorsal sacs. he second pair arise at a level between the

“second and third dorsal sacs, and the third pair touch the second and are

level with the third dorsal sac. hese lateral outgrowths are not, as might
be supposed at first sight, modified appendages, for the rudiments of these lie
on the ventral surface, but they are outgrowths of the body-wall. They are
very conspicuous, as they almost double the length of the animal itself, and
apparently are concerned with the absorption of nutritive material from the
host, since they lie in the spaces between the viscera.

Strings of eggs, either in single or double rows, show through the body-
wall and form a network of interlacing lines in the various parts of the
cephalothorax, the dorsal sacs, and the basal portions of the long outgrowths.

The first antennee are a pair of minute, stumpy, seemingly uniramous
appendages lying under the head a short distance from the anterior end.
Hach is apparently composed of three joints and bears a number of relatively
stout spines ; there are three of these on the basal joint, two on the next, and
two on the terminal joint. The second antenne, while still small, are con-
siderably larger than the first, and are plump, fleshy, uniramous appendages
also showing indications of three joints. They are relatively shorter than the
correspoding members in S. gracilis. Stout spines are also present on
these: two on the basal joint, one on the next, and two on the terminal
joint.

The labrum is large, curved ventrally, and sub-triangular in shape with a
rounded apex and a deep curved indentation in the base. The mouth pre-
sumably lies under this indentation. The mandibles are small and consist of

572 PROF. C. H. O'DONOGHUE? REPORT ON

a soft triangular base, which bears at its antero-median corner a small stiff
blade with three denticles. The maxillee are very small triangular lobes
partly hidden by the mandibles. Hach bears a spine-like prolongation at
its antero-median corner. The first maxillipedes are somewhat triangular in
shape, not unlike the mandibles, save that in place of the blades they bear
spine-like processes. The second maxillipedes are long thin structures,
expanded dorso-ventrally in their anterior region, and bearing spine-like
prolongations at the anterior end. They are approximated in the mid-ventral
line over the front part of their length, but diverge posteriorly. Altogether
the mouth-parts show a considerable resemblance to those of S. gracilis, but
their proportions are different, and they appear to be a little more degenerate.

On the ventral side of the cephalothorax there are a pair of rounded
papilliform processes opposite the bases of the first lateral appendages, and
another and quite similar pair opposite the ends of the last pair of lateral
outgrowths. ‘hese are in the same relative position as the tiny legs in
S. gracilis, as shown in the drawing of Hancock and Norman, but, while each
bears a small spine, they are not nearly so much like thoracic appendages as
those of S. gracilis.

If we adopt the interpretation given by Hancock and Norman (47, p. 52),
two regions can be distinguished in the abdomen: the first an inverted
cone joiming on to the posterior end of the cephalothorax and the second
another inverted cone with a rounded edge at the base borne on the pre-
ceding part. This second cone has a firmer chitinous covering than the
cephalothorax and bears four annular grooves ; thus it is divided up into five
rings, but whether these represent actual abdominal segments or not it is
hardly possible to say. Hancock and Norman group the anterior cone-shaped
portion together with the basal ring of the second cone as the first complete
abdominal segment, although in the present species there is a distinct,
annular, articular groove separating them. The basal ring of the second
cone-shaped portion has two lateral projections upon which the egg-sacs are
borne. On the terminal portion of the abdomen are two small cylindrical
projections each terminating in a short spine.

The egg-sacs ure elongated, cylindrical, soft-walled sacs with rounded ends.
The sacs measured, in one individual, 6°2 mm. long by 1:2 mm. wide. _ The
eggs within them are apparently just crowded together without definite
arrangement, as is characteristic of the family Chondracanthide, and do not
exhibit the linear arrangement that is found in the Caligide.

Notes. Two specimens of this species were found upon one individual of
C. brevicaudatum, both female and about the same size. ‘The one of these
measured 8°5 mm. long by 4:1 mm. broad; its ege-sacs, as noted above, were
6°2 mm. long by 1:2 mm. wide and the length of the longest lateral process
was 14°75 mm. One specimen was partially embedded in an actual hole in
the body-wall of the Nudibranch about 5 mm. behind the urinary aperture,

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 573

and looked as if it were in the process of boring its way down to get into the
body-cavity. ‘his specimen was very noticeable from the outside, and its
long lateral appendages and most of the body projected freely from the
dorsal suface of its host. In examining this individual it was noticed that
two peculiar sac-like structures containing eges of a deep golden-yellow
colour were projecting from the much dilated renal pore. Further exami-
nation and dissection revealed the presence of a second and slightly larger
example of the parasite completely hidden within the body of the host—the
measurements of this specimen are given above. It was lying within the
renal duct at its posterior end, but it was so large and its lateral arm-like
appendages spread about so much among the viscera that it was not possible
to see whether the animal was completely confined within the renal duct,
which had become enormously dilated and ramified, or whether it had broken
through the wall of this structure.

While the specimen on the outside of the Nudibranch contained eggs
within the body, visible through the semitransparent body-wall as a tracery
of lines, it showed no sign of egg-sacs. If these had not been torn off, there-
fore, it would appear as if the production of eggs had not proceeded far in
this individual. In the second specimen, on the other hand, egg-sacs were
present and of large size. As noted above, they projected through the renal
pore, which was much enlarged or perhaps actually broken, and also projected
through the partially closed branchial aperture. They were extremely thin-
walled and delicate, and in the subsequent process of excavating the rami-
fications of the parasite from the body of the host they were badly broken
up—tfortunately not until after they had been measured. When the branchize
were expanded, the egg-sacs must have lain among the branchial plumes as
deseribed in S. gracilis. The body of this individual also contained a number
of eggs showing through the body-wall, and arranged in a series of inter-
lacing lines which passed out into the lateral processes.

No trace of a male individual was found either on or near the females, or
upon or in the body of the host. This is rather remarkable in view of the
fact that Hancock and Norman state that the males, sometimes to the number
of a dozen, are generally to be found associated with the feinales.

The present species is undoubtedly congenerie with S. gracilés Hancock
and Norman, and quite similar to it in a number of ways. It differs in a
number of points, however : the presence of the dorsal sacs, the segmentation
of the abdomen, the more degenerate condition of the thoracic limbs, the
greater length of the lateral appendages, and certain differences in the mouth-

parts inter alia, and so it is here listed as a new species under the name
Splanchnotrophus sacculatus. It is interesting to find a form from the
Antipodes so closely allied to S. gracilis, heretofore only recorded from
the European seas.

574. PROF. C. H. O'DONOGHUE: REPORT ON

List or REFERENCES.

. Apratam, P, S.—“ Notes on some Genera of Nudibranchiate Mollusca, with Notices

of a New Genus and some hitherto undescribed Species in the Collection of
the British Museum.” Ann, & Mag. Nat. Hist. (4) xviii. 1876, p. 182.

. Id. ‘Revision of the Anthobranchiate Nudibranchiate Mollusca, with Descriptions

or Notices of forty-one hitherto undescribed Species.” Proc. Zool. Soe.
London, 1877, p. 196.

. Apams, H., and Apams, A.—The Genera of Recent Mollusca. London, 1858.
. Adams, A., and Rruve, L.—Zoology of the Voyage of H.M.S. ‘Samarang.’ Mollusca.

Brit. Museum, London, 1848.

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. Id. “Notice of a Collection of Nudibranchiate Mollusca made in India by

Walter Elliot, Esq., with Descriptions of several new Genera and Species.’’
Trans, Zool. Soc. London, v. pt. 3, 1864.

. AnaAs, G. F.—“ Descriptions d’espéces nouvelles appartenant & plusieurs genres de

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Sud), accompagnée de dessins fait d’aprés Nature.” Jour, de Conchyl. 3e série,
Tome iv., xii. 1864, p. 43.

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1866, 1867.

. Id. ‘Malakol. Untersuch. IL.” Semper’s Reis. im Archipel d. Philipp. 1871.
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1876, p. 9.

_ Id. “ Malakol. Untersuch. X.” Semper’s Reis. im Archipel d. Philipp. 1876.
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. Id. “ Malakol. Untersuch. XI.” Semper’s Reis. im Archipel d. Philipp. 1878.
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Gesell. in Wien, xxvii. 1878.

. Id. “Neue Nacktschnecken der Siidsee. Pt. 1V.” Journ. Mus. Godeffroy, Heft xiv.

1878, p. 1.

. Id. “ Malakol. Untersuch. XIV.” Semper’s Reis. im Archipel d. Philipp. 1878.
. Id. “Neue Chromodoriden.” Malako, Blatt. i., n. s., 1879.
. Id. “ Beitrige zur Kenntniss der Aeolidiaden. Pt. VI.” Verh. d. k.k. zool.-bot.

Gesell. in Wien, xxviii. 1879.

. Id. “Malakol. Untersuch. Suppl. I.” Semper’s Reis. im Archipel d. Philipp. 1880.

. Id. “Report on the Nudibranchiata.” ‘Challenger’ Expedition, vol. x. 1884.

. Id. “Malakol. Untersuch, XVI.” Semper’s Reis. im Archipel d, Philipp. 1889.

. Id. “System der Nudibranchiaten Gasteropoden. Malakol. Untersuch, XVIII.”

Semper’s Reis. im Archipel d. Philipp. 1892.

. Id. “Die Opisthobranchier der Sammlung Plate.” Zool. Jahrb., Supp. iv. Heft iii.

1898.

32.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 575

. Bereu, R.—“ Malakol. Untersuch, Die Pleurobranchiden.” Semper’s Reis. im

Archipel d. Philipp. 1898.

. Id. ‘“Malakol. Untersuch.” Semper’s Reis. im Archipel d. Philipp. vi. 1904.
. Brarnvittn, H. M. D. pr.—Manuel de Malacologie et de Conchyliolovie. Paris, 1825.
. CHEESEMAN, S. T.—“ Descriptions of three new Species of Opisthobranch Mollusca

from New Zealand.” Proc. Zool, Soc. Lond. 1878, p. 275.

. Cuvier, G.—“ Mémoire sur le Genre Doris.” Ann. du Mus. nation. d’Hist. nat.

Tome iv. 1804.

Daxiy, W. J.—“ The Percy Sladen Trust Expedition to the Abrolhos Islands (Indian
Ocean) Report. I. Introduction; General Description of the Coral Islands
forming the Houtman Abrolhos Group; the Formation of the Islands.” Journ.
Linn. Soc. Lond. vol. xxxiv. No. 226, 1919.

33. Hanus, N. B.—“ Aplysia.” Liverpool Mar. Biol. Comm. Mem. xxiv. 1921.
34, Hurenprre, C. G.—<Symbole Physic, Animalia Evertebrata exclusis. insectis,

35.

36.

37.

38.

39.

40,

”

percensuit Dr. C. G. Khrenberg.
Berolini, 1831.

Ertor, C.—“ Nudibranchiata, with some Remarks on the Families and Genera and
Description of a new Genus, Doridomorpha.” Fauna and Geog. of the Maldive
and Laccadive Archipel. vol. 11. pt. 1, 1903.

Id. ‘On some Nudibranchs from Hast Africa and Zanzibar. Pt. 1.” Proce. Zool. Soe.
Lond. pt. ii. 1903.

Id. “On some Nudibranchs from Hast Africa and Zanzibar. Pt. V.’’ Proc. Zool.
Soe. Lond. pt. ii. 1904.

Id. “Nudibranchs from New Zealand and the Falkland Islands.’ Proc. Mal. Soc.
Lond. vol. vii. 1907.

Id. ‘Notes on a Collection of Nudibranchs from Ceylon.” Spolia Zeylanica,
Colombo, vol. vi. pt. 23, 1909.

Id. Monograph of the British Nudibranchiate Mollusca. Part VIII, Supplementary.
Ray Society, London, 1910. ;

Series prima cum tabularum decade prima.

41. Fiscumr, P.—Manuel de Conchyliologie et de Paléontologie conchyliologique. Paris,
1884.

42. Forbes, Ii.—‘‘On some new and rare British Molluscs,’ Ann, & Mag. Nat. Hist. v.
1840,

43, GistEL, J.—Naturgeschichte des Tierreichs fiir Schulen. 1848.

43

a. GRAvennorst, I. L. C.—Tergestina, oder Beobachtungen und Untersuchungen iiber
einige bei Triest im Meere lebende Arten der Gattungen Octopus etc.
Breslau, 1831.

44, Gray, J. H.—“ A List of the Genera of Recent Mollusca; their Synonyma and Types.”

45
46

47

48.

cE)

Proc. Zool. Soc. Lond. 1847, p. 129.
. Gray, E.—Figures of Molluscous Animals. V. London, 1850.
. Hancock, A.—“On the Anatomy of Dortdopsis, a Genus of the Nudibranchiate
Mollusca.” Trans. Linn. Soc. Lond. vol. xxv. pt. 2, 1865.
. Hancock, A., and Norman, A. M.—“ On Splanchnotrophus, an undescribed Genus of
Crustacea parasitic in Nudibranchiate Mollusca.” Trans. Linn. Soc. Lond.
vol. xxiv. 1863.
van Hassett, J. C.—“ Uittreksel uit eenen Brief yan Dr. J. C. van Hasselt aan Prof.
yan Swinderen.’” Alg. Konst en Letter-Bode, 1 Deel, 1824. Also “ Extrait
une lettre du J. C. van Hasselt au Prof. van Swinderen sur les Mollusques
de Java.” Bull. sci. nat. et de géol. iti. 1824.

. Hepury, C.—A Preliminary Index of the Mollusca of Western Australia.” Journ,

Roy. Soc. Western Australia, 1916.

576 PROF. C. H. O DONOGHUE: REPORT ON Ee

50.

51.

Heptry, C.—“A Check-list of the Marine Fauna of New South Wales. Pt. I.
Mollusca.” Journ. Roy. Soc. New South Wales, vol. xiii. 1918.

Irepate, T.—“ Molluscan Nomenclatural Problems and their Solutions. I.” Proc.
Malac. Soc. Lond. vol. xiii. 1918.

51a. IREDALE, T., and O’Donoauun, C. H.—“List of British Nudibranchiate Mollusca.”

52.

53.

54.

55.

56.

57.

58.

59)

60.

61.

62.

63.

64.

65.
66.

67.

68.
69.

70.

71.

72.

73.

74,

75.

Proe. Malac. Soe. Lond. vol. xv. 1928.

von Jurrine, H.— Versuch eines natiirlichen Systems der Mollusken.” Jahrb, d.
deutsch. Malakozool. Ges., 3. Jahrg. 1876.

Id. Vergleichenden Anatomie des Nervensystems und Phylogenie der Mollusken.
1877.

Kent, W.S.—-A Naturalist in Australia. London, 1897.

Lracu, W. E.—A Synopsis of the Mollusca of Great Britain. London, 1852.

Lnypia, F.—“ Neuer Schmarotzerkrebs auf einen Weichthier.” Zeits. f. wiss. Zool.
iv. 1853.

Macerttiveay, W.—A History of the Molluscous Animals of the Counties of Aberdeen,
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Opuner, N. H.—* Results of Dr. E. Mjéberg’s Swedish Scientific Expeditions to
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Ba. lii. No. 16, 1917.

O’Donoeuur, C. H.—‘ Notes on the Nudibranchiate Mollusca from the Vancouver
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p Orpieny, A.— Mémoires sur des espéces et sur des genres nouvelles de l’Ordre des
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Id. “Remarks on Nudibranchiata inhabiting the Pacifie Islands, with Descriptions
of two new Genera.” Amer, Journ. Conch. vol. ii. 1866.

Id. “Descriptions of Nudibranchiate Mollusca inhabiting Polynesia.” Amer. Journ.
Conch. vol. vi. 1876.

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Id. Tryon’s ‘Manual of Conchology, Structural and Systematic,’ vol. xvi. Wash-
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Quoy et Gaimarp.—Zoologie, I. (Mollusques). Voyage de ‘1’Astrolabe,’ 1883.

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Smiru, H. A.—Report on the Zoological Collections made in the Indo-Pacific Ocean
during the Voyage of H.M.S. ‘Alert.’ Brit. Museum, London, 1884.

Sownrsy, G. B.—Reeve’s ‘Conchologia Ieonica,’ vol. xviii. London, 1870.

Srmrpson, W.—“‘ Descriptions of some of the new Marine Invertebrata from the
Chinese and Japanese Seas.” Proc, Acad. Nat. Sci. Philadelphia, vii. 1855,

Vaysstire, A.—“ Recherches zoologiques et anatomiques sur les Mollusques Opistho-
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Id. “Recherches zoologiques et anatomiques snr les Opisthobranches de la Mer Rouge
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Id. ‘Mollusques de la France et des Régions yoisines, Tome Premier.” Encyclop.
Scient, Paris, 1913,

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 577

EXPLANATION OF THE PLATES.

PuatTE 27.
The photographs on this plate were all taken by Mr. Herring, of the British Museum.

Fig. 1. Lethys giyantea (Sowerby). Dorsal aspect, x about 2/3. The large expanded
lobes at the anterior end of the foot and the anal funnel are clearly shown.

Fig. 2. Tethys denisoni (Smith). Dorsal aspect, x 1. The anal funnel in this specimen
is folded in.

Fig. 3. Dolabrifera pelsartensis, sp. nov. Dorsal aspect, x 1.

Fig. 4. Placobrunchus expansa, sp. noy. Dorsal aspect, X 1. The anterior extension of
the mantle lobe in front of the flattened head shows clearly on the right side,
but is torn and folded on the left. The anterior end of the head with the
tentacles is withdrawn and does not show. The tiny pericardial swelling
appears just behind the head. Owing to the shadow produced by lighting, a
false impression of a solid body is given. In reality this region is mainly com-
posed of the mantle lobes, and the actual body is not quite as wide as the head;
it cannot be distinguished from the lobes.

Fig. 5. Spherostoma dakeni, sp. noy. Veutral aspect, X 1. The mouth shows asa longi-
tudinal cleft at the anterior end, and part of the oral ve#l with one projection
appears on the left-hand side. The margins of the foot in this specimen are
almost approximated. Some of the large branchie show clearly on each side.

Fie. 6. Allotodoris hedleyz, sp. nov. Dorsal aspect, x 1. The granulate appearance of
the back and the nature of the branchial and rhinophoral apertures are clearly
visible. :

Ventral aspect of the specimen illustrated in fig. 6, x 1. The tiny head
is not visible.

Fig. 8. Glossodoris westraliensis, sp. nov. Dorsal aspect, x 1. The dark band near the
margin of the notzum shows faintly.

Lateral aspect, x 1. The broad dark band on the side of the body and
the narrow one on the upper edge of the flange of the foot are visible. The
front end of the foot is clearly bilabiate.

Fig. 10. Aphelodoris affinis Bliot. Dorsal aspect, x 1. In places the darker line around
the noteum is seen, and the dark-coloured branchiz show in the circular spot
towards the posterior end.

Sal

PratT 28.

The photographs on this plate were all taken by Mr. Herring, of the British Museum.

Fic. 11. Heavabranchus imperialis Kent, Dorsal aspect, x 2/3. The branchiz and the
clavus of the right rhinophore are clearly visible, and the relatively enormous
size of the mantle folds can easily be judged.

Fig. 12, Asteronotus fuscus, sp. nov. Dorsal aspect, x 1. This shows the ridges in the
central region cf the dorsum.

Ventral aspect of the specimen illustrated in fig. 12, x1. The relative
sizes of the foot and mantle show clearly. The indentation at the posterior end
is where a portion of the mantle has been broken away.

Fig. 14. Ceratosoma brevicaudatum Abraham. Dorsal aspect, X 1. The tail is bent to the
right, otherwise this gives a good idea of the shape of the animal and also the
position of the rhinophoral and branchial apertures and the tongue-like prolon-
gation of the dorsum.

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 4]

15.

578 PROF. C. H. O'DONOGHUE: REPORT ON

Fig. 15. Dendrodoris mammosa (Abraham). Dorsal aspect, x 1. While the general
appearance is shown well there is no indication of the colour-markings.

3. Nembrotha pwrpureolineata, sp. nov. Dorsal aspect, x 1. This indicates the
general form of the animal, save that the tail is turned to the right, and also
the position of the branchiz.

Lateral aspect of the specimen illustrated in fig. 16, x 1. This shows the
way in which the body increases in height to the level of the gills and then falls
away to the tail.

Fig. 18. Notodoris gardineri Eliot. Dorsal aspect, x 1. The general form of the body
with the tail turned to the right; the rhinophoral apertures and the branchial
valves are clearly shown.

Lateral aspect, x 1. The tail in this specimen is turned up. It shows

clearly the rough nature of the surface, the large size of the branchial valves,

and here and there some of the hard spheres.

Fig. 1

17.

19.

PLATE 29.

All the figures on this plate were drawn from preparations from specimens in the collection
and at the various magnifications shown by the aid of a camera lucida.

Fig. 20. Tethys gigantea (Sowerby). The median tooth and first two pleural teeth of one
side, X 97.
21. —— Three pleural teeth from near the middle of the row, x 97.
The six outermost pleural teeth, x 97.
Fig. 23, Tethys denisoni (Smith). The median tooth and first two pleural teeth of one side,
x 97.

24, —— Three pleurals from near the middle of the row, x 97.
25 The seven outermost pleurals, x 97.

les)
=}
09
bo
(=P)

. Dolabrifera pelsartensis, sp. noy. The median tooth and first two pleural teeth of
each side, x 97.

27. Three pleurals from near the middle of the row, x 97.
28. The four outermost pleurals, x 97.
Fig. 29. Berthella plumula (Montagu). The two innermost pleurals, x 250.

Two pleurals from near the middle of the row, x 250.
Fig. 31. Placobranchus expansa, sp. noy. Three teeth near the inner end of the radula,
seen from the side, x 190.

32. A single tooth from near the outer end of the radula, seen from the side,
x 190.

33. —— A single tooth from near the outer end of the radula, seen from above,
x 190.

34. A single tooth from the sac, seen from the side to show the relatively small

size, X 190.
Fig. 35. Spherostoma dakeni, sp.nov. The jaws, antero-ventral view, X 3.

36 The jaw, lateral view, x 3.

37. —— Two rows of the median tooth and the first pleural tooth on each side,
x97.

38. —— The four pleurals next to the innermost pleural, x 97.

39. — Three pleurals from near the middle of the row, x 97.

40. —-. Three outermost pleurals, x 97.

Fig. 41, Hexabranchus imperialis Kent. The four innermost pleurals, x 70.
The fourth pleural, side view, x 70.

43. —— Three pleurals from near the middle of the row, X 70.

44. —— Four outermost pleurals, x 70.

All the

Fig.

5. Splanchnotrophus sacculatus, sp. noy. Dorsal aspect, X 5.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS. 579

PLatTE 30.
fivures on this plate were drawn from preparations from specimens in the collection
and at the various magnifications shown by the aid of a camera lucida.

5. Alloiodoris hedley?, sp. noy. The four innermost pleurals, x 97.

3. —— Four pleurals from near the middle of the row, x 97.

. —— The four outermost pleurals, x 97.

. Asteronotus fuscus, sp. nov. The three innermost pleurals, x 150.

Three pleurals from near the middle of the row, x 150.

. —— The five outermost pleurals, x 150.
. Glossodoris westraliensis, sp.nov. The fourinnermost pleurals, lateral aspect, x 162.

Three pleurals from near the middle of the row, lateral aspect, x 162.
Three outermost pleurals, lateral aspect, x 162.

. Aphelodoris affinis Hliot. The four innermost pleurals, x 150.

Two pleurals from near the middle of the row, x 150.
The five outermost pleurals, x 150.

. Ceratosoma brevicaudatum Abraham. ‘The four innermost pleurals, x 97.

Four pleurals from near the middle of the row, Xx 97.

. —— The five outermost pleurals, x 97.
. Nembrotha purpureolineata, sp.nov. ‘he labial cuticle, lateral aspect, x 10.

The rachidial tooth,and the pleurals of one side, X 97.

. Notodoris gardinert Eliot. Labial cuticle, front view, x 10.
3. —— Jour inner pleurals, lateral aspect, x 97.

The four outermost pleurals, lateral aspect, x 97.

5. D.S. Dorsal sac;
E.L. Line of eggs; E.S. Eee-sac; L.A. Lateral appendage.

-— Ventral view of the mouth-parts, x 62. The parts are somewhat distorted
by the pressure of the cover-glass, and the first antenna on the left side of the
drawing has been turned forwards. 1 A. First antenna; 2A. Second antenna ;
L. Labrum; M. Mandible; 1Mp. First maxillipede; 2Mp. Second maxilli-
pede; Mx. First mayilla.

O'Donoghue. Journ, Linn. Soc. Zoon. Vou. XXXV,PL. 27,

Huth coll.

ABROLHOS OPISTHOBRANCHIA,.

ees
one eh
Fane

JouRN. Linn. Soc. Zoo... VoL. XXXV, PL. 28.

O'Donoghue.

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ABROLHOS OPISTHOBRANCHIA.

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O'DonoGHUE. JOURN. LINN. Soc., ZOOL. VOL. XXXV. Pt 29.

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Tet, nee re

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS,

O'’DoNxoGHuE, JouRN. LINN. Soc., ZOOL. VOL. XXXV. PL. 30.

OPISTHOBRANCHIATA FROM THE ABROLHOS ISLANDS.

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POE Hg RO WER INGAG Ii eee

Fg Ne Msi js,
Loo

Or [ae eh oet
~ MAY 2 0 1924

A
i]
Vou. XXXV. ZOOLOGY. No. 238.
CONTENTS.
Page
I. On the Genital System of Lyctus brunneus Steph., with a Note
on Lyctus linearis Goeze (Coleoptera), By A.M. Atrsoy, F.E.S.
(Communicated by Dr. A. D. Inns, F.L.S.) (Plates 831-84, and
Nextt gunes) l—4: ie teeers br pices cin. oo: Svsiassenuueidaginens ldnaseucs 581
II. The Foraminifera of Lord Howe Island, South Pacifie. By
KH. Hrron-Auien, F.R.S8., F.L.S., and A. Hartanp, F.R.M.S.
(iPilaiigs) 3 D8 Vig) gases « (ee A ep ne rt rt soo BO

Index, Title-page, Contents, etc.

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Dr. E. J. Salisbury.

GENITAL SYSTEM OF LYCTUS BRUNNEUS. 581

On the Genital System of Lyctus brunneus Steph., with a Note on Lyetus
linearis Goeze (Coleoptera). By A. M. Atrson, F.H.S. (Com-
municated by Dr. A. D. Inms, F.L.S.)

(PLatEs 31-34, and Text-figs. 1-4.)

[Read 3rd May, 1923.)

ConTENTs.

Page
ILMRCINOHON Y adodansoconoouencogadovosnvocgedsoKGO 581
diechniquerandsWethods-manaeeeerseeer eer sooo Gee
Molex Grenitali Sistemi aceeccner errr eerie 585
HemaleiGenitaliSystemla: steerer eererenicceeee ce 585
OGG panicncinon oo sunmammnenD Gn atin soneec ose doblods 590
Notes on the Genital System of Lyctus linearis Goeze.... 590

Observations on Stein’s ‘Female Reproductive Organs in
Coleoptendimmrr sre :):) san vaversercesorracr-faicea tances 591

Note on the Morphology of the Apex of the Abdomen in
LOYCHUS ORTMOALS SMEYVIN, oo0cccoccoonenondoooneboen 592
SWIMM. oo onoococoneEMecOUDDOOnUOEGUDoOGHUON DEE OOO 594
ANGMOWIECEMIGDUS 500 vdoo pene ednccob0oDd5e0ND0 Ape oak 595
FIMO ascovcooseonvsenvoncvouddboegs aotseocc 595
BQVeaaiiom OF WA IME. coocoacscnaacccoo.g0000000K6 596
Referencewle therimoyterytsjcet cir vae: cole ciousl ane sicyence arene 597

INTRODUCTION.

Ty the following description of the genital system of Lyctus brunneus Steph.,
it has been found necessary to add a comparative description of part of the
alimentary tract of both sexes; that of the male discloses nothing remark-
able in the position of these parts, whereas that of the female is curiously
situated, and appears to be only partially adapted to meet its particular
method of oviposition.

Sharp and Muir (40) figure and describe the male genital tube of
L. linearis Goeze (canaliculatus Fab.). Other than this, no reference to the
genital system of either sex of any species of Lyctus has been found.

To complete the examination of the organs ot the female ZL. brunneus, it
was necessary to cut series of sections. Longitudinal and transverse sections
were made of pupee, immature and mature adults. “Immature adults”
refers to beetles removed from their pupal chambers before either their
integument had hardened or they had eaten any wood, i.e. between four
and five days after shedding the pupal exuvie.

The beetles are not sexually mature until they are ready to emerge ;
mating invariably takes place at the earliest possible opportunity after that
event.

LINN. JOURN.—ZOOLOGY, VOL. XXXV. 42

582 MR. A. M. ALTSON ON THE

The only secondary sexual characters observed were a difference between
the ventral pygidial plates (VIII sternite) and between the apical margins of
sternites VII (the 5th visible sternite of systematists) (text-fig. 1). Beyond
these characters, unless the “ ovipositor” or the cedeagus is extruded, or a
pair are found in coition, it is impossible to distinguish the sexes externally,

size being vo guide.

Trext-Fia. 1.

Laan
Secondary sexual characters of Z. brunneus. Male and female ventral pygidial plates.
ots, base of outer sheath of ovipositor; VII, 7th sternite ; 7p, 7th pleurvite ; VIII,

8th sternite. Camera lucida, x 256.

In the description of the male genital tube the nomenclature of Sharp and
Muir (10) is followed, but in the case of the female reference was made to
Packard (8), Berlese (2), Henneguy (3), and Stein (11) for terms applied to
parts homologous to those found in L. brunneus.

TECHNIQUE AND Mernops.

The technique adopted to prepare the specimens for sectioning was similar
to that employed by Awati (1). This method gave excellent results.

Fixation in Carnoy II for 24 hours; three to four hours after immersion
in the fixative the head and most of the legs and the elytra and wings were
cut off.

Dehydration in two washings of 90 per cent. alcohol, three hours ; two
washings in absolute aleohol, three hours ; and one hour in a mixture of
chloroform and absolute alcohol.

The object was next transferred to chloroform, in which it remained for
two days; then into a cold saturated solution of paraffin-wax (58°C.) in
chloroform, where it remained for two to three days. After this period the
solution was warmed by standing it on top of the oven for 10 minutes, and
the object was then transferred into pure molten wax (58° C.) for five to six

hours,

GENITAL SYSTEM OF LYCTUS BRUNNEUS. 583

A block was prepared in the usual way. The sections were stained in
Hhrlich’s Hematoxylin (? hr.) and counter-stained in Picro-nigrosin

(30 secs.).
Mate Genitau System.

The reproductive organs are shown in Pl. 81. fig. 1, and consist of the
usual parts : testes (fs), vasa deferentia (vd), seminal vesicles (sv), accessory
glands (ag), ejaculatory duct (¢), internal sac, median lobe (ml), and tegmen,
comprising the lateral lobes (//) and basal piece (4p).

There are six testicular follicles of unequal size attached to each vas
deferens. They occupy a large area in the abdomen from the dorsal to the
ventral surface, and extend from beyond the basal margin of the abdomen to
the third sternite (Pl. 31. fig. 5). The testicular follicles are white, rounded
and tapering towards their apex. There isa slight cavity in the centre of
the base of each, whence a tube arises which joins the vas deferens, which is
of variable thickness throughout its length. In repose the vasa deferentia
are convoluted, in parts superimposed, and lie on the ventral surface. The
vasa deferentia enter the seminal vesicles on their outer ventro-lateral margin
(PI. 31. fig. 4). The seminal vesicles are joined on their inner lateral surface
for the greater part of their length; they are subcylindrical and rounded at
their apices.

Arising from each seminal vesicle on its inner dorsal margin is a short
tube ; these tubes meet and form the ejaculatory duct. Superimposed and
attached to the dorsal surface of each of the two short tubes is an accessory
gland. The accessory glands are subeylindrical and curved, broadest
towards their base and rounded at their apices.

The ejaculatory duct gradually tapers from its base until it enters the
median lobe through the median foramen (m/f), where it becomes hidden from
view, emptying into the internal sac (7s, Pl. 31. fig. 3). (It was not found
possible to evaginate the internal sac, nor to kill a pair of beetles in coition
with the organs in situ ; but the internal sac was traced in sections of an
immature male.)

The chitinized median lobe, viewed in section, is circular at its base and
shortly becomes elliptical ; towards its apex there arise on the upper dorsal
and upper ventral surfaces a carina situated along the centre, with the
internal sac lying over and on either side of the carina formed on the upper
ventral surface (Pl. 31. fig. 2); the lower or outer ventral surface is convex
at this point. The median orifice (mo, Pl. 81. fig. 3) is situated a little below
the upturned point of the median lobe. Inside the median lobe are series of
muscles and nerve-fibres (P]. 31. fig. 2); one series of muscles is situated
around the internal sac, others are attached to the walls of the lobe.

Encasing the median lobe on either side are the lateral lobes, which are
roughly semicircular in section for the greater part of their length; they are

42*

584 MR. A. M. ALISON ON THE

of a uniform thickness of chitin except towards their base, where the inner
face becomes thin and flexible. Hach lateral lobe contains a group of muscles
and nerve-fibres extending to their apices (Pl. 31. fig. 2). At their base the
lateral lobes become joined together ventrally and dorsally ; at the latter
point they join around the point of articulation (pa). Externally the
median lobe and the lateral lobes bear sensory pits at their apices.

Enclosing the base of the lateral lobes is a thin piece of chitin, the basal
piece (6p), which appears to clip the dorsal edges of the lateral lobes below
the point of articulation. Arising from the surface of the basal piece
towards its posterior margin is the flexible enveloping integument of the
abdomen.

In sections of a male the spermatozoa were found in bundles, held
together at their heads. In balsam mounts of the dissected genitalia these
bundles could also be seen within the testicular follicles when the organs had
been stained in heemalum and indigo-carmine.

Between the dorsal pygidial plate (dp) and the odeagus is a chitinous tube
(ev) which completely envelops the anus (an) and anterior portion of the
rectum (7) (Pl. 81. figs. 2 & 3). Hach side of the chitinous tube (ev)
invaginates to form a pair of stout rods (v7, fig. 3). These rods surround
the cedeagus and abut at their base, which is towards the ventral surface
of the abdomen. From the base of the rods there are three primary sets of
muscles (mj. Two of these sets follow the course of the rods and terminate
around the chitinous tube; the third set runs between the rods and is attached
to the enveloping integument below the base of the cedeagus ; some of these
muscles appear to be attached to the @deagus ventrally.

From this it would appear that the latter muscles are primarily responsible
for giving the cedeagus its necessary downward tilt preparatory to coition.

Following the alimentary tract back from the anus (an, PI. 81. fig. 5) is
the rectum (7), which is quite short and lies dorsal to the cedeagus and
ejaculatory duct. Beyond the rectum is the colon (cl) lying dorsal to the
accessory glands, part of the ejaculatory duct, and the seminal vesicles.
Beyond the colon is the ileum, which disappears below the testicular
follicles and lies towards the ventral surface of the abdomen. ‘There are six
malpighian tubes (mp) arising from the ileum, which at this point is lined
with chitinized setee. Beyond the ileum is the mesenteron (ms), which lies
in the meta- and mesosternum, where it inclines towards the dorsal surface.

Arising from between the metathoracic acetabula (mta) isa “U ”-shaped,
strongly chitinized apodeme (ap); the base of the ‘‘ Y” arises from the meta-
sternum (qg), whilst the apices of the arms are held in position by muscles
attached to the metanotum. rom inside the “ Y ”-piece and from its base
arise two thin chitinous rods (apr). These rods traverse the floor of the
metasternum and terminate above the metathoracic coxe (mec), to which
they are attached by radiating muscle-fibres,

GENITAL SYSTEM OF LYCTUS BRUNNEUS. 085

A pair of glands or sacs (sg) disappearing beneath the cedeagus (oe) are
shown (PI. 31. figs. 1,3, & 5). hese glands unite and form a short narrow
duct. When dissected out of a mature beetle the glands and duct arealways
found distended. The duct terminates abruptly between the 7th and 8th
sternites. In section these glands are seen to be composed of cells containing
very large nuclei. The function of the glands is not known ; no odour other
than that of the wood from which the beetles emerged was ever perceived.
The glands are apparently pygidial glands (Berlese 2).

FemMaLe GeniraL System.

The reproductive organs of the female (Pl. 32. fig. 1) consist of two
ovaries (lo, ro), two oviducts (od), a paired valve (vl), a spermatheca (sp) and
spermathecal gland (spg), and a common oviduct (co) running into the
sheaths of the “ ovipositor,’ which terminates in the bursa copulatrix. In
addition there is a long flexible cloacal stalk (lr) (‘‘ Kloakstiel,” Stein 11)
and two short rods (s7).

Tpxt-FIa. 2.

A, the ovary of an immature female. B, the ovary of a mature unfertilized female,
semi-diagrammatic. c, calyx; ec, egg-chamber; ef, ovarian’ tube; f, filament;
tc, terminal chamber; ve, ripe eggs. Camera lucida, x 54.

The ovaries (Jo, ro) of a mature beetle each consist of fourteen ovarian
tubes (e#). Hach tube consists of from four to five egg-chambers (ec), and a
terminal chamber (to) terminating in a filament (/). The ovarian tubes
arise from an enlargement of the oviducts—the calyx (c). The calices act
as receptacles for the ripe eggs, which pass into them from the basal egg-
chambers. In fig. 1 (Pl. 82) there are three ripe eggs in the left calyx and
two in the right.

It was found that the left ovary was always developed earlier than the right.

oO
f=)

we

586 MR. A. M. ALTSON ON THE

The ovary of an immature female examined consisted of only ten ovarian
tubes (text-fig. 2, A), the other four eventually arising from the apex of the
calyx and between the existing tubes. The ovary of a mature unfertilized
female (text-fig. 2, B) shows a considerable enlargement of the calyx, which
is tightly packed with ripe eggs.

From the calices the eggs pass along short oviducts into a double-chambered
valve (vl). Hach chamber of the valve is lined with brown chitinized setee *
with their free ends lying in the same direction as that in which the eggs
pass (vl, Pl. 32. fig. 4). The chambers of the valve meet to form the
commencement of the common oviduct, and the sets terminate some distance
before a valvular duct (vt) leading into the spermatheca enters the common
oviduct.

The spermatheca is very large and terminates in a single tube—the
spermathecal or accessory gland. In fertilized females the size of the
spermatheca is perceptibly increased. Inside the spermatheca, arising from
the ventro-lateral surface at the point (psp) where it narrows, are found a
series of 27 chitinized sete (s, Pl. 32. fig. 3); their function is obscure.
The spermatozoa are mostly in packets—spermatophores (spp, Pl. 82. fig. 4),
—hbut a considerable number are found loose in the fluid contents of the
spermatheca.

The common oviduct, into which the eggs pass from the valves, is of
muscular structure, and finally disappears into the “ ovipositor.”

The long flexible cloacal stalk (dr, Pl. 32. fig. 2) arises from a point in the
centre and near the base of the ventral pygidial plate (vp). The stalk
extends through the middle of the abdomen into the meta-thorax to a point
beyond the beginning of the “ovipositor,’ where its apex enlarges into a
knob from which two sets of muscle-fibres (m!, m?) arise. The m! set of
muscles consists of series of bundles of long fibres grouped around the
“ovipositor,’ and extending from the apex of the cloacal stalk to a point
between the short rods (sr), where they are attached to the outer sheath (ots).
The m? muscles surround the cloacal stalk to its base, where it is embedded

?

ina mass of muscles. In addition to the muscles (m!, m?), there is another

“ ovipositor ”’ and com-

series (m’), which arise around the beginning of the
pletely surround the outer sheath, extending for about half its length, at
which point they are attached to it. Further, there are numerous muscle-
fibres arising from the same point as the (m°) muscles; these are found
longitudinally traversing the inside of the inner sheath (ins) of the “ ovi-
positor ” (PI. 384. figs. 4-7). And, in addition, there is a series of cancellate
muscles lining the walls of the distal portion of the “ ovipositor,” apparently

* Tn a previous paper, “On the Method of Oviposition and the Hoe of Lyctus brunneus
Steph.” (Journ, Linn, Soc., Zool. xxxv. 1923, p. 217), the writer has stated that it is these
seta-lined valves which impart the striations and process to the deposited egws.

I } ] 88

GENITAL SYSTEM OF LYCTUS BRUNNEUS. 587

to allow for its expansion during the passage of an egg. Another series of
muscles (m*) are found, arising from the lateral walls of the spermatheca and
terminating around the common oviduct at a short distance from its base.
The common oviduct enters the ‘“ ovipositor ” between the m' muscles.
When the ovipositor is in repose—as depicted in Pl. 32,—it consists of
two sheaths, the outer or basal portion (ots), which is a transparent tube
lightly chitinized, and the inner sheath or distal portion (ins), which is a con-
tinuation of the outer sheath. The two portions join at the point where the
common oviduct enters. The surface of the distal portion of the “ ovipositor ”

TEXT-FIG. 3.
Tr — an.

if

\
(

Apex of the ovipositor, ventral. an, anus; Jc, bursa copulatrix; bp, basal piece;

?
fr, forked rod; p, vaginal palp; rd, chitinized rod; vo, vaginal orifice. Camera
lucida, x 128.

is covered with reeumbent brown chitinized setee, which extend as far as the
bursa copulatrix (dc, text-fig. 3). The apex of the “‘ ovipositor ” (text-fig. 3)
is stoutly chitinized, and consists of paired rounded basal pieces (op), between
which is the vaginal orifice (vo). Arising from each of the basal pieces is a

588 MR. A. M. ALTSON ON HE

double-jointed vaginal palp (p), which bears a number of sensory pits. The
inner lateral faces of the palps are concave. Within each basal piece is a
short forked rod (fr), and articulating between the prongs of the fork and
running back some distance is a chitinized rod (rd) (“ Seitenstuck,” Stein 11).

The abdominal nerve-ganglion and commissures are not depicted ; the latter
were only traced to the venter of the spermatheca and to the valve.

The left top corner of PI. 82. fig. 1 depicts a portion of the mesenteron (ms)
terminating in the ileum (/m), from which arise six malpighian tubes (mp).
The ileum, which is lined with chitinized sete at its junction with the
mesenteron, is followed by the colon (cl), which leads into the rectum (7).
The rectum, which is of muscular structure, is visible for a considerable
length before it—like the common oviduet—disappears into the ‘‘ovipositor.”
(In this figure the rectum has been pulled across to the left; it should le
looped over the “ ovipositor” and enter from the right as in PI. 33. fig. 2.)

PI. 34 depicts the ‘‘ ovipositer” in section. Figs. 4, 5, 6, and 7 are camera
lucida drawings of transverse sections from an “immature” female. Fig. 4
shows the rectum (7) entering the inner sheath (ins) of the ovipositor from
the right and assuming a position dorsal to the common oviduct (co), which
enters from under the left. In fig. 5 the rectum (7) is now dorsal to the
common oviduct (co), and both are completely enclosed by the inner sheath
or distal portion (ins) of the “ ovipositor,” which in turn is enclosed within
the outer sheath (ots). In fig. 6 the rectum (7) is seen emerging through the
dorsal wall of the distal portion of the ‘‘ovipositor”’; whilst in fig. 7 the
rectam and anus (an) are now free of the inner sheath, which, at this point,
becomes the integumental covering of the apex of the common oviduct (co)
or bursa copulatrix.

Fig. 8 is a transverse section through the apex of the ‘ ovipositor” of a
fertilized female which had begun egg-laying. The section is about the
beginning of the vaginal orifice (vo) ; it shows the stout outer lateral wall of
the basal pieces (bp) and transverse muscles to allow of expansion.

Figs. 1, 2, and 3 are partially reconstructed camera lucida drawings of
longitudinal sections of the “ ovipositor”’ of a fertilized female that had been
ovipositing. Fig. 1 shows the junction (jo) of the inner and outer sheaths,
and the entry of the common oviduct from underneath to the ventral surface
of the inner sheath and the position of the rectum. Fig. 2 is of a median
section of the “‘ovipositor”’ showing the rectum convoluted (7c). Fig. 3 is
asection through that part of the “ ovipositor ” where the rectum (7) emerges
through the inner sheath.

The relative positions occupied by the genitalia and alimentary tract within
the abdomen, meta- and mesothorax are depicted in Pl. 83. figs. 1, 2, and 3.
No muscles are shown, as their inclusion would obliterate a large part of
these organs. It will be seen that the female possesses an apodeme (ap) of a
similar structure to that described in the male. The figs. 1 and 3 are based

JENITAL SYSTEM OF LYCTUS BRUNNEUS. 589

on reconstructions made from a series of longitudinal sections of a fertilized
female, and depict a left and right lateral view respectively, with the upper-
most ovary and oviduct removed in each case. Fig. 2 is based on recon-
structions made from balsam mounts of cleared specimens and from dissections,
and depicts a dorsal view in which the relatively greater length of the rectum
(r) of the female in comparison with that of the male (Pl. 81. fig. 4, 7) is
made apparent. The ratio is approximately 30: 1.

Pl. 82. fig. 2 shows in longitudinal section the dorsal (dp) and ventral (vp)
pygidial plates with the apex of the “ ovipositor” between, and also part of
the 7th sternite (VII), ¢. e. the 5th sternite of systematists.

The functioning of the parts of the ovipositor appear to be as follows :—

In repose (PI. 32. fig. 1) m!, m?, and m* muscles are in a state of con-
traction, whilst m® and the longitudinal series within the inner sheath are in
a state of expansion. By the contraction of m%, the outer sheath commences
to collapse, the distal portion to issue from between the pygidial plates and
drawing with it the muscular rectum and the common oviduct. By the
simultaneous contraction of the muscles within the inner sheath, the outer
sheath is caused to evaginate and issue from between the pygidial plates.
The outer sheath continues to evaginate and issue until brought in check by
m! muscles. When fully extruded the “ovipositor” is approximately the
length of the female.

The withdrawal of the ovipositor is apparently achieved by the contraction
of the expanded m* muscles, aided by the muscular common oviduct, which
would pull back into its position of repose, bringing the distal portion of the
“ovipositor”’ with it. Simultaneously, the contraction of m! muscles would
pull back the basal part of the outer sheath, whilst m? muscles would draw
into position the apical portion of the sheath.

As there are no muscles attached to the ileum, the colon, or the rectum,
the resumption of the position of repose for this portion of the alimentary
tract must be solely dependent upon the slight contraction of which the
muscular rectum is capable—aided by the fixed mesenteron—and any
tension that might be exerted by the ramifications of the trachea. That such
is apparently the case is shown in Pl. 34. fig. 2. Here the rectum is seen
convoluted—a condition that can only be attributed to an imperfect
musculature.

Other sections of the same beetle disclosed two other points within the
“ovipositor”” where the rectum was similarly convoluted. In this condition
no beetle could defecate unless it extended its “‘ovipositor,’ and thus
straightened the rectum. Whether this is ever doneis not known. But as the
food of these beetles consists of particles of wood, most of which appears in
the frass never to be broken down, it is possible that penetration of the walls of
the rectum and of those of the common oviduct may take place. And it seems
probable that this defectiveness may be the cause of the death of quite a

590 MR. A. M. ALTSON ON THE

number of beetles whose ovaries are found to contain a considerable portion
of their complement of eges. Jt has been observed that many beetles do

expire with the “ ovipositor” extruded—partially in the majority of cases.

When the distal portion of the ovipositor is gently pulled out by means of
a fine pair of forceps from a freshly chloroformed beetle, the basal portion is
seen to extend after the manner of a telescope.

Females have a gland on each side of the seventh abdominal segment cor-
related to the glands or sacs deseribed in the male; and they are of similar
structure. But in the females they do not meet, and from sections they
appear to surround the tracheal trunks leading to the seventh abdominal
spiracles.

Corrion.

It has been mentioned that efforts to kill pairs in coition with the organs
in situ failed to disclose more of the internal sac of the male than is shown
in Pl. 81. fig. 2. And from the examination of the aedeagus the internal sac
does not appear to be capable of evaginating more than half the length of the
median lobe (Sharp & Muir 10).

In coition the beetles are superimposed, and the lateral lobes of the male
assume an almost perpendicular position to the surface of the wood. As the
beetles are very flat, the apices of the lateral lobes come in contact with the
surface of the wood, and, if the female moves, as is frequently the case,
the apices of the lateral lobes become bent and covered with particles of dirt
and dust *.

Novres ON THE GENITAL System or Lyrorvs rrvearis GOEZE.

An attempt has been made to carry out a comparative examination of the
genital system of L. linearis. But no living specimens could be procured,
and the following remarks are based upon the study of preparations made
from the remains of the organs of dead beetles.

Two beetles—both males—were kindly sent by Dr. Hugh Scott, and three
other specimens—one female and two males—were found in the pupal
chambers in oak sent from Paris by M. P. Lesne. The oak was sent under
the impression that it was infested by the larvee of this species ; however, as
no beetles emerged within the normal period, it was broken up and no larvee
or pupee were found.

The dead beetles obtained were first soaked for five days in water to which a
tew drops of formaldehyde were added. ‘he specimens were then washed in
water and dissected. The dissections were treated in the usual manner and
mounted in balsam. The resulting preparations were moderately successful.

* From a control point of view, this is of considerable importance, for the treatment of
the surface of the wood with an oleaginous or viscous substance causes the dirt and dust
to adhere to such an extent as to jam the cedeagus—the lateral lobes hecome stuck to the
median lobe, the median oritice becomes blocked, and the beetle is, as a result, killed.

GENITAL SYSTEM OF LYCTUS BRUNNBEUS. 591

One point observed was that the tergites of linearis were much more
strongly chitinized than in brunneus, in which they are very flexible and only
faintly discernible.

The male—The cedeagus was found to agree with the description and
figures of Sharp and Muir (10), except that the basal margin of the basal
piece was found to terminate in front of the point of articulation, but not so
far forward as in drunnews. Whether the anterior margin of the basal piece
clipped around the lateral lobes was not observed, as no sections were made.
In other respects it was similar, and gave the impression of being a more
robust structure,

Nothing was found of the testes, but the position and appearance of the
remains of the vasa deferentia, seminal vesicles, accessory glands, and ejacula-
tory duct were similar to those of brunneus.

The two rods arising from under the ejaculatory duct as in brunneus, and
the ““U”-shaped apodeme and apodemal rods were present.

The female.—The similarity of the extruded “ ovipositor” as seen in dead
females to that of brunneus has been referred to by others. From the
preparation made, there were observed the setae-lined valve and what would
be the spermatheca and common oviduct. Remains of the ovaries were
present, but the number of ovarian tubes could not be distinguished. The
ovipositor, in repose, and the apex of the distal portion bore a marked
similarity to those of brunneus. Further, the cloacal stalk and short rods, and
the apodeme and apodemal rods were present.

From the appearance of the “ ovipositor ’
alimentary tract must follow a similar course to that of brunneus.

’ in repose, it seems that the

OBSERVATIONS ON STEIN’s ‘ FumaLE REPRODUCTIVE ORGANS IN
COLEOPTERA.’

Stein (11), in his work on the genital system of Coleopterous females,
describes and illustrates the organs of several species which in some par-
ticulars resemble the organs of brunneus. But this resemblance is so slight
in most that no comparative deductions can be drawn. ‘There are, however,
a few points that call for comment.

The “ ovipositor ” in each of these cases consists of a tube (“ Kloakrohr ”’)
which in repose telescopes approximately one-half within the other, but each
varies in the appearance of the distal portion. In all cases the ‘ ovi-
positor” is appareutly shorter, in comparison to the length of the insect, than
that of brunneus and linearis. The rectum, in each of the species referred to,
enters the inner sheath of the ‘
stalk is found, but it is not clear what distance within the abdomen it

‘ovipositor ” as in brunneus. The cloacal

traverses, and in no case is there any indication of the short rods.
In the case of those females that are closely related systematically to the
Lyetids, i.e. Anobium punctatum De Geer (pertinaa F., domesticum Fourc.,

92 “MR. A. M. ALTSON ON THE

Or

striatum Ol.), and Cis bolett Scop.*, the cloacal stalk, according to Stein, is
dorsal to the “oyipositor,”’ and it is therefore in a reversed position in
brunneus and linearis. But in the case of other females somewhat distantly
related systematically, such as Helops (caraboides Panz.) striatus Foure.,
(Hdemera (virescens) lurida Marsh, and Cistela fusca, the cloacal stalk,
according to the same authority, lies below the ‘ ovipositor,” and, as in
brunneus and linearis, it arises from the venter.

Tt might also be observed that in Anobium punctatum De Geer, Stein
describes two accessory glands—in the female, of course. One is unpaired
and the other paired, the former lying ventral to the “ovipositor” and the
latter pair apparently arising from within the inner sheath of the “ ovipositor.”
Pygidial glands, such as have been found in both sexes of brunneus, are not

mentioned by Stein.

Nore on THE MorpxoLogy or THE APEX OF THE ABDOMEN IN
Lycrus pruywevus STEPH.
This note is based upon a close study of the sections previously referred to,
and of a series of balsam mounts of pupze in various stages of development,
as well as various preparations made from the beetles.

TextT-rie. 4,

an :

Apex of the abdomen of three pupz of Z. brunneus from specimens prepared and mounted
in balsam. A, ventral aspect of a female, early stage of development; B, dorsal
aspect of the same, seen through the back of the slide. OC, ventral aspect of a female,
later stage of development; D, dorsal aspect of the same, seen through the back of the
slide. K, ventral aspect ofa male, late stage of development; I*, dorsal aspect of the
same, seen through the back of the slide. am, anus: yen, genitalia; yo, gonopore ;
7p, seventh pleurite; 7-10, tergites; VII-X, sternites. All figures camera lucida
outlines, x 66.

* The opportunity to examine specimens of Cis boleti Scop. has recently occurred, and the
cloacal stalk was found to be ventral to the ovipositor, as in brwnneus.

GENITAL SYSTEM OF LYCTUS BRUNNEUS. 593

Text-fig. 4 depicts the ventral and dorsal aspects of the apex of the abdomen
of three pupz. From these it is apparent that the external appendages of
the genitalia in both sexes arise from a similar position, 7.e.on the ninth
sternite (IX) -and towards the tenth; the gonopore (go) lies medianly between
these appendages, and is situated on the ninth sternite.

In the female L. brunneus the last visible sternite—the fifth of systematists—
is the true seventh sternite (VIL) and corresponds with the seventh tergite (7)
directly above it (Pl. 38. figs. 1 & 3). The pygidium represents part of the
eighth segment with the cloacal stalk (/r) an invagination of the eighth
sternite (VIII), and the paired rods (sr) are lateral invaginations of the
eighth tergite (8). This conclusion is supported by the presence of the m!
and m? series of muscles (Pl. 32. fig. 1), which would seem to be modified
intersegmental muscles. For the position of attachment of the m! set is from
the apex of the cloacal stalk to lateral positions near the beginning of the
ninth seement—the basal portion of the “ ovipositor” ; whilst the m7 set,
which envelop the cloncal stalk, arise from the point of its invagination.
Moreover, a part of the muscles enveloping the paired rods (sr, P1].32. fig. 1)
are attached laterally to the base of the ninth segment, whilst the remainder
of these muscles are attached to the point of invagination of each rod.

The “ ovipositor ” is, therefore, a double tubular prolongation of the ninth
seoment, tergites and sterniles being indistinguishable. Whether the trans-
parent and setze-less basal portion of the “ovipositor” represents a pro-
longation of the “soft membrane” between segments 8 and 9, is not
manifest. Interseegmental membrane is not ‘manifest anywhere else. And,
conversely, with regard to the setze-covered distal portion of the “ ovipositor,”’
sete: are not manifest on any tergite but the seventh.

The apex of the ovipositor bears the anus dorsally between the fused and
reduced tenth tergite and sternite, and it carries the genitalia on the ninth
sternite beyond the anus.

In the male (Pl. 81. figs. 2 & 3) the last visible sternite is the seventh, as
in the female ; and likewise the pygidium represents the eighth seement
The ninth tergite and sternite are not solidly chitinized, and are invaginated
and fused at their beginning to form the lumen enclosing the cedeagus and
the anus, the latter being enveloped in a tube (ev, fig. 3) formed by thie
fused tenth tergite and sternite, as in the female. The invaginated ninth
seement, the sternal portion of which is invaginated more than the tergal
portion, shortly evaginates. Then the tergal portion joins the tergal portion
of the tube [ev (10), fig.3]; whilst the sternal portion, in which the gonopore
is situated medianly, eventually joins the sternal portion of the tube (X,
fig. 2). The ninth sternite is prolonged around the gonopore to form the
extrusible membranous tube carrying the cedeagus, to which it is attached, as
previously stated, around the basal piece. The paired rods (vr, figs. 1 & 3)
that project into the abdomen from the tube (ey) are invaginations of the

594 MR. A. M. ALTSON ON THE

tenth segment, and the three sets of muscles associated with these paired rods
are also intersegmental muscles. The lateral sets following the course of the
rods represent the muscles of the tenth sternite and tergite, whilst the median
set represent those of the tenth and ninth sternite.

The homology between the genitalia of the sexes seems apparent (text-
figs. 3& 4 and Pl. 31. figs. 1 & 3). The sensory-pitted unjointed lateral
lobes homologize with the sensory-pitted jointed vaginal palps; the basal
pieces correspond ; and the evaginated median lobe and extrusible internal
sac homologize with the internal forked rods—* seitenstuck””—and invaginated
uterus. The analogy of their functions is obvious.

In the larva of L. brunneus there are nine pairs of spartans 2 : one prothoracie
and eight abdominal pairs. In the adult there are nine pairs of spiracles :
one prothoracic, one metathoracic—on the soft integument below the wing-
bases—and seven abdominal pairs.

SUMMARY.

Both sexes in L. brunneus are sexually mature when they have emerged
through the wood from their pupal chambers. An external sex-character is
described.

In the male the reproductive organs consist of two testes of twelve
pyriform testicular follicles of unequal size, six arising from each convoluted
vas deferens, which terminate in a pair of jointed subecylindrical seminal

vesicles. A short tube arises from each aa vesicle, and forms the
ejaculatory duct which empties into the wdeagus. Partly superimposed over
the seminal vesicles and emptying into the short tubes are the paired
accessory glands. They are subeylindrical, slightly tapering, and are curved
around to underneath the ejaculatory duct. There is a pair of pygidial
glands.

In the female the reproductive organs consist of two ovaries, each com-
posed of fourteen ovarian tubes with the nutritive cells situated at their
apices. The ripe eggs accumulate in a calyx which continues to form the
oviduct. The oviducts terminate in a chitinized setee-lined paired valve
which forms the source of the common oviduct. At this point the voluminous
spermatheca arises; it terminates in a tubular spermathecal or accessory
gland. The common oviduct disappears into the invaginated double tubular
‘“ovipositor”” at a point.in the metathorax, where the rectum also enters.

In the testicular follicles the spermatozoa are in bundles, in the sperma-
theca they are in packets or spermatophores.

The “ ovipositor ” has been found to consist of a double tubular prolongation
of the ninth abdominal segment, with a reduced tenth segment towards the
apex. When extruded the ‘ovipositor” is approximately the length of
the beetle. Situated at the apex of the “ovipositor” are the genitalia with
the anus a short distance behind.

GENITAL SYSTEM OF LYCTUS BRUNNEUS. 595

Arising as an invagination of the eighth sternite and extending into the
metathorax is the long, flexible, and strongly-chitinized cloacal stalk, from
which arise systems of muscles for exserting and withdrawing the ovipositor.
These muscles are found to be modified intersegmental muscles.

Morphologically the gonopore in both sexes was found to arise medianly
on the ninth sternite, and the homology of the genitalia appears.

In the larva of LZ. brunneus there are nine pairs of spiracles, one thoracic
and eight abdominal ; in the adult there are also nine pairs, two thoracic and
seven abdominal.

The movement which the rectum of the female is subjected to has been
observed to be injurious to this organ, in that, lacking any musculatory
system for its withdrawal, other than its muscular structure, it was found
convoluted inside the “ovipositor” of an egg-laying specimen. ‘his
defectiveness must accelerate death.

' ACKNOWLEDGMENTS.

The economic investigation, of which this paper records part of the results,
was suggested by Prof. H. Maxwell-Lefroy, Imperial College of Science, to
whom the writer has to express his thanks, and to the Committee of the
Scientific Industrial Research Dept. for a grant to carry on the work.

The writer is also indebted to Dr. C. J. Gahan, Keeper of the Department
of Hntomology, British Museum (Nat. Hist.), for identifying specimens
of L. brunneus ; to Dr. Hugh Scott, Cambridge Museum, for specimens of
I. linearis ; and to M. P. Lesne, Mus. Nat. d’Hist. Nat., Paris, for very
kindly sending some oak thought to be infested with L. linearis.

In addition, thanks are due to Dr. A. D. Imms, Rothamsted Experimental
Station, for his advice and for his assistance in connection with the publi-
cation of this paper ; and to Prof. Stewart MacDougall for his efforts to have
the original paper published as a whole.

BIBLIOGRAPHY.

1, Awatr, P. R. (1914). —‘ Mechanism of Suction in Lygus pabulans”: pp. 685-733.
Proe. Zool. Soc. London.

2. Brrurse, A. (1909 et seq.).—‘ Gli Insetti.’ Milan.

3, Henneavy, L. F. (1904).—‘ Les Insectes.’ Paris.

4. Kmrsuaw, J. C., & F. Murr (1922).—“The Genitalia of the Auchenorhynchus
Homoptera”: vol. xv, No. 3, pp. 201-212. Am. Ent. Soc. America.

5. Mra, L. C., & A, Denny (1886),—‘ The Structure and Life-history of the Cockroach.’
London.

6. Murr, F. (1915).—“ Notes on the Ontogeny of the Genital Tubes in Coleoptera”:
pp. 147-152, pl. 12. Psyche, i. Boston, U.S.A.

"7, Murr, F. (1918).— Notes on the Ontogeny and Morphology of the Male Genital Tube
in Coleoptera”: pp. 225-229. Trans. Ent. Soc. London.

8. Packarp, A, S. (1909).—‘ A Text-book of Entomology.’ New York.

596 MR. A. M. ALTSON ON THE

9. Saarp, D. (1918).—“Studies in Rhynchophora”: pp. 209-222. Trans. Ent. Soe.

London.

10. Suarp, D., & I’. Murr (1912).— The Comparative Anatomy of the Male Genital
Tube in Coleoptera”: pp. 477-641, pl. 63. figs. 132 & 132a.
Trans. Wnt. Sec. London.

11, Srnin, F. (1847).—‘ Vergleichende Anatomie und Physiologie der Insekten, I. Mono-
eraphie. Ueber die Geschlechtsorgane und den Bau des Hinterleibes
bei den weiblichen Kifern’: pp. 139, 9 pls. Berlin.

EXPLANATION OF THE PLATES.
PratTE 31,

Fig. 1. Male genitalia. Semi-diagrammatic. ezea 51.
Fig, 2. Transverse section (slightly oblique) through apex of abdomen of immature male.
x 100.
Fig. 8. Right lateral aspect of apex of male abdomen. 114.
Fig. 4, Diagrammatic ventral aspect of seminal vesicles, accessory glands and their junction
with branched ejaculatory duct. x crea 51. |
. Diagrammatic dorsal aspect of genitalia and alimentary tract in situ. Greatly

Fig. 5
magnified.
PLATE 32.
Fig. 1, Female genitalia (camera lucida) and part of alimentary tract (partly diagrammatic).

x 51.

Fig. 2. Longitudinal section through apex of female’s abdomen. x 384,

Fig. 3. Transverse section through spermatheca. C.l. X 384.

Fie. 4. Longitudinal section through base of spermatheca and valves of oviduct. GC 7.
to) to} o

x 384.
PLATE 33.

Arrangement im situ of genitalia and alimentary tract of female.
Partly reconstructed from sections and partly diagrammatic. Greatly magnified.
Fig. 1. Left lateral view. Position of repose.
io. 2. Dorsal view. Position of repose.
Fie. 8. Right lateral view. Position when ovipositor extruded.

PLATE 34.

]. Longitudinal section through ovipositor at junction of inner and outer sheaths.
2, Longitudinal section through middle of ovipositor, showing rectum’ convoluted.
Fig. 3. Longitudinal section through ovipositor, showing rectum and common oviduct
separating.
Fig. 4. Transverse section of ovipositor at junction of inner and outer sheaths.
Fig. 5. Transverse section of middle of ovipositor.
Fig. 6. Transverse section of ovipositor, showing rectum and common oyiduet separating.
Fig. 7. Transverse section of ovipositor showing anus free, and common oviduct completely
enclosed by inner sheath.
Fig. 8. Transverse section through bursa copulatrix,
(Figs. 1, 2, and 3 are of a mature fertilized female. Part (./. and partly reconstructed.
Fies. 4, 5, 6, and 7 are of an immature female, C./, Fig. 8 is of a fertilized female,
0.1, All figures x 840.)

ALTSON. JOURN. LINN. SOC., ZOOL. VOL. XXXV. PL. 31.

GENITAL SYSTEM OF LYCTUS.

ae

ALTSON. JOURN. LINN. Soc., ZOOL. VOL. XXXV. PL. 32.

GENITAL SYSTEM OF LYCTUS,

JOURN. LINN. Soc., ZOOL. VOL. XXXV. PL. 33.

ALTSON.

H
'

a
S
5S

GENITAL SYSTEM OF LYCTUS.

JOURN. LINN. Soc., ZOOL. VOL. XXXV. PL. 34.

ALTSON.

INS =-=————— ==

GENITAL SYSTEM OF LYCTUS.

GENITAL SYSTEM OF LYCTUS BRUNNEUS. OOK

REFERENCE LETTERING.

ag, accessory gland; age, junction of accessory gland and ejaculatory duct; an, anus;
asy, aperture of secreting glands; ap, apodeme ; apr, rod of apodeme; bc, bursa copulatrix ;
bop, basal portion of ovipositor; bp, basal piece ; c, calyx; cl, colon; co,common oviduct ;
em, commissure ; d@, dorsal; dop, distal portion of ovipositor; dp, dorsal pygidial plates (8th
tergite); dr, dorsal rod; ec, egg-chambers ; ej, ejaculatory duct; e/, elytron; ef, ovarian
tubes; ev, enveloping sheath; f, filament; 2, integumental sheath; rs, inner sheath of
ovipositor; zs, internal sac ; jej, junction of ejaculatory duct; jo, junction of outer and inner
sheaths of ovipositor; 7/, lateral lobes; /m, ileum; Jo, left ovary; Jl, cloacal stalk; Zs,
loose spermatozoa; m, muscles; m'—-m', sets of muscles; ma, mesothoracic acetabula;
mec, mesothoracic coxa; mf, median foramen; md, median lobe; mo, median orifice ;
mp, malpighian tubes; ms, mesenteron ; ms/, mesothoracic leg; mta, metathoracie aceta-
bula; mtec, metathoracic coxa; m#l, metathoracic leo; mv, suture between meso- and
metasternum; 2, abdeminal nerve-ganglion; od, oviduct; oe, cedeagus; op, 2-jointed
vaginal palp of ovipositor; ots, outer sheath of ovipositor; ovp, ovipositor; p, palp;
pa, point of articulation ; pl, prothoracie leg; pr, point of extrusion of anus; psp, sete
inside spermatheca ; g, metasternum ; ge, episternum of metasternum; 7, rectum ; re, con-
voluted rectum; ro, right ovary; s, sete; sy, secreting glands; sp, spermatheca; spg,
spermathecal gland ; spp, packets of spermatozoa (spermatophores) ; s7, short rod; s¢, branch
of ejaculatory duct ; stm, stomodeum ; sv, seminal yesicle ; ¢c, terminal chamber; ts, testes ;
v, ventral; vd, vas deferens; vo, vaginal orifice; vp, ventral pygidial plate (VIII sternite) ;
vr, ventral rods; vf, valvular duct; w, wing; 1-10 tergites, I-X sternites.

LINN. JOURN,— ZOOLOGY, VOL, XXXV, 43

re
ints

THE FORAMINIFERA OF LORD HOWE ISLAND. 599

The Foraminifera of Lord Howe Island, Scuth Pacific.
By H. Heron-Atren, F.R.S., F.L.S8., and A. Hartann, F.R.M.S.

(Prarrs 35-37.)

| Read 21st June, 1923.]

Tue highly interesting material which forms the subject of the present

Monograph was collected by Prof. R. Douglas Laurie during the visit of the

British Association to Australia in 1914. We may quote his own description,

which is as follows :—‘It consists of material taken from between tide-

mark at ‘ Middle Beach’ on the east side of the Island, which is the exposed
side, 7. e. without a coral reef. There is a little coral to be seen growing on

this sbore at low tide, but on the other—on the west side of the Island—is a

coral reef quite well developed and protecting a lagoon, abont a mile from

the land.

“‘Middle Beach, observation spot; Lat. 31° 31’ 30’ 8., Long. 159° 15’
28” H.

“ Lord Howe Island is the most southerly Pacific Island with a coral reef.
Norfolk Island, though about 2° 30' further north, has none, nor is there one
on the Australian Pacific coast at similar latitude.

“The whole of the material was collected from one small beach ;.from
two different levels. 58 6, hardened coral sand and weed from rock-pools at
lower half of the beach; 58, loose surface coral sand to an approximate
depth of + inch, from three-quarter high-tide line.”

Prof. Laurie sent us a preliminary sample, with a view to ascertaining the
value of the material, from 58 c, and, this haying promised excellent results,
he subsequently sent us seven bottles, which may be described as follows :—

No. 1. In 70°/, alcohol. This had never been dried. (58 0.)

CS es a This had been previously dried at 50° C. ‘These
yielded about 10 cc. of * floatings.” Much byssus and organic
material. Many species in the finer siftings, and notable specimens of
Nubecularia bradyi, N. schauinslandi, and Discorbina polystomellordes

”

in the coarser grades.
4, A small bottle containing Orbitolites only; ranging from minute speci-
mens up to individuals 6 mm. in diameter. Dried without heat. (55 4.)
Similar material in 70 °/, alcohol. Never dried. (58/.)
6. A single dried specimen of Orbitolites complanata, with the monstrous
rectangular secondary outgrowth. (From 58 6.)

ct

43*

600 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

No. 7. The residue of the preliminary sample from 58.c, dried without heat. This
material was coarse, and yielded practically no “ floatings,” the Fora-
minifera being a good deal worn. It consisted largely of Orbitolites
and Tinoporus (Baculogypsina spherulata) with about twenty other
species of robust types, among which Amphistegina lessonii, Textularia
agglutinans, and its ally Haddonia torresiensis were prominent.

Perhaps the most notable feature of the gatherings was the presence of
two genera new to science, Mifusilina and Craterites, which are fully
described and figured in this monograph.

Another very noteworthy and interesting feature was the profuse occur-
rence of species in the reproductive stages, both by viviparity and ‘* budding.”
We have called attention, in our notes, to Nubecularia lucifuga, Spirillina
campanula (sp. nov.), Discorbina tabernacularis, Gypsina inherens, and others,
containing young brood in the cavity of the test, resulting from the al)sorption
of the internal septa. Beyond this, certain species (e.¢., Bulimina elegan-
tissima) exhibited unquestionable evidence of the reproduction by “ budding ”
from the aperture, described at length and illustrated by Heron-Allen in the
Phil. Trans. of the Royal Society (H-A. 1915, RPF.).

Among the 199 species and varieties described from the material sub-
mitted to us, seven are new to science. Perhaps the most noteworthy
species, beyond these, are Nubecularia schawnslandi, Miliolina kerimbatica,
M. stelligera, Fischerina pellucida, India diaphana, Haddonia torresiensis,
Frondicularia scottii, Uvigerina selseyensis, Ramulina grimaldv, and Discorbina
reniformis.

We may be allowed to say that we have never had material submitted to
us which has been more carefully collected and preserved for examination.
Our only regret is that it should have been collected from reefless areas only.
No complete study of the foraminiferal fauna of the Island can be made
under such reservation. The material shows that a large proportion of the
specimens are not of local origin, but are more or less water-worn shells
which have travelled some distance. The more perfect specimens, of local
origin, are, in the case of a number of species, rather small and pauperate,
probably evidencing existence under difficulties, due, no doubt, to paucity of
food on the exposed side of the Island. A collection made on the eastern
side of the Island, among the reefs and in the still waters of the Lagoon,
would probably have yielded a much longer list.

The general facies of the foraminiferal fauna is, of course, sub-tropical
and of the Indo-Polynesian type. Many of the species recorded have a wide
range, from Hast Atrica, through the Malay and Australasian Seas, to the
Pacific. Perhaps the most significant feature is the marked rarity of certain
genera which might reasonably have been expected to occur in abundance,
e.g., Hauerina, Peneroplis, Alveolina, Polytrema, Operculina, Heterostegina,
and so on, and the entire absence of many typical species which we should
have expected to find in such latitudes and in such conditions.

THE FORAMINIFERA OF LORD HOWE ISLAND. | 601

Sub-Kingdom PROTOZOA.
Class RHIZOPODA.
Order FORAMINIFERA.
Family MILIOLIDA.
Sub-family NuBECULARIINA.
Nuprcuraria Defrance.
1. Nupecunaria LuciruGa Defrance. (PI. 35. fig. 1.)
Nubecularia lucifuga Defrance, 1825, Dict. Sci. Nat. vol. xxxv. p. 210; Atlas
Zooph. pl. 44. fig. 3.
a ses Brady, 1884, FC. p. 134, pl. 1. figs. 9-16.
ie ne Cushman, 1910, ete., FNP. 1917, p. 41, pl. 8. fig. 6.
Frequent, but not very typical, displaying a tendency to run into straight
lines of chambers. In one specimen the broken final chamber shows what

oO.
5

is apparently a young individual fitting closely into the conformation of the
chamber. The young specimen consists of four inflated chamberlets arranged
on a rotaline plan. The surface appears to be pitted, but not perforate.
Texture very thin and delicate.

2, NUBECULARIA BRADYI J/illett.
Nubecularia inflata Brady, 1884, FC. p. 135, pl. 1. figs. 5-8.

3 bradyi (nom. nov.) Millett, 1898, etc., FM. 1898, p. 261, pl. 5.
fig. 6.
» Heron-Allen & Harland, 1914-15, FIA. p. 550, pl. 40.

figs. 8-10.
Abundant and very variable.

3. NUBECULARIA SCHAUINSLANDI (Khumbler). (PI. 35. figs. 2-5.)
Miliolina schauinslandi Rhumbler, 1906, FLC. p. 41, pl. 3. figs. 20, 21.
Quinqueloculina ,, Cushman, 1910, etc., FNP. 1917, p. 56, pl.8. figs. 7, 8

(after Rhumbler).

Frequent and very variable. The test, so far as external appearances go,
starts with a perfectly normal and well-developed individual comparable
with Miliolina rotunda or M. labiosa, followed by a straight or curving series
of irregularly formed chambers terminating in a large, gaping, irregular
orifice, narrowed down by a variable number of projecting teeth. It appears
to be nothing more than a strongly-developed Nubecularia bradyi, and is
certainly much more closely allied to that form than to the milioline series of
the initial chambers. The more regularly shaped specimens suggest Vevillina
coronata (Millett), but are very different from Sidebottom’s specimens
(Manchester Mem. vol. xlix. 1905, No. 11), the types of which are

602 MESSRS. EB. HERON-ALLEN AND A, EARLAND ON

in our collection. The points of difference lie in the extremely regular
disposition of the chambers in Nevillina, and particularly in the distinctive
crown-like aperture. Millett’s original type-specimen of Biloculina coronata,
which is also in our collection, is, on the contrary, much more likely to be
referable to Nubecularia schawinslandi than to Nevillina coronata, its only
point of identity with Wevllina being the fact that the projecting teeth
round the aperture actually meet and coalesce instead of remaining distinct
and separate as in Wubecularia schawinslandi. Otherwise the whole structure
and appearance of the specimen is distinctly nubecularine and has no feature
in common with Nevillina coronata.

Sub-family MILIOLININ &.
Brrocunina ad? Orligny.

4, BILOCULINA RINGENS, var. STRIOLATA Brady.
Biloculina ringens, var. striolata Brady, 1884, FC. p. 148, pl. 3. figs. 7, 8.
Millett, 1898, etce., FM. 1898, p. 262, pl. 5.

0? ” ”
fig. 8.
e fs is Heron-Allen & KEarland, 1914-15, FKA.
p. 551.
Rare. The specimens agree entirely with Brady’s description and figure.

5. Brnocutina sarst Schlumberger.
Biloculina sarst Schlumberger, 1891, BGE. p. 166, pl. 9. figs. 55-59, text-
figs. 10-12.
i » Cushman, 1921, FP. p. 471, pl. 97. fig. 1, text-fig. 48.

Rare, The specimens are fairly large and distinctive.

6. BrLocuLINA DEPRESSA dd? Orbigny.
Biloculina depressa @Orbigny, 1826, TMC. p. 298, no. 7; Modéle, no. 91.
Brady, 1884, FC. p. 145, pl. 2. figs. 12, 15-17; pl. 3.
figs. 1, 2.

09 ”
Very rare.

7. BILOCULINA DEPRESSA, var. SERRATA Bailey.
Biloculina depressa, var, serrata Bailey, 1861, New Spp. Mier. Org. Para River,
S. America. Boston Journ. Nat. Hist.
vol. vii. p. 350, pl. 8. fig. E.
Cushman, 1910, etc., FNP. 1917, p. 75, pl. 29.
fig. 2.
A single specimen, the serration being confined to the aboral half of the

shell.

THE FORAMINIFERA OF LORD HOWE ISLAND. 603

8. BinocuLina ELONGATA d’ Orbigny.
Biloculina elongata WVOrbigny, 1826, TMC. p. 298, no. 4.
Brady, 1884, FC. p. 144, pl. 2. fig. 9.
3 » Heron-Allen & Harland, 1918, CL. p. 22, pl. 1. fig. 4.

Very rare, but fairly typical.

oy) ”

9. Binocutina opposrra Deshayes.
Biloculina opposita. Deshayes, 1831, CCT. p. 252, pl. 3. figs. 8-10.
Bronn, 1837, LG. p. 1143, pl. 42. fig. 30.
(Eneyel. Methodique, vol. 11. p. 138, fide Broun.)
5 oblonga d’Orbigny, 1839, FC. p. 1638, pl. 8. figs. 21-28.

Three small but distinctive specimens which agree remarkably well with
Deshay es’ figures, except as regards the tooth, which is not strongly fureate
as in his specimens, but less prominent as in d’Orbigny’s B. oblonga, which :
is, in other respects, identical with Deshayes’ earlier figure. ‘This form may
be regarded as an abbreviated and inflated variety of B. elongata.

39 ”

SPIROLOCULINA d’ Orbigny.

10. SPIROLOCULINA ANTILLARUM @? Orbigny.
Spiroloculina antillarum VOrbigny, 1839, FC. p. 166, pl. 9. figs. 3, 4.
Brady, 1884, FC. p. 155, pl. 10. fig. 21.
55 55 Heron-Allen & Harland, 1908, etc., SB. 1911, p. 301.
Abundant. One of the most typical forms in the gatherine. The final
pair of chambers is usually so turgid as almost to enclose all the preceding

” ”

chambers.

11. SPIROLOCULINA CANALICULATA d’ Orbigny.
Sptroloculina canaliculata dOrbigny, 1846, FFY. p. 269, pl. 16. figs. 10-12.
Jones, Parker, & ee 1866, ete., MCF. p. Ae
pl. 3. figs. 39, 40.
oh 5 Cushman, 1921, FP. p. 395, pl. 80. fig. 3
Very rare and weak, and exhibiting a tendency to separation of the
chambers, so as to leave lacunae between the adjacent chambers, as in

bby Ee)

S. acutimargo Brady.

12. SPrroLOcULINA PLANULATA (Lamarck).
Miliolites planulata Lamarck, 1804, AM. p. 352, no. 4.
Spiroloculina ,, Brady, 1884, FC. p. 148, pl. 9. fig. 11.
3 » Heron-Allen & Karland, 1914-15, FKA. p. 555.
A single large specimen which on one side is plane and on the other feebly
costate, a variation referred to in FIA. (vt supra).

604 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

=~

13. SPrROLOCULINA TENUIROSTRA KKarrer. (PI. 35. figs. 6, 7.)
Spiroloculina tenwirostra Karrer, 1867, FO. p. 308, pl. 2. fig. 5.
acutimargo Brady, 1884, FC. p. 154, pl. 10. fig. 14 (only).
- % Egger, 1893, FG. p. 222, pl. 1. figs. 26-28.

A single specimen, identical with Hgger’s figure of S. acutimargo Brady.
Under this name Brady figures three distinctive forms. Fig. 12 is clearly
S. afica Terquem. Figs. 13 and 15 are what we may regard as S. acutimargo
proper. Hig. 14 is a quite distinctive little shell which Cushman (C. 1921,
FP. p. 398), under S. acutemargo, attributes to the genus Masslina. It is the
same as Hoger’s figure and is in our opinion attributable to S. tenuirostra
Karrer, the distinctive features of which are a sharp angular edge to the thin
embracing chambers, almost meeting over the faces of the shell. In our
specimen the embracing character is so marked that the two final chambers
with their flanges envelop the entire face, and the separate chambers are only
visible where the surface of the test is abraded. We have observed this form

in many tropical coral sands.

”

Minrotina Williamson.

14. MrinioLina ciRCULARIS (Bornemann).
Triloculina circularis Bornemann, 1855, FSH. p. 349, pl. 19. fig. +.
Miliolina » Brady, 1884, FC. p. 169, pl. 4. fig. 3; pl. v. figs. 13, 14 (?).
“ », Heron-Allen & Harland, 1914-15, FKA. p. 557.
Abundant, and very variable in size. Both biloculine and triloculine forms,
as figured by Millett (M. 1898, etc., FM. 1898, p. 499, pl. 11. figs. 1-3), the
biloculine being relatively rare.

15. Miniotina VALvuLARIs (feuss).
Triloculina valuularis Reuss, 1851, FSUB. p. 85, pl. 7. fig. 56.
Miliolina ss Brady, 1884, FC. p. 161, pl. 4. figs. 4, 5.
‘3 re Heron-Allen & Harland, 1913, CL. p. 27.
Fairly frequent, but small.

16. Minronina pinatata (d’ Orbigny).
Quinqueloculina dilatata dOrbigny, 1839, FC. p. 192, pl. 11. figs. 28-30.
Schlumberger, 1893, MGM. p. 75, text-figs. 29, 30,
pl. 3. figs. 70-74; pl. 4. figs. 87-90.
Miliolina » Heron-Allen & Harland, 1914-15, FKA. p. 559.

Two large characteristic individuals.

39 te)

17. Miniotina LABiosa (d’ Orbigny).
Triloculina labiosa VOrbiguy, 1839, FC. p. 178, pl. 10. figs. 12-14.
Miliolina », Brady, 1884, FC. p. 170, pl. 6. figs. 3-5,

a », Mallett, 1898, ete., FM. 1898, p. 502, pl. 11. figs. 8, 9.

Rare, but typical.

THE FORAMINIFERA OF LORD HOWE ISLAND. i 605

18. MinioLina suBRoTUNDA (Jontagu).
Vermiculum subrotundum Montagu, 1803, TB. pt. 2, p. 521.
Miliolina subrotunda Brady, 1884, FC. p. 168, pl. 5. figs. 10, 11.
Frequent and small.

19. Miniouina seminupa (Reuss).
Quinqueloculina seminuda Reuss, 1866, FABS. p. 125, pl. 1. fig. 11.
Miliolina subrotunda (Montagu), var., Wright, 1885-6, BLP. p. 319, p. 26. fig. 5.
> senvnuda Heron-Allen & Harland, 1914-15, FIA. p. 560.
Not uncommon, variable in the number of strize, which tend to spread over
the entire surface of the shell, thus linking the species with I. webbiana

(d’Orb.).
20. Mrmionina wEBBIANA (d? Orbigny).
Triloculina webbiana VOrbigny, 1839, FIC, p. 140, pl. 3. figs. 13-15.
Maliolina fichtehana Brady, 1884, FC. p. 169, pl. 4. fig. 9.
5 suborbicularis Millett, 1898, etc., FM. 1898, p. 502, pl. 11. fig. 13.
es webbiana Heron-Allen & Harland, 1914-15, FKA. p. 560.
Rare.
21. Minronina TRIGONULA (Lamarck).
Mitiolites trigonula Lamarck, 1804, AM. vol. y. p. 351, no. 3.
Triloculina ,, _ d@Orbigny, 1826, TMC. p. 299, pl. 16, figs. 5-9; Modeéle,
no. 93.
Miliolina i Brady, 1884, FC. p. 164, pl. 3. figs, 14-16.
Frequent. All the specimens are of the very elongate ty pe, comparable
with MW. affins (d’Orb.) (F. 1905, SOM. p. 59, pl. 1. fig. 2).
22. MInioLina TRICARINATA (d? Orbigny).
Triloculina tricarinata VOrbigny, 1826, TMC. p. 299. no 7; Modéle, no. 94.
Miliolina * Brady, 1884, FC. p. 165, pl. 5. fig. 17.
Triloculina s Cushman, 1921, FP. p. 454, figs. 35, 36.

Rare. Both long and short forms occur.

23. MILIOLINA BERTHELINIANA Brady.
Miliolina bertheliniana Brady, 1884, FC. p. 166, pl. 114. fig. 2.
9 x Heron-Allen & Harland, 1914-15, FKA. p. 563,
pl. 41. figs. 32-35.
Very rare. Both long and short forms occur.

24. Minionina cuttRaTa Brady.
Miliolina eultrata Brady, 1879, etc., RRC. 1881, p. 45.

55 Bs Brady, 1884, FC. 161, pl. 5. figs. 1, 2.
st i Heron-Allen & Harland, 1914-15, PKA. p. 564, pl. 42.

figs. 1-10.
A single very long and delicate individual.

606 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

25. MILIOLINA BOSCIANA (a Orbigny).
Quinqueloculina bosciana d’Orbigny, 1839, FC. p. 191, pl. 11. figs. 22-24.

Miliolina ue Millett, 1898, ete., FM. 1898, p. 267, pl. 6. fig. 1.
5 Ap Heron-Allen & Karland, 1914-15, FKA. p. 566.

Small, but not uncommon.

26. MinioLiIna opLtonea (Montagu).
Vermiculwn oblongum Montagu, 1803, TB. p. 522, pl. 14. fig. 9.
Milrolina oblonga Brady, 1884, FC. p, 160, pl. 5. fig. 4.
* ea Heron-Allen & Harland, 1914-15, FIA. p. 566.
Not uncommon. Both the square and the rounded types occur, the
rounded predominating. One of the latter is characterized by a long stopper-
like tooth, such as we noted from Ulare Island (H.-A. & H. 1913, OL. p. 25).

27. MrnioLina RorunDA (d’ Orbigny).
Triloculina rotunda VOrbigny, 1826, TMC. p. 299, no. 4.
Miliolina 33 Millett, 1898, etc., FM. 1898, p. 267, pl. 5. figs. 15, 16.
Me ., Heron-Allen & Harland, 1914-15, FKA. p. 568, pl. 42.
figs. 27-30.
Two distinct forms occur, one moderately frequent of the triloculine type
figured by Millett (wt supra), the other of the distinctive d’Orbignyan type,
large, thick-shelled, and finely striate. :

28. MILIOLINA VULGARIS (dOrbigny).
Quinqueloculina vulgaris V@Orbigny, 1826, TMC. p. 302, no. 35.
.3 re Schlumberger, 1893, MGM. p. 65, pl. 2. figs. 65, 66,
& woodcuts 13, 14.
Miliolina vulgaris Heron-Allen & Karland, 1914-15, FKA. p. 569.
Large, typical, and frequent.

29. MILIOLINA SEMINULUM (Linné).
Serpula seminulum Linne, 1767, SN. p. 1264, no. 791.
Miliolina ,, Brady, 1884, FC. p. 157, pl. 5. fig. 6.
a es Heron-Allen & Earland, 1914-15, FKA. p. 569,
Common. Varying greatly in size and oceasionally reaching very large
dimensions.

30. MinioLina (TRILOCULINA) L&VIGATA (d? Orbigny).
Triloculina levigata V@Orbigny, 1826, TMC. p. 300, no. 15.

= 44 Terquem, 1878, FEP. p. 57, pl. 5. (10) figs. 20-21.
99 os Schlumberger, 1893, MGM. p. 205, pl. 1. figs. 45-47.
3 5 Heron-Allen & Harland, 1922, WGA. pl. 1. figs. 11-14.

Two quite typical specimens. The surface-texture is rather rough.

THE FORAMINIFERA OF LORD HOWE ISLAND. 607

31. MILIoLINA TRIANGULARIS (d’ Orbigny).
Quinqueloculina triangularis VOrbigny, 1826, ‘IMC. p. 302, no. 34; 1546,
: FEV. p. 288, pl. 18. figs. 7-9.
Miliolina 3 Jones, Parker, & Brady, 1866, ete., MFC. 1895,
p. 118, pl. 4. fig. 1, and pl. 6. figs. 2, a—b.

Typical specimens. Rare.

32. MiILioLINaA AUBERIANA (d? Orbigny).
Quingueloculina auberiana @Orbigny, 1839, FC. p. 193, pl. 12. figs. 1-3.
Miliolina - Brady, 1884, FC. p. 162, pl. 5. figs. 8, 9.
55 35 Heron-Allen & Harland, 1914-15, FKA. p. 571.

Common and generally small. Large and typical specimens are rare.

33. MILIOLINA AUBERIANA, var. SEMIRBTICULATA, noy. (PI. 35. figs. 8-10.)
Having the characteristic form of J/. auberiana, but with the peripheral
margins of the chambers reticulated. Cushman’s M. kerimbatica, var. philip-
pinensis (C. 1921, FP. p. 438, pl. 89. figs. 2, 3) is a similar form, but the
periphery and median wing are rounded instead of sharp as in JZ. wuberiana.

34, Mrnionina cUVIERIANA (d’ Orbigny).

Quinqueloculina cuvieriana @Orbigny, 1839, FC. p. 190, pl. 11. figs. 19-21.
Brady, 1884, FC. p. 162, pl. 5. fig. 12.
Heron-Allen & Earland, 1914-15, FKA. p. 571,

pl. 42. figs. 33-36.

Miliolina “i

” ”

One weak specimen.
35. Mettotina Bicostava (d’ Orbigny).
Quinqueloculina bicostata @Orbigny, 1839, FC. p. 198, pl. 12. figs. 8-10.
Miliolina bicostata Goés, 1894, ASF. p. 112, pl. 20. fig. 855.
: Heron-Allen & Harland, 1914-15, FKA. p. 572, pl. 42.
figs. 42-45.
Large and well-developed and fairly frequent. The specimens differ from
the Kerimba individuals in the aperture, which is normal, round, and furnished
with the usual tooth, agreeing in these respects with the figure of Goés.

” ”

36. Mrtrotina unvosa (Karrer).
Quinqueloculina undosa Karrer, 1867, FO. p. 361, pl. 3. fig. 3.
Miliolina wndosa Brady, 1884, FC. p. 176, pl 6. figs. 6-8.
Heron-Allen & Harland, 1914-15, FKA. p. 572, pl. 43.

figs. 1-4.
Frequent and typical.

oF by)

37. Minrotina uNDULATA (a Orbigny).
Quingueloculina undulata d’Orbigny, 1826, TMC. p. 302, no. 27.
Schlumberger, 1895, MGM. p. 71, pl. 1. figs. 53, 54;
pl. 2. figs. 60, 615; text-figs. 23, 24.

29 ”

608 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

Miholina undulata Heron-Allen & Harland, 1914-15, FKA. p. 573, pl. 43.
figs. 5-8.
Very rare and rather small, but quite typical.

38. Mintouina reTicuLatTa (d’ Orbigny).
Triloculina reticulata dOrbigny, 1826, TMC. p. 299, no. 9.
Miliolina 5 Brady, 1884, FC. p. 177, pl. 9. figs. 2-4.
Heron-Allen & Harland, 1914-15, FKA. p. 573, pl. 43.
figs. 9, 10.
Frequent and large, of the quinqueloculine rounded type, with coarse

29 9

markings.

39. Minronina PARKERI Brady.
“Quinqueloculina with oblique ridges,” Parker, 1858, MIS. p. 53, pl. 5. fig. 10.
Miliolina parker’ Brady, 1884, FC. p. 177, pl. 7. fig. 14.
Heron-Allen & Harland, 1914-15, FKA. p. 574, pl. 48.
figs. 11, 12.

Very rare, but large and typical specimens.

bh) De)

40, Minionina KERIMBATICA Heron-Allen Harland.
Miliolina kerimbatica Heron-Allen & Earland, 1914-15, FKA. p. 578, pl. 43.
figs. 13-23.
Quingueloculina ,, Cushman, 1921, FP. p. 437.

A single large and typical individual.

41. Mrnrouina Fusca Brady.

Quingueloculina fusca Brady, 1870, FTR. p. 286, pl. 11. fig. 2.
Schulze, 1874, etc., R. 1875, p. 134, pl. 6. figs. 19, 20.
Heron-Allen & Harland, 1914-15, FKA. p. 576.

Rare, but large and well-developed.

”

Miliolina by

42, MinioLina contorra (ad Orbigny).
Quinqueloculina contorta d’Orbigny, 1846, FFYV. p. 298, pl. 20. figs. 4-6.
Miliolina nS Goés, 1894, ASF. p. 111, pl. 20. figs. 851, 852.
Cushman, 1921, FP. p. 432, pl. 90. fig. 1.

29 29

Frequent and large.

43, MrILioLIna FERUSSACI (d’ Orbigny).
Quingueloculina ferussacii @Orbigny, 1826, TMC. p. 301, no.18; Modéle,
no. 32.
Miholina "5 Brady, 1884, FC. p. 175, pl. 118. fig. 17,
pa nA Heron-Allen & Karland, 1914-15, FIA. p. 578.
Frequent, and all of the feeble smooth type figured by Millett (M. 1898,
etc.. FM. 1898, p. 508, pl. 12. fig. 7 [fig. 6 in text ]).

THE FORAMINIFERA OF LORD HOWE ISLAND. 609

44, Mrnionina LINNEZANA (d Orbigny).
Triloculina linneiana d’Orbigny, 1839, FC. p. 172, pl. 9. figs. 11-13.
Miliolinu inneana Brady, 1884, FC. p. 174, pl. 6. figs. 15-20.
Heron-Allen & Harland, 1914-15, FKA. p. 579.

+} ”

A few very large and typical specimens.

45. MInroLina STRIATA (da? Orbigny).
Quinqueloculina striatu @Orbigny, 1826, TMC. p. 301, no. 4.
i » Lerquem, 1882, FEP. p. 184, pl. 20. (28) figs. 10-12.
Miliolina » Heron-Allen & Earland, 1914-15, FKA. p. 579, pl. 44.
figs. 13-17.

Rare ; very small and weak.

46. MinroLina sTELLIGHRA (Schlumberger).
Quinqueloculina stelligera Schlumberger, 1893, MGM. p. 68, pl. 2. figs. 58, 59.
Miliolina Pf Heron-Allen & Harland, 1913, CI. p. 31, pl. 1.
figs. 14, 15; 1916, FWS. p. 215, pl. 39, figs. 28-31.

Rare, but typical.

Sub-family HAUERININ&.
Haverina da’ Orbigny.
47, Haumrina compressa a’ Orbigny.
. Hauerina compressa @’Orbigny, 1846, FFV. p. 119, pl. 5. figs. 25-27.

Brady, FC. p. 190, pl. 11. figs. 12, 13.
a 3 Heron-Allen & Harland, 1914-15, FKA, p. 588.

A single undoubted specimen.

” ”

48. HAUERINA ORNATISSIMA (Aarrer).
Quingueloculina ornatissema Karrer, 1868, MFKB. p. 151, pl. 3. fig. 2.
Hauerina 53 Brady, 1884, FC. p. 192, pl. 7. figs. 15-22.
5 s Heron-Allen & Harland, 1914-15, FKA.-p. 590.

Two rather poor specimens, not developed to the outspreading stage.

PLANISPIRINA Seguenca.

49, PLANISPIRINA AURICULATA Myger.
Planispirina auriculata Nigger, 1893, FG. p. 245, pl. 3. figs. 13-15.
Heron-Allen & Harland, 1914-15, FKA, p. 590,
pl. 46. figs. 3-7.
Rare, but the two forms differing in relative length and breadth oecur,
as described in our Kerimba Monograph (ut supra).

3” »”

610 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

Sub-family FrscHERININ &.
FIscHERINA Terquem.

50. FiscHEeriIna PELLUCIDA JZillett.
Lischerina pellucida Millett, 1898, etc., FM. 1898, p. 611, pl. 13. figs. 14, 15.
Pr i Heron-Allen & Harland, 1914-15, FKA. p. 591.
One excellent and typical specimen.

Sub-family PENEROPLIDIN&.
Cornuspira Schultze.

51. CoRNUSPIRA INVOLVENS Reuss.
Operculina involvens Reuss, 1849-50, FOT. p. 370, pl. 1. (46) fig. 20 (not 30).
(Cornuspira, 1861; Model, no 15.)
Cornuspira » {Brady, 1884, FC. p. 200, pl. 11. figs. 1-3.
3 * Heren-Allen & Earland, 1914-15, FKA. p. 593.
Minute, but not infrequent. Both megalo- and microspheric forms, the
first predominating as usual.

52. CORNUSPIRA SELSEYENSIS Heron-Allen § Harland.
Cornuspiwra ? Harland, 1905, FBS. p. 199, pl. 13. figs. 2-4.
Cornuspira selseyensis Heron-Allen & Earland, 1908, etc., SB. 1909, p. 319,
pl. 15. figs. 9-11.
Small and rare.

Prneropuis Alontfort.

53. PENEROPLIS PERTUSUS (Morskal).
Nautilus pertusus Forskal, 1775, Descriptio animalinm, p. 125, no. 65.
Peneroplis ,, Brady, 1884, FC. p. 204, pl. 13. figs. 16, 17.
in $3 Heron-Allen & Harland, 1914-15, FKA. p. 601.

Very rare and extremely pauperate in structure.

54, PENEROPLIS CARINATUS d’ Orbigny.
Feneroplis carinatus V’Orbigny, 1839, FAM. p. 33, pl. 3. figs. 7, 8.
‘a = Brady, 1884, FC. p. 205, pl. 13. fig. 14.
Ms i Heron-Allen & Harland, 1914-15, FICA. p. 602.

Also rare, but less pauperate than the pertusus type.

OrprroLirEs Lamarck.

re

55. ORBITOLITES DUPLEX Carpenter.
Orbitolites duplea Carpenter, 1856, ete., RF. 1856, p. 220, pl. 5. fig. 10;
pl. 9. fig. 10,
bf 3 Brady, 1884, FC. p. 216, pl. 16. fig. 7.
20 os Heron-Allen & Karland, 1914-15, FKA. p, 605.

THRE FORAMINIFERA OF LORD HOWE ISLAND. 611

Comparatively infrequent as compared with O. complanata. A few speci-
mens exhibit a crumpled development of the marginal rows of chambers,
approaching the structure of Orbitolites complanata (Marginopora vertebralis) ,
var. plicata Dana (1846, in Wilkes’s U.S. Exploring Exp. vol. vii. p. 706,
pl. 60. fig. 9), which is better known under Brady’s varietal name, var.
laciniata, but there is no indication that these specimens represent a repro-
ductive process as in Dana’s variety.

56. ORBITOLITES COMPLANATA Lamarck.

Orbitolites complanata Lamarck, 1802, Syst. Anim. sans Vert. p. 376.

e ‘ Brady, L884, EC. p. 218, pl. 16. figs. 1-6; pl. 17
figs. 1-6.
a Pa Heron-Allen & Harland, 1914-15, FKA. p. 606.

Hxtremely common, but never reaching any great dimensions. No
specimens of var. plicata or viviparous individuals were observed.

Crarerites Heron-Allen & Harland.
CRATERITES, gen. nov.

This new genus is founded for the reception of a single specimen, which,
while evidently closely connected with Orbitolites, cannot by any stretch of
generic definition be included with hitherto recorded types. The closest
search through the material available has not resulted in the discovery of
other specimens or of anything which might be regarded as an earlier or
more advanced condition of growth, and we feel compelled, against our usual
practice, on such very insufficient data, to institute a new genus for what is
unqestionably a novel and interesting type of construction. Future research
may render it necessary to amplify or amend our description, or may possibly
relegate the specimen to the position of a freak ora pathological develop-
ment of Orbitolites complanata.

57. CRATERITES RECTUS, sp. noy. (Pl. 35. figs. 11, 12.)

The type-specimen is free, but the base, which is roughly quadrigonal in
form, suggests that it was once sessile on some other object. The basal layer
consists of a nubecularine mass of chambers without a trace of spiral disposition.
Arising from this outspread basal layer is a thick trunk nearly cireular in
section, composed of five or six superimposed rings of chamberlets, orbito-
line in appearance, but devoid of marginal pores. From this point the trunk
rapidly increases to nearly double its diameter at its narrowest point by the
addition of further superimposed rings of orbitoline chambers without mar-
ginal pores. The greatest diameter of the test is reached at a point which is
about twelve layers of chambers above the rim of the base. From this
widest point the upper surface of the test is completed by a thin and highly
conyex cover, which bears no trace of septation, but is entirely covered with

612 MESSRS. E. HERON-ALLEN AND A. EBARLAND ON

coarse perforations, like the rose of a watering-can. Where this oral layer,
which closely resembles the marginal edge of Orbitolites complanata, has
been broken away at a point on the edge, a similar cribrate septum is visible
underneath.

The specimen bears a superficial resemblance to the Mycetozoan genus
Craterium.

The genus represents a morphological variation of the normal structure of
Orbitolites complanata. In that type the chamberlets form ‘a disc, the sides
of which are imperforate, while the edge of the dise is covered with cribrate
apertures. In Craterites the disc is originally perforate on one side and
imperforate round the edge. Growth ensues by the addition of a series of
chambers, which must necessarily be superimposed over the aperture. Hence
we get a series of discs superimposed on one another and with a circular cribrate
covering over all in Craterites, while in Orbitolites the original dise increases
rapidly in diameter by the addition of rings of concentric chamberlets,

Size :—
Height from edge of base to top of cover, 1°20 mm.
5¢ as bottom of cover, *70 mm.

Diameter at base, 95 mm.
narrowest part of trunk, 55 mm.
widest part of trunk and junction of cover, ‘95 mm.

2?

ed

ALVEOLINA d’ Orbiany.
58. ALVEOLINA MELO (Fichtel § Moll).

Nautilus melo Fichtel & Moll, 1798, TM. p. 118, pl. 24.
Alveolina ,, Heron-Allen & Karland, 1914-15, FKA. p. 607.

A single large specimen.

Family ASTRORHIZID As.
Sub-family ASTRORHIZIN”®.
Tripia Heron-Allen § Harland.

59. IrntpIA DIAPHANA Heron-Allen §° Harland.

Tridia diuphana Heron-Allen & Harland, 1914-15, FKA. 1914, p. 371, pl. 36;
1915, p. 607.

The species is evidently abundantly attached to coral and_nullipore
fragments. Several good specimens were obtained, both sessile and detached,
the latter exhibiting the dried protoplasmic body beneath the chitinous
membrane,

THE FORAMINIFERA OF LORD HOWE ISLAND. 613

Denpropurya Strethill Wright.

60. DENDROPHRYA RADIATA Strethill Wright (?).
Dendrophrya radiata Strethill Wright, 1861, Ann. & Mag. Nat. Hist. ser. 3,
vol. viii. p. 122. (No figure.)
2 ,, Brady, 1884, FC. p. 238, pl. 27a. figs. 10-12.
Two large fragments which we ascribe to Dendrophrya, to which they bear
a closer resemblance than to any other species. They represent fragments
of an organism constructed of irregularly branching flattened tubes, formed
of fine brown sand deposited upon a chitinous membrane. The interior of
the tubes is more or less rough (semi-labyrinthic) with projecting material.
The internal calibre of the tube is very large, the walls being comparatively
thin. In the absence of further material we are unable to assign these frag-
ments to any of the recorded species, and hesitate to give them a new name.
It may be that our organism represents fragments of Cushman’s suggested
species Dendrophrya ramosa or its var. robusta. (C. 1921, FP. p. 56, pl. 18.
fios. 7, 8.)
Sub-family RHABDAMMININ &.
SAGENINA Chapman.

61. SAGENINA FRONDESCENS (Brady).
Sagenella frondescens Brady, 1879, etc., RRC. 1879, p. 41, pl. 5. fig. 1.
Sagenina ss Chapman, 1899, FFA. p. 4, pl. 1. figs. 1, 2; pl. 2. figs. 1, 2.
ue 3 Heron-Allen & Harland, 1914-15, FKA. p. 611.

Large and typical colonies attached to small stones.

Family LITUOLIDA.
Sub-family LirvuoLin &.
HAPLOPHRAGMIUM Reuss.

62. HAPLOPHRAGMIUM COMPRESSUM (rods.
Lituolina irregularis, var. compressa Goés, 1882, RROS. p. 141, pl. 12.
figs. 421-423.
Haplophragmium emacratum Brady, 1884, FC. p. 305, pl. 33. figs. 26-28.
compressum Goés, 1896, DOA. p. ol.
Heron-Allen & Harland, 1914-15, FKA. p. 613,
pl. 46. figs. 20, 21.

A single small and coarsely agglutinate specimen.

”

” ”

63. HAPLOPHRAGMIUM CANARIENSE (d’ Orbijny).
Nonionina canariensis d’Orbigny, 1839, FIC. p. 128, pl. 2. figs. 33, 34.
Haplophragmium canariense Brady, 1884, FC. p. 310, pl. 35. figs. 1-5.
Heron-Allen & Marland, 1914-15, FKA. p. 614.
E 44

tk) thd

LINN. JOURN.—ZOOLOGY, VOL, XXXVo

614 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

Common, and attaining large dimensions. Characterized by a thin white
regular test, deeply ferruginous, and very neatly constructed of fine sand
with a very considerable proportion of sponge-spicules.

64, HAPLOPHRAGMIUM ANCEPS Brady.

Haplophragmium anceps Brady, 1884, FC. p. 3138, pl. 35. figs. 12-15.
Millett, 1898, etce., FM. 1899, p. 361, pl. 5. fig. 10.
a » Heron-Allen & Harland, 1918, CI. p. 47, pl. 3. fig. 4.

Two small, but typical specimens.

” ”

PLacopsinina d’ Orbigny.
gn!

65. PLACOPSILINA CENOMANA d’ Orbigny.
Placopsilina cenomana d’Orbigny, 1850, ete., PP. vol. ii. p. 185, no. 758.
Lituola (Placopsilina) cenomana Carpenter, Parker, & Jones, 1862, ISF. p. 148
pl. 11. fig. 14.
Placopsilina ,, v Brady, 1884, FC. p. 315, pl. 36. figs. 1-3.
A number of fragments agreeing in structure with this species, but all
“free.” It is possible that they may have been originally sessile upon some

perishable organism.

Drrrusinina Heron-Allen 5 Harland.

DIFFUSILINA, gen. nov.

Test sessile, squamous, composed of very finely comminuted sand and mud
enveloping a thin labyrinthie layer of chambers. External surface smooth
and finished, white to grey in colour, furnished with a few sparsely distributed
pustules of more loosely aggregated material.

The fragments of calcareous alga furnished many examples of this new
type. It is at first very difficult to recognize owing to its colour harmonizing
with that of its host. Its squamous form and habit of growing only in
depressions and crevices of the alga add to the difficulty of detecting it, but
when once recognized its presence can hardly be overlooked. It may
possibly be a widely distributed type.

The affinities of Difusilina are not easily discoverable. The neatly con-
structed test and high proportion of cement in the finished exterior surface
indicate the Lituolide, but it has no close relationship to any previously
recorded type. Wesuggest Bdelloidina as its nearest, but still a distant, ally.

66. DirrusiLINA HUMILIS, sp. nov. (PI. 35. figs. 13-16.)

Test sessile, of irregular outline, squamous, adapting its growth to
depressions and crevices on the surface of its host (difusus=poured out,
spilled). Colour resembling its host, white to grey. Hxternal surface
smoothly finished and flat; marginal edges thinned out so as to appear

THE FORAMINIFERA OF LORD HOWE ISLAND. 615

almost continuous with the surface of attachment and presenting no sign of
marginal apertures. Nearly all the specimens exhibit a varying number
(1-4) of pustular processes on the superior surface. These rise slightly
above the smoothly finished agglutinate surface, and appear to consist of the
same fine sand and mud as the rest of the test, but without agelutinating
cement. It appears probable that they represent apertures for the extension
of protoplasm.

The test when laid open is seen to contain a single layer of intricately
ramifying flat tubes, without septa, separated from one another by thin walls
of agglutinated material and from the surface of the alga by a thin floor or
pavement layer of similar material. The tubes are completely filled with a
mass of dark protoplasm. The protoplasmic body is voluminous, dark in colour,
and ramifies through the entire structure in a single layer separated from
the surface of the alga by a floor of material similar to that used in the
construction of the upper layer of the test.

Size very variable. Compact specimens 1-3 mm. in diameter, but narrow
specimens following a crevice in their growth may probably attain four or
five times this size.

Happonia Chapman.

67. HADDONIA TORRESIENSIS Chapman. (PI. 35. figs. 17-22.
Haddonia torresiensis Chapman, 1897, ‘On Haddonia, a new Genus of the
Foraminifera from Torres Straits,’ Journ. Linn. Soc.
Lond. vol. xxvi. (Zoology) (1898) pp. 452-456, pl. 28.
& text-fig. p. 453. ;
us » Heron-Allen & Harland, 1914-15, FKA. p. 616, pl. 46.
fig. 22.

Abundant and extremely variable both as regards size and relative irregu-
larity of growth. This organism, which is so abundant in this material,
appears to be unquestionably referable to Chapman’s genus ; but a study of
the material Jeads us to differ from his conclusions as to its relationships.
He placed his genus among the labyrinthic Lituoline ; we feel inclined to
transfer it to the Textulariidee, and we have even some doubt as to its generic
value. A study of a series of specimens indicates that at Lord Howe Island
growth starts as a free and regularly formed Teatularia (agglutinans or
gramen). After attaining almost full growth the chambers begin to run
wild, and they may then assume practically any form—curving or straight
lines, acervuline masses, remain free, attach themselves to large sand-grains,
or become sessile upon other organisms, but often so lightly as to become
detached without damage. ‘The shell-structure is of coarse calcareous and
siliceous grains with a large proportion of caleareous cement. Chapman
states that ‘‘the test commences either with a straight or a sinuous series of
chambers, or, more rarely, with a flat coil of a single whorl, after which the

44*

616 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

chambers are arranged in a more or less rectilinear manner.” In only one
or two instances have we met with a coiled initial series. But for the estab-
lished existence of Haddonia as a genus, and the fact that the Lord Howe
specimens generally coincide with Chapman’s diagnosis, we should have
regarded our specimens as biological freaks, due perhaps to brackish-water
conditions affecting the growth of the characteristic Tevtularia of the district,
as such conditions of life have been proved to do.

Sub-family TROCHAMMININS.
TrocHAMMINA Parker §; Jones.

68. TROCHAMMINA SQUAMATA Parker & Jones.
Trochammina squamata Parker & Jones, 1860, RMF. p. 304, table.
Brady, 1884, FC. p. 337, pl. 41. fig. 3.
Heron-Allen & LEarland, 1913, CI. p. 50, pl. 3.
figs. 7-10.
Smail, pauperate, and very rare.

99 99

” ”

69. TROCHAMMINA OCHRACEA (Williamson).
Rotalina ochracea Williamson, 1858, RFGB. p. 50, pl. 4. fig. 112; pl. 5.
: fig. 113.
Trochammina ,, Millett, 1898, ete., FM. 1899, p. 368, pl. 5. fig. 12.
Heron-Allen & Earland, 1914-15, FKA. p. 619, pl. 46.
figs, 27, 28.
One extremely pauperate individual, showing the flange described by us
from Kerimba (ut supra), and two small but typical specimens.

29 ”

70. TROCHAMMINA ROTALIFORMIS Wright.
Trochammina inflata, var., Balkwill & Wright, 1885, DIS. p. 331, pl. 138.
figs. 11, 12.
rotaliformis Heron-Allen & Harland, 1913, CI. p. 52, pl. 3.
figs. 11-13,
Frequent and well-developed.

”

71. TROCHAMMINA VESICULARIS (oés.
Trochammina vesicularis Goés, 1894, ASF. p. 31, pl. 6. figs. 235 -237.

One large and typical specimen of this high-domed type.

Carteriva Brady.

72. CARTERINA SPICULOTESTA (Carter).
Rotalia spiculotesta H. J. Carter, 1877, ‘‘ Description of a new Species of
Foraminifera (Rotalia spiculotesta),”’ Ann. & Mag. Nat.
Hist. ser. 4, vol. xx. p. 470, pl. 16; 1879, ser. 5, vol. ii.
p. 414; 1880, SGM. ser. 5, vol, v. p. 452.

THE FORAMINIFERA OF LORD HOWE ISLAND. 617

Carterina spiculotesta Brady, 1884, FC. p. 346, pl. 41. figs. 7-10.
S a Sidebottom, 1906, ete., RFD. 1905, p. 6, pl. 1. fig. 1,

A single undoubted fragment, representing a terminal chamber. The
individual when perfect must have been of considerable size.

Family TEXTULARIID A.
Sub-family Tux tuLARIIN A”.
TEXTULARIA Defrance.

73. TEXTULARIA FoLIuM Parker 5 Jones.
Textularia folium Parker & Jones, 1865, NAAF. pp. 870 & 450, pl. 18. fig. 19.

35 » Brady, 1884, FC. p. 357, pl. 42. figs. 1-5
6 », | Chapman, 1907, REV. p.127, pl. 9. tig. 4.

A single specimen.

714. TEXTULARIA INCONSPICUA, var. JuGOsA (Brady).
Tevtularia jugosa Brady, 1884, FC. p. 358, pl. 42. fig. 7.
*, inconspicua, var. jugosa Millett, 1898, etc., FM. 1899, p. 558, pl. 7.
fig. 2.
a 6 i Heron-Allen & Earland, 1908, etc.,SB.1911,
p. 310, pl. 9. fig. 12; 1914-15, FKA.
p. 624.
One typical specimen.

75. TEXTULARIA RHOMBOIDALIS Millett.
Textularia rhomboidalis Millett, 1898, etc., FM. 1899, p. 559, pl. 7. fig. 4.

bos % Sidebottom, 1904, ete, RFD. 1905, p. 8, pl. 2.
figs. 2,23
Ps Bs Heron-Allen & Harland, 1914-15, FKA. p. 624.

Very abundant and very variable, the marginal edges ranging from quite
straight to strongly scalloped, owing to the projection of the extremities of
the chambers. The median line is often deeply excavated. Small specimens
are often so compressed as to be with ditheulty separated from Boliwina.

Limbation of the sutural lines often gives a highly decorative appearance to
the shell.

76. TEXTULARIA CRISPATA Brady.
Textularia crispata Brady, 1884, FC. p. 359, pl. 113. fig. 2.
ae 5 Heron-Allen & Harland, 1914-15, FKA. p. 624, pl. 47.
figs. 5, 6.

Two specimens, one very large and quite typical, the other small.

615 MESSRS. BE. HERON-ALLEN AND A. EARLAND ON

77. TEXTULARIA SAGITTULA Defrance.
Textularia sagittula Defrance, 1824, Dict. Sci. Nat. vol. xxxii. p. 177; vol. li.
p. 844; Atlas Conch. pl. 13. fig. 5.
re 3 Brady, 1884, FC. p. 361, pl. 42. figs. 17, 18.
“i Ms Heron-Allen & Harland, 1914-15, FKA. p. 625.
Very rare.

78. 'TEXTULARIA SAGITTULA, var. FIsStuLOSA Brady.

Textularia sagittula, var. fistulosa Brady, 1884, FC. p. 362, pl. 42. figs. 19-22
3 "0 - Millett, 1898, etc., FM. 1899, p. 561, pl. 7.
fig. 9.
Also rare and small.
79. TEXTULARIA RUGOSA (Reuss).
Plecanium rugosum Reuss, 1869, FOG. p. 453, pl. 1. fig. 3.
Textularia rugosa Brady, 1884, FC. p. 625, pl. 42. figs. 23, 24.
st » Heron-Allen & Harland, 1914-15, FKA. p. 625, pl. 47.
figs. 7-9.
Large, but not very strongly marked.

80. TEXTULARIA AGGLUTINANS d’ Orbigny.
Textularia agglutinans VOrbigny, 1839, FC. p. 144, pl. 1. figs. 17, 18, 82-34.
i ze Brady, 1884, FC. p. 363, pl. 43. figs. 1-3.
ss ‘ Heron-Allen & Karland, 1914-15, FKA. p. 626.
Abundant and subject to much variation in the direction of irregular
growth, suggestive of incipient progress towards Haddonia, q. v.
81. TexTULARIA CANDEIANA d@’ Orbigny.
Teatularia candeiana @Orbigny, 1839, FC. p. 148, pl. 1. figs. 25-27.

5 % Sidebottom, 1904, ete., RFD. 1905, p. 7, pl. 2. fig. 1.
Ks x Heron-Allen & ‘Boaendl 1914-15, FKA. p. 627, pl. 47.
figs. 10-16.

Not uncommon, but not very strongly developed or typical.

82. TexTULARIA ABBREVIATA dd? Orbigny.
Textularia abbreviata d’Orbigny, 1846, FFY. p. 249, pl. 15. figs. 9-12 (not

7-12).

Mi agglutinans, var. abbreviata Parker & Jones, 1865, NAAF. p. 369,
pl. 17. fig. 76.

ws abbreviata Heron-Allen & Earland, 1922, T.N. p. 120.

One large stoutly built specimen.

83. TEXTULARIA GRAMEN a? Orbigny.
Textularia gramen WOrbigny, 1846, FFV. p. 248, pl. 15. figs. 4-6.
51 5 Brady, 1884, FC. p. 365, pl. 43. figs. 9, 10.
aM vs Heron-Allen & Earland, 1914-15, FIA. p. 627.

THE FORAMINIFERA OF LORD HOWE ISLAND. 619

Common, and exhibiting the same tendency to Haddonia-formation as
2 c=)
T. agglutinans.

84; TEXTULARIA CoNICA d’ Orbigny.
Textularia conica d@’Orbigny, 1839, FC. p. 145, pl. 1. figs. 19, 20.
A » Brady, 1884, FC. p. 365, pl. 43. figs. 138, 14; pl. 113. fig. 1.
rt ;, Heron-Allen & Earland, 1914-15, FKA. p. 629.

Common.

85. TEXTULARIA Contca, var. JuGosA Jones § Millett.
Teetularia sagittula, var. jugosa Jones; Millett, 1898, etce., FM. 1899, p. 561,
pl. 7. fig. 8.
A few small specimens comparable with Millett’s figure (ut supra), which
appears to us to be referable to 7. conica rather than to 7. sagittula.

VERNEUILINA d’ Orbigny.

86. VERNEUILINA SPINULOSA Reuss.
Verneuilina spinulosa Reuss, 1849-50, FOT. p. 374, pl. 2. (47) fig. 12.
PP 3 Brady, 1884, FC. p. 384, pl. 47. figs. 1-3.
Heron-Allen & Harland, 1914-15. FKA. p. 630.
Not uncommon. All the specimens are of the broad heavy type, destitute
of spines.

” ”

87. VERNBEUILINA POLYSTROPHA (/teuss).
Bulimina polystropha Reuss, 1845-46, VBK. pt. 2, p. 109, pl. 24. fig. 53.
Vernewilina ,, Brady, 1884, FC. p. 386, pl. 47. fig. 9.
5 < Heron-Allen & Earland, 1929, VP. (passim).

Minute and very rare.

88. VERNEUILINA PROPINQUA Brady.
Verneuilina propingua Brady, 1884, FC. p. 387, pl. 47. figs. 8-12 (not 13, 14).
ry ie Cushman, 1910, ete., FNP. 1911, p. 56, fig. 92.
A single minute but typical specimen.

Tritaxia Leuss.
89. TRIraAxIA LEPIDA Brady.
Tritaxia lepida Brady, 1879, ete., RRC. 1881, p. 55.
g » Brady, 1884, FC. p. 389, pl. 49. fig. 12.
i » Millett, 1898, ete., FM. 1900, p. 12, pl. 1. fig. 15.
Frequent.
Pavonina d@ Orbigny.
90. PAVONINA FLABELLIFORMIS d’ Orbigny.
Pavonina flabelliformis V@Orbigny, TMC. p. 260, no. 1, pl. 10. figs. 10, 11;
Modeéle, no. 56.

620 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

Pavonina flabelliformis Brady, 1884, FC. p. 374, pl. 45. figs. 17-21.

Heron-Allen & Harland, 1914-15, FKA. p. 632,
pl. 48. figs. 1-6.

One broken specimen which, when perfect, must have been of very large

0 B)

size.

SprropLecta Hhrenberg.

91. SPIROPLECTA BIFORMIS (Parker 5 Jones).
Textularia EEE, var. beformis Parker & Jones, 1865, NAAF. p. 370,
pl. 15. figs. 23, 24.
Spiroplecta biformis Brady, 1884, FC. p. 376, p. 45. figs. 25-27
Heron-Allen & Harland, 1914-15, FKA. p. 634.

ted 39

One typical specimen.
GauprRYINA a? Orbigny.

92. GAUDRYINA ScABRA Brady.
Gaudryina scabra Brady, 1854, FC. p. 381, pl. 12, fig. 5.
Heron-Allen & Harland, 1914-15, FKA. p. 635, pl. 48.
figs. 7-14.
Rare. The specimens are small, but compare in all respects with those
figured by us (ut supra). We there suggested that they might be multiform
variations of Verneuilina polystropha, and the Lord Howe specimens confirm

29 ”

that view.

93. GAUDRYINA RUGOSA (d’ Orbigny).
Gaudryina rugosa dOrbigny, 1840, CBP. p. 44, pl. 4. figs. 20, 21.

. Brady, 1884, FC. p. 381, pl. 46, figs. 14-16.

Heron-Allen & Karland, 1914-15, FKA. p. 635; 1922,

DN. p. 122.
Very common and finely developed. As in the New Zealand area, two

distinct types occur, to which we have called attention in our ‘Terra Nova’
Report (wt supra), one long and the other short.

ty) bh)

” ”

Subfamily BULIMININ &.
Butimina d’ Orbigny.

94, BULIMINA ELEGANTISSIMA d’Orbigny. (PI. 35. figs. 23, 24.)

Bulimina elegantissima d’Orbigny, 1839, FAM. p. 51, pl. 7. figs. 18, 14.
Brady, 1884, FC. p. 402, pl. 50. figs. 20-22.
i i Heron-Allen & Harland, 1914-15, FKA. p. 639.
Rare, but characteristic, and an interesting series; two specimens exhi-
biting budding-off tests, of one and two chambers respectively, precisely
similar to those figured by Heron-Allen (1915, RPF), and by Sidebottom

” tb)

THE FORAMINIFERA OF LORD HOWE ISLAND. 621

under the type-species B. elegantissima (S. 1904, etce., RFD. 1905, p. 11,
pl. 2. figs. 7-12), also a specimen in which the terminal septal face and
terminal septa have been absorbed.

95. BULIMINA WILLIAMSONIANA Brady.

Bulimina williamsoniana Brady, 1884, FC. p. 408, pl. 51. figs. 16, 17.
Millett, 1898, etc., FM. 1900, p. 279, pl. 2. fig. 8.
Heron-Allen & Harland, 1914-15, FK A. p. 641.

29 be)
2” ”

One very minute specimen.

Bonivina d’ Orbigny.

96. Botivina punctTarA d’ Orbigny.

Bolivina punctata d’Orbigny, 1839, FAM. p. 68, pl. 8. figs. 10-12.
Brady, 1854, FC. p. 417, pl. 52. figs. 18, 19.
i 3 Heron-Allen & Harland, 1914-15, FKA. p. 644.

Common, but rather weakly developed.

” ”

97. BoLivINA TEXTILARIOIDES Reuss.
Bolivina textilarioides Reuss, 1862, NHG. p. 81, pl. 10. fig. 1.
Brady, 1884, FC. p. 419, pl. 52. figs. 23-25.
55 5). Heron-Allen & Harland, 1914-15, FKA. p. 645.

Rare and small. A variety also occurs in greater numbers, characterized
by a broad compressed shell with rounded aboral extremity, five or six pairs
of chambers, separated by strongly limbate sutural lines. The parallel
arrangement of these lines indicates its affinity to B. textilarioides, but it differs
considerably from the type. The form is not infrequent in tropical shallow-

water gatherings.

” 2?

98. Bonivina DILATATA Feuss.

Bolivina dilatata Reuss, 1849-50, FOT, p. 381, pl. 3. (48) fig. 15.
Brady, 1884, FC. p. 418, pl. 52. figs. 20, 21.
5 55 Heron-Allen & Harland, 1914-15, FEA. p. 645.

Rare and far from typical.

” 3?

99. Bottvina tTorTUOSA Brady.
Bolivina tortuosa Brady, 1884, FC. p. 420, pl. 52. figs. 31-34.
Heron-Allen & Harland, 1913, CI. p. 66, pl. 5. fig.1; 1914-15,
FKA, p. 645.

Large and well-developed, but rare.

bb) ”

100. Bouivina RoBusta Brady.
Bolivina robusta Brady, 1884, FC. p, 421, pl. 53. figs. 7-9.
p Heron-Allen & Harland, 1914-15, FKA. p. 646.

One very small but otherwise typical specimen

622 MESSRS. &. HERON-ALLEN AND A. EARLAND ON

101. Bourvina timBata Brady.
Bolivina limbata Brady, 1884, FC. p, 419, pl. 52. figs. 26-28
Heron-Allen & Harland, 1918, CI. p. 67, pl. 5. figs. 2, 3;
1914-15, FKA. p. 646, pl. 50. figs. 1-4.
Rare, but large and well-developed. Most of the specimens are bifarine
in their later development.

” oy)

102. BoLiIviIna LIMBATA, var. ABBREVIATA, noy. (Pl. 36. figs. 25-27.)

Test compressed, consisting of 4 to 7 pairs of chambers with rounded
marginal edges, heavily limbate but with flush sutures. Aboral extremity
rounded. The shell reaches its maximum breadth at about the third pair of
chambers, and therefore the sides are almost parallel. Aperture large,
extending over the septal face of the final chamber. Surface of the chambers
prominently perforate between the clear sutural lines.

This is a very easily recognized little form, and not uncommon in shallow-
water tropical gatherings. It may represent an intermediate type between
B. teatilarioides (which it resembles in the straight sutural lines, almost
vertical to the long axis) and the typical B. limbata. It never, so far as we
have observed, has that tendency to develop a bifarine terminal portion so
characteristic of the latter species.

103. Bouivina VARIABILIs (Williamson).
Textularia variabilis (typica) Williamson, 1858, RFGB. p. 76, pl. 6. figs. 162,
163.
Bolivina "A Heron-Allen & Harland, 1914-15, FKA. p. 647.
The most typical Bolivina of the gathering, frequent, well-developed, and
exhibiting a considerable range in the extent of the surface-markings.

104. Boutvina puicara d’ Orbigny.
Bolivina plicata dOrbigny, 1839, FAM. p. 62, pl. 8. figs. 4-7.
es ni Goés, 1894, ASF. p. 51, pl. 9. figs. 487, 488.
Two specimens, very strongly marked.
105. Boutvina inFLATA Heron- Allen § Farland.

Bolivina inflata Heron-Allen & Earland, 1918, CL. p. 68, pl. 4. figs. 16-19 ;
1916, F.W.S. p. 240.

Not uncommon and quite typical.
Sub-family CassIDULININ®.
CassipuLina d’ Orbigny.

106. CassIDULINA CRASSA d? Orbigny.
Cassidulina crassa d’Orbigny, 1839, FAM. p. 56, pl. 7. figs. 18-20.

HE FORAMINIFERA OF LORD HOWE ISLAND. 623

Cassidulina crassa Brady, 1884, FC. p. 429, pl. 54. figs. 4, 5.
Cushman, 1910, ete., FNP. 1911, p. 97, fig. 141.

ub} ”

Rare and very small.

107. CassIDULINA SUBGLOBOSA Brady.

Cassidulina subglobosa Brady, 1884, FC. p. 480, pl. 54. fig. 17.
Cushman, 1910, etc., FNP, 1911, p. 98, fig. 152.
Heron-Allen & Harland, 1914-15, FKA. p. 652.

” ”

” ”

Frequent, occurring in two distinet forms, one very small and hyaline, the
other much larger (up to three times the size) and with a matt surface.
The second form presents the same external characteristics even in the
immature condition, and we are unable to offer any explanation of this
condition. They are not dead or eroded shells.

108. CasstpuLiIna (ORTHOPLECTA) CLAVATA Brady.
Cassidulina ( Orthoplecta) clavata Brady, 1884, FC. p: 432, pl. 113. fig. 9.
Chapman, 1901, FFA. (1902) p. 402 (list),
no. 114.
Heron-Allen & Harland, 1914-15, FKA.
p. 654, pl. 50. figs. 21, 22.

One small specimen of this very rare shallow-water tropical type.

29) 2? ”

HHRENBERGINA Reuss.

109. EHRENBERGINA SERRATA Reuss.
Ehrenbergina serrata Reuss, 1849-50, KOT. p. 377, pl. 48. fig. 7.
Cushman, 1910, ete., FNP. 1911, p. 101, fig. 155.
ss » Heron-Allen & Karland, 1922, TN. p. 140.

Rare. The specimens are small, with extremely pronounced, raised,
limbate sutures. Immature specimens are hardly distinguishable from the
figure of Cassidulina elegans Sidebottom (8S. 1910, J. Quekett Mier. Cl.
ser. 2, vol. xi. p. 106, pl. 4. fig. 1), to which, perhaps, we should have
ascribed them in the absence of the mature and more typical individuals.

by) ”

Family LAGENIDZ.
Sub-family LAGENIN A”.
LAGENA Walker & Boys.

110. Lacena tingata ( Williamson).
Entosolenia lineata Williamson, 1848, BSGL. p. 18, pl. 2. fig. 18.
Lagena » Brady, 1884, FC. p. 461, pl. 57. fig. 18.
Heron-Allen & Farland, 1914-15, FKA. p. 656.

One typical specimen.

9 »

624 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

111. Lacena squamosa (Montagu).
Vermiculum squamosum Montagu, 1803, TB. p. 526, pl. 14. fig. 2.
Lagena squamosa Cushman, 1910, etc., FNP. 1913, p. 16, pl. 6. fig. 1.
a us Heron-Allen & Earland, 1922, TN. p. 151, pl. vi. figs. 1, 2.
One abnormal, thick-shelled, and weakly marked individual.

112. Lacuna Lavis (Montagu).
Vermiculum leve Montagu, 1803, TB. p. 524.
Lagena levis Williamson, 1848, BSGL. p. 12, pl. 1. figs. 1, 2.
a », Brady, 1884, FC. p. 455, pl. 56. figs. 7-14, 30.
One abnormal specimen, the aboral half of which is very irregularly
formed.

113. Lagena L&vicata (leuss).
Fissurina levigata Reuss, 1849-50, FOT. p. 366, pl. 1. (46) fig. 1.
Lagena i Brady, 1884, FC. p. 473, pl. 114. fig. 8.
bp s Heron-Allen & Harland, 1914-15, FKA. p. 661.

One specimen with a hooded aperture resembling Sidebottom’s fig. 7 on
pl. 17 (S. 1912, LSP. p. 400), which he considers to be ailied to L. mil-
lettzi Chaster (C. 1892, FS. p. 61, pl. 1. fig. 10). The Lord Howe specimen
has a normal aboral base. A good many small specimens also of the
typical form.

114. Lagena ANNECTENS burrows § Holland.
Lagena annectens Burrows & Holland, in J., P., & B. 1866, ete., MFC. 1895,
p. 208, pl. 7. fig. 11.
4p a Heron-Allen & Earland, 1922, TN. p. 155.
One small weak specimen.

115. Lacena mareinata ( Walker 5 Boys).
Serpula (Lagena) marginata Walker & Boys, 1784, TMR. p. 2, pl. 1. fig.
Lagena marginata Brady, 1884, FC. p. 476, pl. 59. figs. 21-28.
Heron-Allen 6: Harland, 1914-15, FKA. p. 663.

Many small fully marginated specimens.

Sy

” 2

116. Lagena BICARINATA ( Terquem).
Iissurina bicarinata Terquem, 1882, FEP. p. 31, pl. 1. (9) fig. 24.
Lagena i Millett, 1895, etc., FM. 1901, p. 624, pl. 14. fig. 13.
i 4 Heron-Allen & Harland, 1914-15, FKA. p. 665.

One minute specimen.

117. LacENna chatTHRATA Brady.

Lagena elathr ata Brady, 1884, FC. p. 485, pl. 60. fig.
a3 Heron-Allen & Harland, 1913, CI. p. on pl. 7. fig. 10.

One very feebly costate specimen.

THE FORAMINIFERA OF LORD HOWE ISLAND. 625

Sub-family NoposaRIINA.
FronprcuLaria Defrance. ‘

118. Fronprcvunarra scorrt Heron- Allen § Earland. (P1. 36. figs. 28, 29.)
Frondicularia scottii Heron-Allen & Harland, 1922, TN. p. 175, pl. 6.
figs. 30-32.

One small and feeble example which we have no hesitation in ascribing
to this species. It lacks the longitudinal grooves and decorations of the
type, but it has the same truncate edges and thick shell-wall, which obscures
the internal structure. This is brought out by wetting the specimen. The
occurrence of this individual in a shore sand is very noteworthy; it is
practically identical in size with the pauperate specimen from Raine Island,
recorded by us wt supra, but is even weaker, having no limbation of the
sutural lines. We figure both the Lord Howe and Ranie Island specimens.

CRISTELLARIA Lamarck.

119. CrIsTELLARIA ROTULATA (Lamareh).
Lenticulites rotulata Lamarck, 1804, AM. p. 188, no. 3; 1830, EM. p. 330,
pl. 466. fig. 5.
Cristellaria ,, Brady, 1884, KC. p. 547, pl. 49. fig. 18.
i i Heron-Allen & Harland, 1914-15, FKA. p. 671.
Rare and rather small, but typical.

120. CRISTELLARIA GIBBA a? Orbigny.
Cristellaria gibba, VOrbigny, 1839, FC. p. 40, pl. 7. figs. 20, 21.
cy » Brady, 1884, FC. p. 546, pl. 69. figs. 8, 9.
45 » Cushman, 1910, etc., FNP. 1913, p. 69, pl. 35. fig. 1.

One weak specimen.

Sub-family PoLYMORPHININ®.
PotyMorPHINA a’ Orbigny.

121. PoLyMORPHINA COMMUNIS d? Oriigny.
Polymorphina (Gauttulina) communis dOrbigny, 1826, TMC. p. 266,
nos, 14, 15, pl. 12. figs. 1-4; Modéles, nos. 61, 62.
hi communis Brady, 1884, FC. p. 568, pl. 72. fig. 19.
ea ‘ Heron-Allen & Marland, 1914-15, FKA. p. 673.

A single small specimen.
122. PoLYMORPHINA PROBLEMA d’ Orligny.

Polymorphina (Guttulina) problema VOrbigny, 1826, TMC. p. 266, no. 14;
Modéle, no, 61,

626 MESSRS. E. HERON-ALLEN AND A. HARLAND ON

Polymorphina problema Brady, 1884, FC. p. 568, pl. 72. fig. 20 ; pl. 73,
fig. 1.
Heron-Allen & Earland, 1922, TN. p. 182.

” ”

A single small specimen.

123. PoLyMORPHINA REGINA Brady, Parker, § Jones.
Polymorphina regina Brady, Parker, & Jones, 1870, GP. p. 241, pl. 41. fig. 32.
‘ », Brady, 1884, FC. p. 571, pl. 73. figs. 11-13.
se ,, Cushman, 1910, etc., FNP. 1913, p. 91, pl. 41. figs. 6, 7.

One very large broken specimen and one of normal size.

Uvicurina a? Orbigny.

124, Uviarrina pyemaa d’ Orbigny.
Uvigerina pygmc@a @Orbigny, 1826, TMC. p. 269, pl 12. figs. 8,9; Modéle,
no. 67.
Brady, 1884, FC. p. 575, pl. 74. figs. 11, 12.
- ie Heron-Allen & Harland, 1914-15, FKA. p. 675.
Three rather small specimens with very strong costation.

” oP)

125. Uvicrrina porrecta brady.

Unigerina porrecta Brady, 1879, etc., RRC. 1879, p. 274, pl. 8. figs. 15, 16.
Brady, 1884, FC. p. 577, pl. 74. figs. 21-23,
Heron-Allen & Harland, 1914-15, FKA. p. 675.

” oh)
”

Frequent and typical.

126. Uvicerina sELSEYENSIS Heron-Allen § Earland.
Uvigerina selseyensis Heron-Allen & Harland, 1908, ete., SB. 1909, p. 437,
pl. 18. figs. 1-3.
Cushman, 1910, etc., FNP. 1913, p. 93, pl. 42. fig. 5.
Several specimens resembling the Hocene fossils recorded by us (ut supra),
in the shape and arrangement of the chambers, but with a rough, feebly

” ”

hispid or striate surface.

127. Uvierrtna ancuLosa Walliamson.
Uvigerina angulosa Williamson, 1858, REGB. p. 67, pl. 5. fig. 140.
Brady, 1884, FC. p. 576, pl. 74. figs. 15-18.
=! os Heron-Allen & Harland, 1914-15, FKA. p. 676.
Frequent and large. Two separate forms occur, one very long, the other
short and rapidly increasing in breadth.

” oy)

Sacrina Parker 5 Jones.

128. SAGRINA COLUMELLARIS Brady.

Sagrina columellaris Brady, 1884, FC. p. 581, pl. 75. figs. 15-17.
Cushman, 1910, etc., FNP. 1913, p. 104, pl. 47. figs. 2,3,
Heron-Allen & Harland, 1914-15, FKA. p. 676,

oh) oh)

+}e) 29

Rare, but typical,

lor}
i)
1

THE FORAMINIFERA OF LORD HOWE ISLAND.

129. Sacrina RAPHANUS Parker 5; Jones.
Uvigerina (Sagrina) raphanus Parker & Jones, 1865, NAAF. p. 364, pl. 18.
figs. 16, 17.
Sagrina raphanus Brady, 1884, FC. p. 585, pl. 75. figs. 21-24.
5 Heron-Allen & Earland, 1914-15, FKA. p. 677.

Rare, large, of the entosolenian type.

Sub-family RaMULININA.
RaMULINA Rupert Jones.

130. RaMULINA GRIMALDIL Schlumberger. (Pl. 36. fig. 32.)
Ranulina grimaldi Schlumberger, 1891, Mém. Soe. Zool. France, vol. iv.
pp. 509-511, pl. 5.
Ramnulina? Heron-Allen & Harland, 1922, TN. p. 187, text-fie.

The occurrence in this material of a specimen practically identical in
formation with the one recorded by us (ut supra) from New Zealand seems to
render it necessary to give it a specific name, a task which we have hitherto
avoided. Rather than make a new species on such insufficient material, we
have decided to use Schlumberger’s name. The organism described and
figured by Flint as I. proteiformis (F. 1899, REA. p. 321, pl. 68. fig. 7)
appears to bea simpler form of the same kind and possibly identical with
Schlumberger’s species.

Family GLOBIGERINID AN.

GLOBIGERINA d? Orbigny.
131. GLOBIGERINA BULLOIDES d’ Orbigny.
Globigerina bulloides V@Orbigny, 1826, TMC. p. 277, no. 1; Modeles, nos. 17, 76.
Brady, 1884, FC. p. 593, pls. 77 & 79. figs. 3-7.
, Heron-Allen & Harland, 1914-15, FKA. p. 678.

The specimens are small and few in number.

oh)

132. GLOBIGERINA TRILOBA Reuss.
Globigerina triloba Reuss, 1849-50, FOYT. p. 374, pl. 2. (47) fig. 11.
Brady, 1884, FC. p. 595, pl. 79. figs. 1, 2; pl. 81. figs. 2, 3.
= » Heron-Allen & Harland, 1914-15, FIA. p. 678.
More frequent, larger, and better developed than G. bulloides.

” 29

133. GLOBIGERINA CRETACEA, var. EGGERI Heron-Allen §- Farland.
Globigerina cretacea, var. eggeri Heron-Allen & Harland, 1922, TN. p. 188,
pl. 7. figs. 6-8,

Two small but typical individuals,

628 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

134. GLOBIGHRINA RUBRA d’ Orbigny.
Globigerina rubra d’Orbigny, 1839, FC. p. 82, pl. 4. figs. 12-14.
i » Brady, 1884, FC. p. 602, pl. 79. figs. 11-16.
ay », Heron-Allen & Harland, 1914-15, FKA. p. 679.

Large and not infrequent, but devoid of the characteristic colouring.

Family ROTALITD Ai.
Subfamily SPIRILLININ &.
Sprritina Hhrenberg.
135. SPrRILLINna vivipaRra Hhrenberg.
Spivillina vivipara Khrenberg, 1841, SNA. p. 449 , pl. 3. fig. 41.
M ur Brady, 1884, FC. p. 630, pl. 85. figs. 1-5
a a Heron-Allen & Harland, 1914-15, FKA. p. 683, pl. 51.
figs. 19-28.
Frequent, and presenting the usual wide range of breadth of tube and
number of convolutions.

136. SprRILLIna LimBaTa Brady.
Spirillina limbata Brady, 1879, RRC. p. 278, pl. 8. fig. 26.
ie » Heron-Allen & Harland, 1914-15, FIKA. p. 684.

Rare and not large, but otherwise characteristic.

137. SprRILLINA DECORATA Brady.
Spirillina decorata Brady, 1884, FC. p. 633, pl. 85. figs. 22-25.
Fi a Heron-Allen & Earland, 1914-15, PKA, p. 685.
Very rare and small.

138. SPIRILLINA DECORATA, var. UNILATERA Chapman.
Spirillina decorata, var. wnilatera Chapman, 1901, etc., FFA. 1902, p. 410,
pl. 36. fig. 17.

A single specimen, exactly resembling Chapman’s figure.

139. SPrRILLINA CAMPANULA, sp. nov. (Pl. 36. figs. 33-41.)

‘Test minute, thin-walled, free or sessile, campanulate or bell-shaped, but
more or less laterally compressed, the apex of the bell obtusely rounded.
The exterior surface of the test, viewed as an opaque object, is dull, and the
sutural lines are flush and only visible as internal markings. Viewed as a
transparent object under a high power, the surface is slightly rough or scaly,
and the interior of the bell is filled almost to the rim with an unseptate tube,
which, starting from a rather large proloculum occupying the apex of the
bell, is coiled in 3-5 convolutions round a solid central axis or columella.
Viewed from the base of the bell, the umbilicus is solid, smooth, and rather

THE FORAMINIFERA OF LORD HOWE ISLAND. 629

deeply depressed, showing no sign of coiling or of an oral aperture. As in
Spirillina lucida Sidebottom, the oral extremity of the tube is compressed and
sealed into the outer wall of the test. The protoplasmic body, which is dark
yellow in colour and full of minute granules of darker tint, is voluminous
and occupies the whole of the convolutions.

Spirillina campanula is interesting as presenting an entirely novel plan
of growth. The genus is normally a plano-spirally coiled tube, but in most
of the species a certain degree of asymmetry is observable between the two
faces of the spire, one being raised and the other depressed. This tendency
is particularly noticeable in S. revertens Rhumbler and S. lucida Sidebottom,
and is still more pronounced in S. groomit Chapman, but none of these can
compare with the extraordinary convexity of S. campanula.

The species is probably abundant at Lord Howe Island, as many specimens
were found in spite of the very small quantity of fine material available for
examination. All the specimens were free except one, which was sessile on
a fragment of coralline. The perfect manner in which the rim of the bell
with its excised edges adapted itself to the shape of the host suggests that
such asessile habitat may be normal, at any rate, up to a stage in the existence
of the organism. Such conditions would account for the peculiar form of
the rim and the varying degree of compression of the bell. Buta careful
search of the small quantity of coralline available did not provide further
specimens.

One large individual has lost allinternal septa by absorption, and the entire
cavity of the bell is packed with young indiyiduals, each consisting of a
proloculum encircled by a complete coil of tube. At least ten individuals
can be counted in the cavity.

The size and relative proportions are very variable. Five specimens
measured for height and maximum breadth gave the following measure-

ments :—
JalGH ING coonceocn 15 OTS. «lS = OOH LIL} seam,
Breadth ...... CIS NS) ILA Me) 115) rae,
Four others measured gave :—
Maximum breadth...... “135 cil LL 095) enna.
Thickness at rim ...... OW, Os OS7 OS . mam,

Sub-family RovaLrin a.
Pavevtina Williamson.

140. PATELLINA coRRUGATA Williamson.
Patellina corrugata Williamson, 1858, RFGB. p. 46, pl. 3. figs. 86-89.
a Mi Brady, 1884, FC. p. 634, pl. 86. figs. 1-7.
an 35 Heron-Allen & Earland, 1914-15, FIA. p. 687.

Common and typical. All of the rather low-domed, scaly type.
LINN, JOURN.—ZOOLOGY, VOL, XXXV, 45

630 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

CymBaLopora Hagenow.

141. CymBatopora Pony (d’ Orbigny).
Rosalina pocyi WV Orbigny, 1839, FC. p. 92, pl. 3. figs. 18-20. Rotalia squam-
mosa, p. 91, pl. 3. figs. 12-14.
Cymbalopora poeyt Brady, 1884, FC. p. 636, pl. 102. fig. 13.
Hs » Heron-Allen & Karland, 1914-15, FKA, p. 687.

One small and weak, and one large and typical specimen.

142. CympaLopora mILLETTII Heron-Allen § Farland.
Cymbalopora bulloides Millett, 1898, etc., FM. 1903, p. 697, pl. 7. fig. 4
mullettvi Heron-Allen & Earland, 1915, FKA. p. 689, pl. 51.
figs, 32-35.
ts he Heron-Allen, 1915, RPF. p. 253, pl. 16. fig. 36 ; pl. 17.
figs. 46-48, 50, 51.
One perfect specimen with the characteristically marked balloon, and a
number of specimens of the apical rotaline mass of chambers.

DiscorBina Parker 5 Jones.

143. DiscorBina cora (d’Orbigny).
Rosalina cora d’Orbigny, 1839, FAM. p. 45, pl. 6. figs. 19-21.
» ay Heron-Allen. & Earland, 1915, FKA. p. 690; 1922, FGA.
p- 183, pl. 1 (aumbered 2). figs. 33-35.
Fairly typical examples of this very primitive form of D. globularis occur,
but they are rare.

144. Discorpina nitipa ( Williamson).
Rotalina nitida Williamson, 1858, RFGB. p. 54, pl. 4. figs. 106-108.
Rotalia ,, Brady, 1884, FC. p. 627.
Discorbina,, Heron-Allen & Karland, 1914-15, FKA, p. 691.

Very rare, minute and weak.

145. Discorpina concinna Brady.
Discorbina concinna Brady, 1884, FC. p. 646, pl. 90. figs. 7, 8
Ms » LHgger, 1893, FG. p. 388, pl. 15. figs. 99-9 4,
a », Heron-Allen & Harland, 1914-15, FKA. p. 691.
Frequent and typical.

146. DiscorBINA ISABELLEANA (d@’ Orbigny).
Rosalina isabelleana @Orbigny, 1839, FAM. p. 48, pl. 6. figs. 10-12.
Discorbina ,, Brady, 1884, FC. p. 646, pl. 88. fig. 1
i Hy Heron-Allen & Earland, 1914-15, FKA. p. 692.

Frequent. The specimens are good, but small.

THE FORAMINIFERA OF LORD HOWE ISLAND. 631

147. DiscorBINA VILARDEBOANA (d’ Orbigny).
Rosalina vilardeboana W’Orbigny, 1839, FAM. p. 44, pl. 6. figs. 13-15.

‘3 33 Brady, 1884, FC. p. 645, pl. 86. fig. 12; pl. 88. fig. 2.
3 4 Heron-Allen & Harland, 1914-15, FKA, p. 692.

Frequent and generally small, but some of the specimens attained very
large size.

148. DiscorpinA Rosacea (d’Orligny).
Rotalina rosacea d Orbigny, 1826, TMC. p. 273, no. 15; Modeéle, no. 39.
Discorbina ,, Brady, 1884, FC. p. 644, pl. 87. figs. 1, 4.
M » Heron-Allen & Karland, 1914-15, FKA, p. 692.
Frequent and typical.

149. Discorpina BAccATA Feron-Allen §- Harland.
Discorbina baccata Heron-Allen & Harland, 1913, CL. p. 124, pl. 12. figs. 1-8 ;
1922, TN. p. 200. i
A single small but typical specimen.

150. Discorsina TURBO (d’ Orbigny).
Rotalia (Trochulina) turbo @Orbigny, 1826, TMC. p. 274, no. 39; Modéle,

no. 73.
Discorbina turbo Brady, 1884, FC. p. 642, pl. 87. fig. 8.
- » Heron-Allen & Harland, 1914-15, FKA. p. 693.

A single small specimen.

151. DiscorBINA MEDITERRANENSIS (d’Orbigny).
Rosalina mediterranensis VOrbigny, 1826, TMC. p. 271, no. 2.
Discorbina aH Heron-Allen & Harland, 1913, CI. p. 118, pl. 9.
figs. 12-14, pl. 10. fig. 1; 1914-15, FKA. p. 693.
Common and very typical.

152. DiscorBINA GLOBULARIS (d’ Orbigny).
Rosalina globularis d’Orbigny, 1826, TMC. p. 271, no. 1, pl. 13. figs. 1-4 ;
Modeéle, no. 69.
Discorbina ,, Brady, 1884, FC. p. 643, pl. 86. figs. 8, 13.
" ry Heron-Allen & Harland, 1914-15, FIKA. p. 694, pl. 51.
figs. 36-39.
Very common, presenting every range of variation from depressed forms
hardly separable from D. cora to highly inflated specimens separable only
from D. valvulata by the absenee of limbation.

153. DiscorBINA ARAUCANA (d’ Orbigny).
Rosalina araucana V@Orbigny, 1839, FAM. p. 44, pl. 6. figs. 16-18.
Discorbina araucana Brady, 1884, FC. p. 645, pl. 86. figs. 10, 11.
mk rs Heron-Allen & Harland, 1914-15, FKA. p. 695.

ommon and fairl y t nical Varying considera bl 1n size.
C ) 5S d
45

632 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

154. DiscorBINA VALVULATA (d’Orbigny).
Rosalina valvulata dOrbigny, 1826, TMC. p. 271, no. 4; 1836, FIC. p. 136,
pl. 2. figs. 19-21.
Discorbina ,, Brady, 1884, FC. p. 644, pl. 87. figs. 5-7.
Ne a Heron-Allen & Earland, 1914-15, FKA, p. 695.

Common. Very fine and typical specimens.

155. Discorpina RENIFORMIS [eron-Allen § Harland.
Discorbina reniformis Heron-Allen & Harland, 1914-15, FKA. p. 698, pl. 52.
figs. 7-14.
A single quite characteristic specimen of this very obscure species. It
appears to be very widely distributed over the Indo-Pacific area, though it
is always rare.

156. Discorpina RuGOSsA (d’ Orligny).
Rosalina rugosa V@Orbigny, 1839, FAM. p. 42, pl. 2. figs. 12-14.
Discorbina ,, Brady, 1884, FC. p. 652, pl. 87. fig. 3; pl. 91. fig. 4.
eS » Heron-Allen & EKarland, 1914-15, FKA. p. 197.

Rare, but very fine and typical specimens.

157. DiscoRBINA POLYSTOMELLOIDES Parker § Jones.
Discorbina polystomelloides Parker & Jones, 1865, NAAF. p. 421, pl. 19. fig. 8.
. ey Brady, 1884, FC. p. 652, pl. 91. fig. 1.
Heron-Allen & Harland, 1914-15, FKA. p. 698,

pl. 52. figs. 19-23.

Common and attaining comparatively enormous dimensions, the largest
having a maximum diameter of 3mm. The specimens vary considerably in
their surface ornament, some having developed secondary shell-structure in
the form of beads and ribs to an extent that we have never previously
observed in this species.

9 ”

158. Discorpina rimosa Parker & Jones.
Discorbina rimosa Carpenter, Parker, & Jones, 1862, ISF. p. 205.
Millett, 1898, etc., FM. 1903, p. 702, pl. 7. fig. 7.
5 » Heron-Allen & Harland, 1914-15, FKA. p. 700.

The specimens referable to this species merely tend to confirm our views
expressed (ut supra) as to the inexpediency of separating this species from
D. polystomelloides. They could very well be included as pauperate speci-
mens of the latter, in view of the wide range of variation exhibited by that

oP) 2”

species.
159. DiscoRBINA RARESCENS brady.

Discorbina rarescens Brady, 1884, FC. p. 651, pl. 90. figs. 2, 3, & ?4.
Heron-Allen & Harland, 1914-15, FIA. p. 700.

One small but perfectly typical specimen,

” 29

THE FORAMINIFERA OF LORD HOWE ISLAND. 633

160. DiscorBInA puLVINATA Brady. (PI. 36. figs. 42-46.)
Discorbina pulvinata Brady, 1884, FC. p. 650, pl. 88. figs. 10, a, b.
= ie Sidebottom, 1904, ete., RFD. 1908, p. 14, pl. 5. tig. 4.
ws ss Heron-Allen & Harland, 1914-15, FKA. p. 703.
Frequent, and the specimens vary between relatively smooth and highly

corrugate on the superior face. Nearly all of them exhibit “budding” in
various stages, from a single minute primordial chamber in the umbilicus of
the oral side to specimens in which the base and internal septa have disap-
peared by absorption. Two individuals with young brood clustered round
the orifice were observed, which we figure.

161. DiscoRBINA PATELLIFORMIS Brady.
Discorbina patelliformis Brady, 1884, FC. p. 647, pl. 88. fig. 3; pl. 89. fig. 4.
a aks Sidebottom, 1904, ete., RFD. 1908, p. 14, pl. 5. fig. 3.
Heron-Allen & Harland, 1914-15, FKA. p. 703,
pl. 52. fig. 32.
Very common and attaining a large size. The specimens are perfectly
preserved and very handsome. One “budding” pair of small individuals

oh) ”

was observed.

162. DiscorBINA HARMERI [Teron-Allen § Earland.
Discorbina harmeri Heron-Allen & Karland, 1922, TN. p. 204, pl. 7. figs. 9-11.
One minute but typical specimen. It is interesting to note tle occurrence
of this recently described species in a new area, though it is evidently not
firmly established there.

163. DiscoRBINA TABERNACULARIS Brady. (PI. 86. figs. 47-49.)
Discorbina tabernacularis Brady, 1884, FC. p. 648, pl. 89. figs. 5-7.
5 $0 Heron-Allen & Harland, 1914-15, FIA, p. 704.
Common and quite typical. Many “budding” and associated pairs ; also
individuals with the base and septa eroded. A specimen of the latter where
the internal cavity was filled with young individuals (fig. 49).

164. DigscorBINA LAURIEI, nom. noy. (PI.36. figs. 50-52; PI. 37. figs. 53-55.)
Discorbina tabernacularis Sidebottom, 1910, RFBP. p. 25, pl. 3. fig. 12.

The little test figured by Sidebottom though doubtless allied to /). taber-
nacularis cannot be taxonomically referred to that species. It has many
points of distinction: the paucity of chambers, which rarely exceed two con-
volutions of five chambers each; its blunted, or rounded, apex, with a
prominent primordial chamber ; its greatly inferior but constant size; and
perhaps, most of all, its constant habit of “budding” pairs. Specimens
which do not present this feature are so uncommon as to be noticeable. It

634 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

is moderately common in the material, as it is, indeed, in shallow-water
gatherings from most tropical and temperate seas. Liebus and Schubert’s
variety, D). tabernacularis, var. levis (Jahrb. k.-k. geol. Reichsanst. 1902,
vol. lii. p. 301, fig. 5), has many points of resemblance, but differs in the
absence of the ghar acteristic radial beading which decorates the base of
LD). lauriei, as it does the base of J). taber eee

165. DiscoRBINA PYRAMIDALIS, sp. nov. (PI. 87. figs. 56-61.)

Test free, shaped like a four-sided pyramid with slightly excavated sides,
and salient angles, consisting of 3-5 convolutions, each of four chambers, the
widest portion of each chamber being at the angle, and the sutural divisions
running down the middle of each side. Shell-wall thick ; the base excavate,
studded with beads, arranged radially, with the aperture of the final chamber
in the umbilical recess. In all the specimens examined in balsam the
primordial chamber is large and spherical, occupying as much as a quarter
of the pointed apex of the pyramid.

Several “twinned” pairs occur. The specimens are usually about the
same size. ‘There is a considerable range both in size and height of the free
individuals; in fact, there appear to be two fairly distinctive forms, one small
and more or less regularly pyramidal (7. e., the height being roughly equal to
the diameter of the base), the other with a height double that diameter.
These may represent the megalo- and microspheric stages of the organism.
Among other variations noticeable are (1) a single three-sided specimen,
(2) several specimens in which the angular edges of the pyramid are curving
lines, so that, when viewed from the top, a spiral structure is suggested.
The shell is thick and free from surface ornamentation.

This species is evidently closely allied to Millett’s D. corrugata (M. 1898,
ete., FM. 1903, p. 700, pl. 7. fig. 5). We have good examples of this, from the
Arakan coast of Burmal!. Millett’s original types, which are in our possession,
bear, in his handwriting, as locality “Sagami Bay, Japan.” Millett’s form
has five salient angles and a much less polished exterior than D. pyramidalis.
We have typical specimens of D. pyramidalis from Suva, Fiji Is. (12 fms.,
very rare), and we have noted its occurrence at ‘Challenger’ Station 185,
“Raine Id.” (155 fms., also very rare), so that we may assume that the
form is widely distributed in the tropical Pacific. It is of constant occur-
rence in a series of shore-sands which we possess from St. Vincent’s Gulf
and Spencer Gulf (8. Australia), and it also occurs in two shore-sands from
W. Australia (Fremantle and Rottnest Island). The lack of suitable
material from the remaining shores of Australia renders the records incom-
plete, but it appears highly probable that the species will be found to be a
typical form in Australian shore-gatherings.

Size. Basal diameter at edges *08—16 mm. (small type) to 28 mm. (large
type). Height 05-21 mm. (small type) to °35 mm. (large type).

THE FORAMINIFERA OF LORD HOWE ISLAND. 635

PLANORBULINA d@’ Orbigny.

166. PLANORBULINA MEDITERRANENSIS d’ Orbigny.
Planorbulina mediterranensis d’Orbigny, 1826, TMC. p. 280, pl. 14. figs. 4-6 ;
Modeéele, no. 79.
Brady, 1884, FC. p. 656, pl. 92. figs. 1-3.
Heron-Allen & Harland, 1914-15, FKA. p. 705.

” ”?
” oh)

A single large specimen.

167. PLANORBULINA LARVATA Parker § Jones.
Planorbulina vulgaris, var. larvata Parker & Jones, 1859, ete., NF. 1860,
p. 294.
larvata Brady, 1884, FC. p. 658, pl. 92. figs. 5, 6.
Heron-Allen & Earland, 1914-15, FKA. p. 706.

9
9 ”?

One somewhat doubtful specimen.

TRUNCATULINA dOrbigny.

168. TRUNCATULINA LOBATULA ( Walker 5 Jacob).
Nautilus lobatulus Walker & Jacob, 1798, AEM. p. 642, pl. 14. fig. 35.
Truncatulina lobatula Brady, 1884, FC. p. 660, pl. 92. fig. 10 (ete.).
- a Heron-Allen & Earland, 1914-15, FKA. p. 706.

Common and presenting the usual range of variation.

169. TRUNCATULINA VARIABILIS d@’ Orbigny.
Truncatulina variabilis V@Orbigny, 1826, TMC. p. 279, no. 8.
Brady, 1884, FC. p. 661, pl. 93. figs. 6, 7.
Heron-Allen & Karland, 1914-15, FKA. p. 706; 1922,
FGA. p. 137, pl. 1 (numbered 2). figs. 38, 39.

oy) ”

” by)

Very.common, ranging from normal variations up to the wild-growing
Soldanian types.

170. TRUNCATULINA REFULGENS (Montfort).
Cibicides refulgens Montfort, 1808-10, CS. vol. i. p. 122, 31™e genre.
Truncatulina ,, Brady, 1884, FC. p. 659, pl. 92. figs, 7-9
Heron-Allen & Harland, 1914-15, FKA. p. 707.

” ”

Very common and highly typical.

171. TRUNCATULINA UNGERIANA (a Orbigny).
Rotalina ungeriana VOrbigny, 1846, FFV. p. 157, pl. 8. figs. 16-18.
Truncatulina ,, Brady, 1884, FC. p. 664, pl. 94. fig. 9.

3 », Heron-Allen & Karland, 1914-15, FKA. p. 708.

Frequent and well-developed.

(oa)

636 MESSRS. E. HERON-ALLEN AND A. BARLAND ON

172. TRUNCATULINA PR&CINOTA Karrer.
Rotalia precincta Karrer, 1868, MF KB. p. 189, pl. 5. fig. 7.
Truncatulina ,, Brady, 1884, FC. p. 667, pl. 95. figs. 1-3.
“4 », Heron-Allen & Harland, 1914-15, FKA. p. 709.
Very rare, but typical.

Poutvinutina Parker § Jones.

173. PULVINULINA REPANDA (Fchtel 5 Moll).

Nautilus repandus Fichtel & Moll, 1798, TM. p. 35, pl. 3. figs. a—d.
Pulvinulina repanda Brady, 1884, FC. p. 684, pl. 104. fig. 18.
a bs Heron-Allen & Harland, 1914-15, FKA. p. 713.

Frequent and some of the specimens are large, but nearly all broken or
water-worn.

174. PULVINULINA LATERALIS (Tergquem).
Rosalina lateralis Terquem, 1878, FIR. p. 25, pl. 2. (7) fig. 11.
Pulvinulina ,, Brady, FC. p. 689, pl. 106. figs. 2, 3.
i », Heron-Allen & Farland, 1914-15, FKA. p. 714, pl. 53.
figs. 6-11.

One large and typical specimen.

175. PULVINULINA CoNcENTRICA Parker & Jones.
Pulvinulina concentirica Parker, Jones, & Brady, 1864, RES. p. 470, pl. 48.

fig. 14.
be a Brady, 1844, FC. p. 686, pl. 105. fig. 1.
es my Heron-Allen & Harland, 1914-15, FKA. p. 714.

Small, but frequent and typical.

176, PULVINULINA CANARIENSIS (@’ Orbigny).
Rotalina canariensis d’Orbigny, 1839, FIC. p. 130, pl. 1. figs. 34-36.
Pulvinulina —,, Brady, 1884, FC. p. 692, pl. 108. figs. 8-9.
ss es Cushman, 1921, FP. p. 338, pl. 66. fig. 1
Rare, but quite typical.

177. PuLVINULINA cRASSA (d’ Orbigny).
Rotalina crassa VOrbigny, 1840, CBP. p. 32, pl. 3. figs. 7, 8.
Pulvinulina ,, Brady, 1884, FC. p. 694, pl. 103. figs. 11, 12.
», Cushman, 1921, FP. p. 338, pl. 67. fig. 3.
Frequent and well-developed.

178. PULVINULINA TRUNCATULINOIDES (d’ Orbigny).
Rotalina truncatulinoides d’Orbigny, 1839, FIC. p. 182, pl. 2. figs. 25-27.
Pulvinulina 5 Heron-Allen & Farland, 1922, TN. p. 216.

THE FORAMINIFERA OF LORD HOWE ISLAND. 637

Very rare. This is the type with the sunken umbilicus and pronounced
sutures as contrasted with the smooth and rounded type P. micheliniana.
We have gone into this matter on several occasions, and particularly
ut supra.

179. PULVINULINA ELEGANS (d’ Orbigny).
Rotaha (Lurbinulina) elegans @’Orbigny, 1826, TMC. p. 276, no. 54.
Pulvinulina elegans Brady, 1884, FC. p. 699, pl. 105, figs. 4-6.
a » Heron-Allen & Harland, 1914-15, FKA. p. 717.

One small and broken specimen.

Rorauia Lamarck.

180. Rorania BECCARII (Linné).
Nautilus beccarti Linné, 1767, SN. p. 1162; 1788, SN. p. 3370.
Rotaha (Turbinulina) beccarti d@Orbigny, 1826, TMC. p. 275, no. 42; Modéle,
no, 74,
» beccarti Brady, 1884, FC. p. 704, pl. 107. figs. 2, 3.

One large but very weak specimen, hardly separable from the next species.

181. Roranta pERLUCIDA Heron-Allen § Harland.
Rotalia beccarti (pars) Balkwill & Wright, 1885, DIS, p. 351,
perlucida Heron-Allen & Karland, 1913, C1. p. 139, pl. 13, figs. 7-9 ;
1914-15, FIA. p. 718.

Not uncommon. Hxtremely depressed and pauperate.

”

Sub-family TINOPORIN&.
GypsIna Carter.

182. GYPSINA INHARENS (Schultze). (Pl. 37. figs. 62-64.)
Acervulina inherens Schultze, 1854, OP. p. 68, pl. 6. fig. 12.
Gy psina se Brady, 1884, FC. p. 718, pl. 102. figs. 1-6.
be is Heron-Allen & Harland, 1914-15, FKA. p. 724.

Very common, and, as usual, very variable in the size of the chamberlets.
One specimen was observed characterized by the extremely smail size of
the constituent chambers, some of which are broken and disclose tightly
packed young individuals within (fig. 62).

Among the noticeable variations are two forms: one, white with very
large and loosely aggregated chambers, often extending in acervuline
projections, and sometimes fairly smooth and coarsely perforate, at others
covered with densely aggregated spines of secondary shell-matter. The
shape may possibly be due to the irregular surface of the host from which
the specimens have become detached. In another variant, possibly more
nearly related to G. rubra, the form is wild-growing, large-chambered, the

638 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

chambers often running in irregular chains, thick-walled, coarsely perforate,
but without exogenous shell-matter, colour inclined to pale red running to
dirty white. The final chamber of an extended series is often subglobular
in shape and of large size.

The first variety suggests Holocladina pustulifera Carter (C. 1880, SGM.
p. 447, pl. 18, figs. 4, a-g), but lacks the terminal perforation of the spine,
upon which Carter lays a stress which appears to us to be superfluous.

183. Gypsmva RUBRA (d’ Orbigny).
Planorbulina rubra d’Orbigny, 1826, TMC. p. 280, no. 4.
» Fornasini, 1908, SON. p, 44, pl. 2. fig. 3.
Gypsina » Heron-Allen & Harland, 1914-15, FKA. p. 725, pl. 53.
figs. 35-37.
Many large fragments. The exogenous shell-growth is not so abnormally
developed as is usually the case in this form.

184. GypsINa VEsIcuLARIS (Parker 5 Jones).
Orbitolina vesicularis Parker & Jones, 1859, etc., NF. 1860, p. 31, no. 5.
Gypsina i Brady, 1884, FC. p. 718, pl. 101. figs. 9-12.
Heron-Allen & Earlend, 1914-15, FKA. p. 726.

A few large water-worn specimens.

” 2)

185. Gypsina GLOBULUS (Reuss).
Ceriopora globulus Reuss, 1847, Haidinger’s Naturw. Abh. vol. ii. p. 33, pl. 5.
1H,
Gypsina 3 Brady, 1884, FC. p. 717, pl. 101. fig. 8.
3 Heron-Allen & Earland, 1914-15, FKA. p. 727.

Large, but infrequent.

BacuLoeypsina Sacco.

186. BACULOGYPSINA SPHZRULATA (Parker § Jones). (PI. 37. fig. 65.)
Orbitolina spherulata Parker & Jones, 1859, ete., NF. 1860, p. 33.
Tinoporus baculatus Carpenter, 1856, etc., RF. 1860, p. 564, pl. 18, figs. 2-6.

Bs ‘A Brady, 1884, FC. p. 716, pl. 101. figs. 4-7.
Baculogypsina baculata Silvestri, 1905, B. p. 69, fig. 2.
i spherulatus Cushman, 1921, FP. p. 359, pl. 75. fig. 6.
Extremely abundant ; attaining large dimensions and exhibiting great
variety in the number, arrangement, and development of the spines or lobose
processes. These frequently have a furcate appearance, but minute exami-
nation shows that this appearance is due to the close proximity of separate
processes. Some of the specimens are almost spherical, and in these, as

a rule, the processes are short and inconspicuous, though often occurring

in greater numbers than usual, and occasionally concentrated into closely

THE FORAMINIFERA OF LORD HOWE ISLAND. 639

juxtaposed groups. The finer material provided an abundant supply of
immature specimens, which when mounted in balsam exhibited a large
primordial chamber with chitinous lining, followed by a single coil of about
twelve globular chambers of gradually increasing size, in one plane. The
chambers in the conyolulion are separated by dense, solid shell-substance.
On completion of rather more than a single convolution, the chambers
become less spherical and are irregularly heaped over their predecessors as
in Gypsina. The spinous processes, perforated by secondary canals, are seen
to originate from a solid layer of shell-substance on the outer wall of the
primary coil. The general colour of the specimen is much less conspicuous
than in most ‘Tinoporus gatherings”? which have been examined. The
species generally gives a yellowish-orange tint to the sands in which it is
predominant, but the Lord Howe specimens are nearly white.

The complicated synonymy of this organism is fully discussed in the
papers quoted above, and in others referred to in those papers.

PoLytREMA isso.

187. Potyrrema mintaceum (Pallas).
Millepora miniacea Pallas, 1766, Elenchus Zoophytorum, p. 251.
Polytrema miniaceum Brady, 1884, FC. p. 721, pl. 100, figs. 5-9; pl. 101.
fig. 1.
“ a Heron-Allen & Harland, 1922, TN. p. 221, pl. 8; 1922,
FGA. p. 139, pl. 2 (numbered 1).
Curiously rare. The specimens call for no particular comment. A few
specimens of the free early stage were observed.

188. PoLyTRBMA MINIACEUM, var. ALBA Carter.
Polytrema miniaceum, var. album Carter, 1877, CB. p. 218, pl. 13. figs. 14, 15.

mn “ , alba Brady, 1884, FC. p. 721, pl. 101. figs. 2, 3.
35 a » » Heron-Allen & Harland, 1922, TN. 226.

As usual, rarer than the type, but unmistakable.

Family NUMMULINID Ai.
Sub-family PoLYSTOMELLINA.
Nontontwa d’ Orbigny.

189. Nonionrva DEPRESSULA ( Walker 5 Jacob).
Nautilus depressulus Walker & Jacob, 1798, AEM. p. 641, pl. 14. fig. 33.
Nonionina depressula Brady, 1884, FC. p. 725, pl. 109. figs. 6, 7.
Heron-Allen & Harland, 1914-15, EKA. p. 730.

” ”

Very rare and minute.

640 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

190. NoNIONINA UMBILICATULA (J/ontagu).
Nautilus umbilicatulus Montagu, 1803, TB. p. 191; Suppl. p. 78, pl. 18.
fig. 1.
Nonionina umbilcatula Brady, 1884, FC. p. 726, pl. 109. figs. 8, 9.
Heron-Allen & Harland, 1914-15, FKA. p. 730.

” 9

A single small specimen.

191. Nonronina pAuPERATA Balkwill 5 Wright.
Nonionina pauperata Balkwill & Wright, 1885, DIS. p. 353, pl. 13. figs. 25, 26.
Heron-Allen & Earland, 1908, etc., SB. 1911, p. 342,
pl. 11. figs. 16, 17; 1914-15, FKA. p. 732.

” ”

Frequent and typical.
PotystomMELLA Lamarck.

192. PoLysToMELLA sTRIATO-PUNCTATA (fiehtel § Moll).
Nautilus striato-punctatus Fichtel & Moll, 1798, TM. p. 61, pl. 9. figs. a—c.
Polystomella striato-punctata Brady, 1884, FC. p. 733, pl. 109. figs. 22, 23.
ey f Heron-Allen & Farland, 1914-15, FKA. p. 733. ”

A single small but typical specimen.

193. POLYSTOMELLA cRISPA (Linné).
Nautilus crispus Linné, 1788, p. 3370, no. 3.
Polystomella crispa Brady, 1884, FC. p. 736, pl. 110. figs. 6. 7.
a » Heron-Allen & Earland, 1914-15, FKA. p. 733.

Not uncommon, but small and weakly developed.

194. PoLySTOMELLA MACELLA (Lichtel § Moll).
Nautilus macellus Fichtel & Moll, 1798. TM. p. 66, pl. 10. figs. e-g.
Polystomella macella Brady, 1884, FC. p. 737, pl. 110. figs. 8, 9, 11.
»» Heron-Allen & Karland, 1914-15, FKA. p. 734.

Frequent and typical.

195. PoLysromELia mituerti Heron-Allen § Harland.
? Polystomella verriculata Millett, 1898, etc., FM. 1904, p. 604, pl. 11. fig. 3.
milletia Heron-Allen & Harland, 1914-15, FKA. p. 735, pl. 53.
figs. 38-42.

A single small and starved but unmistakable specimen.

9

Sub-family NUMMULITINA,
AMPHISTEGINA d’ Orbigny.

196. AMPHISTEGINA LESSONII d? Orbigny.
Amphistegina lessonii d’Orbigny, 1826, TMC. p. 304, no. 3, pl. 17. figs. 1-4;
Modeéle, no. 98.
Brady, 1884, FC. p. 740, pl. 111. figs. 1-7.
Heron-Allen & Earland, 1914-15, FKA. p. 736.

” ”

THE FORAMINIFERA OF LORD HOWE ISLAND. 641

Common, and presenting all the usual modifications of biconvexity. The
specimens are for the most part strongly developed. No specimens of the
tuberculate varieties, so abundant in some tropical gatherings, were seen.

197. AMPHISTEGINA LESSONII, var. GIBBA d’ Orbigny.
Amphistegina gibba d’Orbigny, 1826, TMC. p. 304, no. 6.
Ry mamillata d’Orbigny, 1846, FEV. p. 208, pl. 12. figs. 6-8.
3 lessonvi, var. gibba Heron-Allen & Harland, 1914-15, PKA.
p- 737.
This variety is fairly frequent and typical. It is characterized by its
marked plano-convexity.
OPERCULINA a? Orbigny.
198. OPERCULINA COMPLANATA (Defrance).
Lenticulites complanata Defrance, 1822, Dict. Sci. Nat. vol. xxv. p. 453.
Operculina * d’Orbigny, 1826, TMC. p. 281, no. 1, pl. 4. figs. 7-10;
Modeéle, no. 80.
Brady, 1884, KC. p. 743, pl. 112. figs. 3-6, 8.
i a Heron-Allen & Harland, 1914-15, FICA. p. 737.
Extremely rare ; represented by two moderate-sized water-worn specimens
and one very minute individual. The rarity of this form is noteworthy.

9 2

Hererostecina d’ Orbigny.
199, HETEROSTEGINA DEPRESSA dl’ Orbigny.
Heterostegina depressa @Orbigny, 1826, TMC. p. 305, pl. 17. figs. 5-7;
Modéle, no. 99.
Brady, 1884, KC. p. 746, pl. 112. ee 14-20..
Heron-Allen & Harland, 1914-15, PKA. p. 738.

” ”
” ”

Rare, but large and well-developed.

List of Works referred to in Abbreviated References in this Report.

B. 1837, LG. HH. G. Browy.—Lethiea Geognostica. Stuttgart, 1837-8.

B. 1855, FSH. J. G. Bornemann.—Die mikroskopische Fauna des Septarienthones von
Hermsdorf bei Berlin. Zeitschr. d. Deutsch. geol. Ges. vol. vil. pp. 807—
37], pls. 12-21.

B. 1864, RES. H. B. Brapy.—On the Rhizopodal Fauna of the Shetlands. Trans. Linn.
Soe. (London), vol. xxiv. pp. 463-475, pl. 48.

B. 1870, FTR. G. S. Brapy, D. Rosrrrson, & H. B. Brapy.—The Ostracoda and
iceman of Tidal Rivers. Ann. & Mag. Nat. Hist. ser. 4, vol. vi.
pp. 273-806, pls. 11, 12.

B., P., & J. 1870, GP. H. B. Brapy, W. K. Parkur, & T. R. Jonzs.—A Monograph of
the Genus Polymorphina, Trans. Linn. Soc. (London), vol. xxvii.
pp. 197-253, pls. 89-42.

B. 1879, ete., RRC. H. B. Brapy.—k eticularian Rhizopoda of the ‘Challenger’ Expe-
dition. Q. Journ. Micr, Sci. n.s. vol. xix. pp. 20-63, pls. 8-5; pp. 261-
299, pl. 8; vol. xxi. 1881, pp. 87-71,

642 MESSRS. BE. HERON-ALLEN AND A. EARLAND ON

B

B.

Q

F.

. 1884, FC. H. B. Brapy.—Scientific Results of the Voyage of H.M.S. ‘Challenger’
(Zoolegy), vol. ix. Report on the Foraminifera. 2 vols. 4to. Text and
Plates. London, 1884.

& W. 1885, DIS. F. P. Batxwrtt & J. Wricur.—Report on some Recent Fora-
minifera found off the Coast of Dublin and in the Irish Sea. Trans. Roy.
Trish Acad. vol. xxviii. (Science), pp. 317-868, pls, 12-14.

. 1856, ete,, RF. W. B. Canpryrnr.—Researches on the Foraminifera. Phil. Trans.

Roy. Soe. pt. i. 1856, elvi. pp. 1812-36; pt. 11. ibid. pp. 647-669 ; pt. iii.
1859, vol. exlix. pp. 1-41; pt. iv. 1860, pp. 585-594.

.P. & J. 1862, ISF. W. B. Carpenter, W. K. Parker, & T. R. Jonzs.—Introduction

to the Study of the Foraminifera. London, 1862. (Ray Society.)

. 1877, CB. H. J. Carrrr.—On the Locality of Canpenteria balaniformis, with Descrip-
tion of a new Species, etc. Ann. & Mag. Nat. Hist. ser. 4, vol. xix.
pp. 209-219, pl. 18.

. 1880, ete., SGM. H. J. Canrer.—Report on. Specimens dredged up from the Gulf of

Manaar. Ann. & Mag. Nat. Hist. ser. 5, vol. v. pp. 487-457; vol. vi.
pp. 85-61, 120-156. Supplementary Report. Ibid. vol. vii. pp. 861-385.

. 1892, FS. G. W. Cuasrer.—Report upon the Foraminifera of the Southport Society of

Natural Science District. First Rep. Southport Soe. of Nat. Sci. 1890-91
(Southport, 1892), pp. 54-72, pl. 1.

. 1899, FFA. F. Cauapman.—On some New and Interesting Foraminifera from the

Funafuti Atoll, Ellice Islands. Journ. Linn. Soc. London (Zoology),
vol. xxviii. (1902), pp. 1-27.

. 1901, FFA. F. Cuapman.—On the Foraminifera collected round the Funafuti Atoll

from shallow and moderately deep water. Journ. Linn. Soc. London
(Zoology), vol. xxviil. (1902), pp. 379-417, pls. 35, 36.

. 1907, RFV. F. Caapman.—Recent Foraminifera of Victoria; some Littoral Gatherings.
Journ, Quekett Micr. Club, ser. 2, vol. x. (1909), pp. 117-146.

. 1909, SNZ. F. Crapman.—Report on the Foraminifera from the Sub-Antarctic Islands

of New Zealand. In ‘Sub-Antarctic Islands of New Zealand,’ Welling-
ton, N.Z., 1909, pp. 312-371.
. 1910, ete., FNP. J. A. Cusuman. A Monograph of, the Foraminifera of the North
Pacific Ocean. Smithsonian Inst. U.S. Nat. Mus. Bull. 71, pt. 1, 1910;
pt. 2, 1911; pt. 8, 1913; pt. 4, 1914; pt. 5, 1915; pt. 6, 1917.

. 1921, FP. J. A. Cusoman.—¥oraminifera of the Philippine and adjacent Seas. Bull.

U.S. Museum, No. 100, vol. iv. Washington, D.C., 1921.
. 1831, CCT. G. P. DrsHayes.—Coquilles caractéristiques des terrains, Paris, 1831.
. 1841, SNA. C. G. Eurenpere.—Verbreitung und Ninfluss des mikroscopischen Lebens
in Sud- und Nord-Amerika. Abh, d. kel. Ak. d. Wiss. (Berlin), 1843
(for 1841), pp. 291-446, pls. 1-4, and Bericht, pp. 139-142, with
Appendix, pp. 202-209.
. 1893, FG. J. G. Eacur.—loraminiferen aus Meeresgrundproben, gvelothet yon 1874
bis 1876, von S.M.Sch. ‘Gazelle’ Abh. d. kel. bayerisch Ak. d. Wiss.
(Munich), ii. Cl. vol. xviii. pt. 2, pp. 195-458, pls. 1-21.
. 1905, FBS. A. Eartanp.—The Foraminifera of the Shore Sand at Bognor, Sussex.
Journ. Quekett Micr. Club, ser. 2, vol. ix. no. 57, pp. 187-282.
.& M. 1798, TM. L. von Ficuren & J. P. C. von Moxu.—Testacea microscopica ete.
Vienna, 1798.
. 1899, RFA. J. M. Frinr.—Recent Foraminifera. A Descriptive Catalogue of Speci-
mens dredged by the U.S. Fish Commission Steamer ‘ Albatross,’
Washington, U.S.A., 1899. :
1905, SOM. OC. Fornastnt.—Illustrazione di Specie Orbignyane di Miliolidi istituite nel
1826. Mem, Ac. Sci, Ist. Bologna, ser. 6, vol. ii. pp. 59-70.

THE FORAMINIFERA OF LORD HOWE ISLAND. 643

F, 1908, SON. C. Fornasint.—lIllustrazione di Specie Orbignyane di Nodosaridi, etc.,
istituite nel 1826. Mem. Ac. Sci. Ist. Bologna, ser, 6, vol. v. pp. 41-54.

G. 1882, RRCS. A. Goiis.—On the Reticularian Rhizopoda of the Caribbean Sea. K.
Svenska Vet.-Ak. Handl. vol, xix. no. 4 (Stockholm, 1882), pp. 1-151,
pls. 1-12. :

G. 1894, ASF. A. Goiis.—A Synopsis of the Arctic and Scandinavian recent Marine
Foraminifera hitherto discovered. K. Svenska Vet.-Ak. Handl. (Stock-
holm), vol. xxy. no. 9.

G. 1896, DOA. A. Goiis.—Reports on the Dredging Operations carried on by the U.S.
F.-C.5. ‘ Albatross,’ ete. Bull. Mus. Comp. Zool. Harvard, vol. xxix.
no. 1. Cambridge, Mass., 1896.

G. 1906, FLL. G. C. Goueu.—The Foraminifera of Larne Lough and District. Dep. of
Agriculture, etc. Fisheries, Ireland, Sci. Invest. 1905, no. 3 (1906).

H.-A, & BE. 1908, ete., SB. [. Heron-Atten & A. Eartanp.—The Recent and Fossil

* Foraminifera of the Shore Sands at Selsey Bill, Sussex. Journ. R.

Micr. Soc. 1908, pp. 529-543; 1909, pp. 806-336, 422-446, 677-698 ;
1910, pp. 401-426, 693-695; 1911, pp. 298-348, 4386-448.

H.-A. & E. 1913, Cl. E. Heron-Ariten & A, Eartann.—Clare Island Survey, pt. 64.
Foraminifera. Proc. R. Irish Academy, vol. xxxi. pt. 64.

H.-A. & E, 1914, FKA. E. Heron-Atien & A. Hartanp.—On the Foraminifera of the
Kerimba Archipelago, ete. Trans. Zool. Soc. London, vol. xx. pt. i.
pp. 363-390, pls. 35-57; vol. xx. pt. il. pp. 543-794, pls. 40-43.

H.-A. 1915, RPF. E. Heron-Arirn.—Contributions to the Study of the Bionomics and
Reproductive Processes of the Foraminifera. Phil. Trans. Roy. Soc.
(London) B, vol. cevi. 1915, pp. 227-279, pls. 18-18.

H.-A. & BE. 1916, FWS. E. Herox-Atten & A. EARLAND.—Foraminifera of the West
Coast of Scotland. ‘Trans. Linn. Soc. London, ser. 2 (Zoology), vol. xi.
pp- 197-800, pls. 389-48.

H.-A. & E. 1922, TN. E. Heron-Atren & A, HARLAND.—British Antarctic (‘Terra
Nova’) Exp. 1910, Zoology, vol. vi. No. 2, Protozoa, Pt. ii. Foraminifera,
pp. 20-284, pls. 1-8. British Museum (Nat. Hist.).

H.-A. & EH. 1922, FGA. E. Hrron-Arien & A. Eartann.—Foraminiféres des Sables
Rouges du Golfe d’Ajaccio. Bull. Soc. Sci. Hist. et Nat. de la Corse.
Bastia (Corsica), pp. 109-149, pls. 1, 2 (numbered 2, 1).

J. & P. 1860, RFM. T. Ruperr Jones & W. K. Parker.—On the Rhizopodal Fauna of
the Mediterranean compared with that of the Italian and some other
Tertiary Deposits. Q. Journ. Geol. Soc. vol. xvi. pp. 292-307, and table.

J., P., & B. 1866, ete, MFC. T. R. Jonus, W. K. Parker, H. B. Brapy (and others).—
A Monograph of the Foraminifera of the Crag. London, 1866-1897,
(Paleontographical Soc.), pt. 1. 1866; pt.ii. 1895; pt. iii. 1896; pt. iv.
1897.

K, 1867, FO. F. Karrer.—Zur Foraminiferenfauna in Osterreich. Sitz. Ak, Wiss. Wien,
vol. ly. Abth. i. pp. 331-368, pls. 1-8.

K. 1868, MFKB. F. Karrer.—Die Miocene Foraminiferenfauna yon Kostej] im Banat.
Thid. vol. lviii. Abth. i. pp. 111-1938, pls. 1-5.

L, 1767-1788, SN. C.von Linné. Systema Nature. Ed. XII. Leipzig, 1767; Ed. XIII.

; (a J. F. Gmelin) Leipzig, 1788-93.

L. 1804, ete, AM. J. B. pr Lamarck.—Annales du Muséum (Paris), vol. v. 1804;
yol. vill. 1806; vol. ix. 1807.

L. 1830, ete., EM. J. B. pe Lamarck.—Eneyclopédie Méthodique. “ Vers,” vol. ii. 1830.

‘M, 1803, TB. G. Monracu.—Testacea Britannica, or Natural History of British Shells.
In 8 yols, 4to, London, 1803. Supplement (Plates), 1808,

644 MESSRS. E. HERON-ALLEN AND A. EARLAND ON

M. 1808-10, CS. D. pk Montrorr.—Conchyliologie Systématique etc., 2 vols, Paris,
1808-10.
M. 1898, ete., FM. F. W. Mirterr.—Report on the Recent Foraminifera of the Malay
Archipelago contained in Anchor-Mud, collected by Mr. A. Durrand,
F.R.M.S. J. R. Mier. Soc. 1898-1904.
v’0. 1826, TMC. A. D. p’Ornieny.—Tableau Méthodique de la Classe des Céphalopodes.
Ann. Sci. Nat. (Paris), vol. vii. pp. 245-814, pls. 10-17.
vO, 1826 (Modéles). A. D. p’OrBrany.—Modéles de Céphalopodes Microscopiques vivants
et fossiles. Paris, 1826 (2nd issue, with new Catalogue, 1848).
pO. 1889, FC. A. D. p’OrBigny.—Foraminiféres. In R. de la Sagra: Hist. Phys. Pol. et
Nat. de l'Ile de Cuba. Text, 8vo, plates, fol. Paris, 1839.
p’O. 1889, FAM. A. D. p’OrBrany.—Voyage dans l’Amérique Méridionale, vol: v. pt. 5.
Foraminiféres. Paris, 1839.
pO. 1889, FIC. A. D. D’ORpiawy.—Foraminiferes. In Barker-Webb & Berthelot, Hist.
Nat. des Iles Canaries, vol. ii. pt. 2, pp. 119-146, 3 pls. Paris, 1839.
vO, 1840, CBP. A. D. p'OrpigNy.—Mémoire sur les Foraminiféres de la Craie Blanche
du Bassin de Paris. Mém. Soc. Géol. France, vol. iy. pp. 1-41, pls. 1-4.
pO. 1846, FFV. A.D. p’Orprieny.—Foraminiféres Fossiles du Bassin Tertiaire de Vienne.
4to, 21 plates. Paris, 1846.
vO, 1850, ete., PP. A. D. p’Orprany.—Prodrome de Paléontologie stratigraphique
uniyerselle des Animaux mollusques et rayonnés. Paris. Vol. i. 1849;
vol. 11. 1850; vol. iii. 1852.
P. 1858, MIS. W.K. Parxkrr.—On the Miliolitidee of the East Indian Seas. Pt. i.
Miliola, Trans. Micr. Soc. Lond., n. s., vol. vi. pp. 58-59, pl. 5 & figs,
P. & J. 1859, etc., NF. W. IK. Parker, T. R. Jones, and others.—On the Nomenclature
of the Foraminifera, Ann. & Mag. Nat. Hist. 1859 to 1872. Pts. 1-16.
P.& J. 1865, NAAF. W.K. Parker & T. R. Jones.—On some Foraminifera from the
North Atlantic and Arctic Oceans, ete. Phil. Trans. Roy. Soc. (London),
vol. cly. pp. 325-441, pls. 12-19. e
R., 1849-50, FOT. A.E. Reuss. Neue Foraminiferen aus der Schichten des vesterreich-
ischen Tertiirbeckens. Denkschr. Math. Nat. Kl. d. k. Ak. Wiss. Wien,
vol. i. 1850, pp. 865-890, pls. 45-51 *.
R. 1845-46, VBK, A. E. Reuss.—Die Versteinerungen der bdhmischen Kriedeformation
Stuttgart, 4to, 1845-46.
R. 1862, NHG. A. EH. Reuss.—Die Foraminiferen des Norddeutschen Hils und Gault.
Sitzb. Ak. Wiss. Wien, vol. xlvi. Abth. i. (1863), pp. 5-100, pls. 1-18.
1866, FABS. A. E. Rrvuss.—Die Foraminiferen, Anthozoen, und Bryozoen des
Deutschen Septarienthones. Denkschr. Ak. Wiss. Wien, vel. xxv.
(1865), pp. 117-214, pls. 1-11.
R. 1869, FOG. A. E. Reuss.—Zur Fossile Fauna der Oligocinschichten yon Gaas. Sitzb.
Ak. Wiss. Wien, vol. lv. Abth. i. pp. 17-182, pls. 1-8.
. 1906, FLC. L. Ruaumprirr.—Foraminiferen von Laysan und den Chatham Inseln,
Spengel’s Zoologischer Jahrbticher, vol. xxiv. pt. i. pp. 21-80, pls. 1-5.
8. 1854,OP. M.S. Scurunrzr.—Ueber den Organismus der Polythalamien (Foraminiferen)
nebst Bemerkungen iiber die Rhizopoden in Allgemeinen. Leipzig,
1854.

R.

a

es)

* This paper is variously referred to as being dated 1849 and 1850. It was read in May
1849, and all the separate copies were dated on a special title-page 1849, but the volume of
which it forms part was issued in 1850, and is so dated,

THE FORAMINIFERA OF LORD HOWE ISLAND. 645

. 1874, ete, R. F. KE. Scpunze.—Rhizopodenstudien. Archiv fiir Mikr. Anat. nos. 1, 2,

1874; nos, 3-5, 1875; no. 6, 1876.

. 1891, BGR. C. ScurumperGcer.—Révision des Biloculines des Grands Fonds. Mém.

Soc. Géol. France, vol. iv. pp. 542-579 (pp. 155-198 in the Reprints),
pls. 9-11.

. 1893, MGM. C. ScurumBerGEr.—Monographie des Miliolidées du Golfe de Marseille.

Mém. Soe. Zool. France, vol. vi. pp. 57-80 (pp. 199-228 in the Reprints),
pls. 2-4.

. 1904, ete., RFD. H. Stprsotrrom.—Report on the Recent Foraminifera from the Coast

of the Island of Delos (Grecian Archipelago). Memoirs of the Man-
chester Literary and Philosophical Society (Manchester), pt. 1, 1904;
pt. 2, 1905; pt. 3, 1906; pt. 4, 1907 ; pt. 5, 1908; pt. 6, 1909.

. 1905, B. A. StrvestRr.—Osservazioni critiche sul genere Baculogypsina Sacco, Att.

R. Ace. Nuovi Lincei, 1905, pp. 65-82, text-figs.

. 1910, RFBP. HH. Siprsotrom.—Report on the Recent Foraminifera from the Bay of

Palermo (Sicily), 14-20 fms. Mem. & Proc. Manchester Lit. & Phil. Soc.
vol. liv. pt. 3, no. 16.

. 1912, ete., LSP. H. Stprporrom.—Lagene of the South-west Pacific Ocean, H.M.S.

‘ Waterwitch,’ 1895. Journ. Quekett Micr. Club, ser. 2, vol. xi. no. 70,
pp. 375-434, pls. xiv.-xxi. Supplementary Paper: ib. ser. 2, vol. xii-
1913, no. 73, pp. 161-210, pls. xv.—xviil.

T. 1878, FIR. O. Trrquem.—tLes Foraminiféres ete. du Pliocéne Supérieur de I’'Ile de
Rhodes. Mém. Soc. Géol. France, sér. 3, vol. i. Mém. iii.
T. 1882, FEP. O. Terquem.—Les Foraminiféres de l’Eocéne des Environs de Paris. Mém
Soc. Géol. France, sér. 3, vol. ii. Mém. iii.
W.& B. 1784, TMR. G. Watker & W. Boys.—Testacea Minuta Rariora ete. London,
: 1784, 3
W.& J. 1798, AEM. G. Watxker & E. Jacos.—In G. Adams, Essays on the Microscope,
2nd Ed. London, 1798.
W. 1848, BSGL. W.C. Wittramson.—On the Recent British Species of the Genus
Lagena. Ann. & Mag. Nat. Hist. ser. 2, vol. i. pp. 1-20, pls. 1-2.
W. 1858, RFGB. W. C. Witrramson.—On the Recent Foraminifera of Great Britain.
London, 1858 (Ray Soc.).
W. 1877, RFDA. J. Wricur.—Recent Foraminifera of Down and Antrim. Proc. Belfast
Naturalists’ Field Club, 1876-77, Appendix iv. pp. 101-108, pl. 4.
W. 1885-86, BLP. J. Wrieut.--Foraminifera of the B.N.F.C. Cruise off Belfast Lough,

June 1885; also, Foraminifera found by Dr. Maleomson at Rockport,
Belfast Lough. Rep. Belfast Nat. Field Club, 1885-6, Appendix,
pp. 317-326, pl. xxvi.

LINN. JOURN.— ZOOLOGY, VOL. XXXV. 46

646

15.

14-16.

. Bolivina limbata, var. abbreviata, nov. Side views.

MESSRS. E. HERON-ALLEN AND A. EARLAND ON

EXPLANATION OF THE PLATES.

PLATES 35-37.

. Nubecularia lucifuga Defrance. Showing young individuals in fractured chamber.

x 55.
schauinslandi (Rhumbler). x 35.

”

. Spiroloculina tenuirostra Warrer. Fig. 6, Central portion abraded. x 145.
. Miliolina auberiana, var. semtreticulata, var. nov. Front view. \
is x 59.

i Hy Pe Back view.
” ” ” ” Oral view.

. Craterites rectus, gen. nov. et sp. nov. Side view, showing the expanding base,

layers of chamberlets, and cribrate oral surface. 30.

55 5 gen. noy. et sp. nov. Superior view of cribrate oral surface,
showing similar structure underneath through fractures on
surface-layer. xX 30.

Diffusilina humilis, gen. noy. et sp.nov. A perfect specimen attached to Nullipore,

showing pustular processes on the surface, which is otherwise
un-abraded. x 28.

- » gen. nov. et sp. noy. Specimens showing partially abraded
surface, exposing the labyrinthic protoplasmic structure
(shown black). 28.

. Haddonia torresiensis Chapman. Young textularian specimen, showing commence-

ment of wild later growth. x 30.

x oa . Fully-developed specimens, showing textularian
commencement to wild-growing portion. x 30.

a s i Showing the oral extremity. x 30.

e iw 5 Specimens incorporating large sand-grains. x30.

. Bulimina elegantissima VOrbigny. Fig. 23. The bud is a single chamber; Fig. 24,

developed into a young individual of several chambers. x 146.

Edge view. { x 145.

7 ” ” ”

28. Frondicularia scottii Heron-Allen & Earland. Specimen from Lord Howe Island.
x 145.
29. 5 * Specimen from ‘Challenger’ Stn.
185 (Raine Island). x 145.
30-31. Uvigerina selseyensis Heron-Allen & Earland. x 145.
82. Ramulina grimaldit Schlumberger. X 26.
33-34. Spirillina campanula, sp. nov. Side views.
35. 5 ~ Apical view} Viewed as opaque specimens.
36. Fo 3 ; Basal view.
37-38 a as “s Balsam mounts. Side views.

3 . om ) on Apical view. Scaly surface.

40. is iF A a of Basal view.

41. 5h a oD ss 4 Specimen in which the septa
haying been entirely absorbed, the central cavity is filled with a
mass of young individuals, which are shown in optical section.
All x 540.

42. Discorbina pulvinata Brady. Superior view.

43-45, 50 on Basal view, showing development of young indi-

viduals around oral aperture.

ORE Se

HERON-ALLEN & EARLAND.

FORAMINIFERA FROM

Journ Linn.Soc, ZOOL VoL. XXXV. Piss

IRD HOWE ISLAND.

HERON-ALLEN & EARLAND.

LORD HOWE ISLAND.

FORAMINIFERA FROM

HERON-ALLEN & EARLAND.

FORAMINIFERA FROM

Journ. Livny Soc, Z00L VoL XXXYV. PL. 36

LORD HOWE ISLAND.

‘Se

HERON-ALLEN & EARLAND.

338

37
FORAMINIFERA F pos “ORD HOWE ISLAND

OLXK XV. Pi.3a7

7,
{

RN LINN Soc, ZOoL V

JOU

HERON-ALLEN & EARLAND.

LORD HOWE ISLAND.

FORAMINIFERA FROM

,
ee

THE FORAMINIFERA OF LORD HOWE ISLAND. 647

Fig. 46. Discorbina pulvinata Brady. Basal view of specimen in which the internal septa
and central portion of the base have been absorbed
after process of reproduction. All x 145.

47-48, 7 tabernacularis Brady. Adult budded pairs. 145.
49, ) * 9 Basal view of specimen, showing young
individuals. (The marginal edge is broken away.) X 145.
KO A ey lauriei, nom. noy. Superior view.
51. . Basal (oral) view.
52-55. % 9” *s Specimens showing budding young in various stages
of development. All x 145.
56, ” pyramidalis, sp. nov. Side view, short (typical) form.
57. 0 ,; 0 5 Jong form.
58. =) on 5 Basal side view.
59. on on % Apical side view.
60. 7 a os Basal view.
61, Ss 15 i Budding pair. All x 145,

62. Gypsina inherens Schultze. Specimen showing young individuals in broken
chamber. xX 145.

63. 3 in Ks Specimen suggesting Holocladina (see text). X 55.
64, 3 me i Wild-growing specimen. X 26.

65. Baculogypsina spherulata (Parker & Jones). A young individual in balsam,
showing the transition from the early rotaline plan of growth
to the acervuline later stages. The rotaline chambers are
shown dark owing to the chitinous lining. Axial canals
perforate the spines and communicate with the primordial
chamber. x 105.

{Synonyms and native names are printed in ¢talics. A star * denotes the first

publication of a name; a dagger + denotes a fossil.

The absence of an

authority is due to omission by the respective authors. |

Aaptos aaptos, O. Schimdt, 479, 508.

— adriatica, Gray, 508.

Abrolhos Islands: Amphipoda and Isopoda,
1; Echinoderms, 229-251 ; Opistho-
branchiata, 521; sea-pens, 21; sponges,
477.

Acamarchis, Lamx., 297.

— dentata, Lamx., 297.

— neritina, Lamx., 297.

Acanthodoris pilosa, 569.

Acantholeberis curvirostris, Zilljeb., 425,
4357.

Acarapis woodi, Hirst, 382.

Acari, new species, 363-392.

Acarus marginatus, Herm., 366.

Acervulina inherens, Schultze, 637.

Acroperus harp, Baird, 423, 425.

Actheres, 90.

Actinodoris, Khrenb., 553.

— sponsa, Khrenb., 553.

Actinometra annulata, Bell, 235.

— nigra, P. H. Carp., 232.

— stelligera, P. H. Carp., 282.

Actinophrys, Ehrend., sp., 457, 473.

Adults, phyletic succession, 83.

Aetea, Lamu., 297, 308, 309, 316.

— anguina, 297.

— ligulata, Busk, 304.

— truncata, Landsb., 297, 309.

Aeteopsis, Boeck, 297.

— elongata, Boeck, 297.

Adthusa ciliatifrons, Fav., 47, 48.

Aathusina, 48.

— gracilipes, Miers, 47, 78.

Alcedo ispida, Zinn., its brilliant colouring
aposematic, 288.

Alecto parvicirra, J. Miller, 234.

Algex, food of Daphnia, 416.

Alicus rostratus, Triig., 883.

Allantion, Sandun*, 461; mentd., 452, 475.

— tachyploon, Sandon *, 461; mentd., 452,
457, 467.

Alloniscus, Dana, 107; four spp., 108.

—alluaudi, Dollf., 108.

— brevis, Budde-Lund, 108.

— elegans, Dollf., 108.

— guttatus, Dollf., 108.

— nacreus, Collinge *, 107, 108, 113.

— pallidulus, Budde-Lund, 108.

— pigmentatus, Budde-Lund, 108. |

— tigris, Dollf., 108.

Alloidoris, Bergh, 547.

—hedleyi, O’Don.*, 523, 547, 549, 550,
577, 579.

— lanuginata, Abrah., 549.

— marmorata, Bergh, 547, 548, 549, 550.

Alona affinis, Leyd., 425.

— rustica, 7. Scott, 425.

Alonella excisa, 425.

— nana, 425.

Alonopsis elongata, Sars, 414, 423, 42 4
425.

Altson, A. M., Method of oviposition and
the ege of Lyctus brunneus, Steph.
217-227; Genital system of Lyctus
brunneus, Steph., 581.

Alveolina, Ord., 612.

— melo, Heron-All. § Earl., 612,

650 INDEX.

Alysidium, Bush, 298.

— lafontii, Busk, 298.

— parasiticum, Busk, 298.

Amastigia, Bush, 298, 300, 825, 329, 339,
347, 358.

— abyssicola, 335.

— antarctica, 338.

— benemunita, 331, 334, 348.

— caberioides, 335.

— crassimarginata, 384.

— funiculata, 335.

— gaussi, 338.

— kirkpatricki, Levensen *, 295, 328, 335,
337, 338, 341, 358,.359, 361.

—nuda, Busk, 330, 331, 332, 338, 335,
359, 360, 361.

— pateriformis, 339.

— rudis, 332, 333, 384, 345, 360, 361.

— solida, 338.

Amblypneustes pallidus, Valenciennes, 249 ;
distrib., 229, 252.

Ameletus ornatus (Zaton), wing-veins,
145-160.

— perscitus, Haton, wing-veins, 153.

Ameeba actinophora, Awerb., 452, 457, 468,
470.

— diploidea, 457.

——limax, Duj., 457, 468.

— limicola, hwmbd., 469.

— radiosa, Ehrenb., 457.

—— sp., 457.

—— striata, Penard, 452, 469.

—~ terricola, Greef, 452, 457, 469.

— vespertilio, Penard, 452, 469.

Amorphina Duchassaingi, Tops., 508.

Amphimetra jacquinoti, A. H. Clark, 284 ;
distrib., 231.

Amphipoda from Abrolhos, 1.

Amphioxus, Yarr., 90. -

Amphistegina, Orb., 640.

— lessonii, Orb., 640.

— — var. gibba, Orb., 641.

— mamillata, Orb., 641.

Amphithoé setosa, Hasw., 12.

Anabzena, A. Juss., food of Daphnia, 416.

Anchinoé fictitioides, Dendy, 504, 479, 519.

— fictitius, J. Steph., 505.

Ancorina aaptos, O. Schmidt, 508, 509.

— adriatica, Vosm., 508. :

— australiensis, Lendenf., 475, 492.

— brevidens, Dendy *, 493, 478, 479, 519.

Andersonia, Boulenger, 298.

Anderssonia, Kluge, 298, 329.

— antarctica, Kluge, 298, 338.

Anderssonia, Strebel, 298.

Angasiella, Angas § Cross, 568.

Anguinaria, Liam., 298.

— spatulata, Lam., 298.

— truncata, Landsb., 504.

Angularia, Busk, 298.

Anisonema minus, Sandon*, 462, 475;
mentd., 452, 457.

Anisoptera, mentd., 143.

Antedon, Frem., 86, 93, 95, 96, 97.

— gyges, Bell, 234.

— serripinna, P. TH. Carp., 235.

Antenne of Chlorops, 406.

Antennophorus uhlmanni, Haller, 367.

Anthenea australiv, Doder/ein, 238; dis-
trib., 230, 232.

— globigera, Déderlein, 239; distrib., 230,
232.

— pentagonula, Perie, mentd., 239.

Antiopa cristata, 569.

Aphanizomenon, Morr., food of Daphnia,
416.

' Aphelodoris, Bergh, 556.

— affinis, Alot, 556, 558, 577, 579.

— antillensis, Bergh, 556.

Aplysia, Linn., 527, 528.

— denisoni, Smith, 533.

Apocheima, Schaeff., 88.

Apus cancriformis, Schaef, in Scotland,
420.

Avcella arenaria, Grreef, 452, 454, 471.

— vulgaris, Lhrenb., 452, 471.

Archidoris incerta, 570.

— tuberculata, 569.

Arhina, Collinge, 108.

Armadillidiun celatum, Miers, 103, 104.

— purpurascens, Kroy., 105.

Armadillo horridus, Budde-Lund, 108.

Armina cygnea, 525.

Arrhenurus insulanus, Koen., 888.

Arthropoda, mentd., 285.

Ascoleucetta, Dendy *, 477.

— compressa, Dendy *, 481, 477, 518.

Asexual reproduction in Ptychodera, 393.

Assulina minor, Penard, 472.

— muscorum, Gireef, 453, 457, 472.

— seminulum, Leidy (pars), 472.

Astacus, Gron., 35, 39.

INDEX. 651

Asterias pectinata, Linn., 238.

Asterina, Nardo, mentd., 244.

— burtonii, Gray, 243; distrib., 232.

— gunnii, Gray, 243; distrib., 229-230.

Asteroidea from West Australia, 235-245.

Asteronotus, Ehrenb., 550.

— bertrana, 570.

— fuscus, O’Don.*, 551, 523, 552, 577,
579.

— hemprichii, Lhrend., 550.

Asteropus simplex, Dendy *, 478, 494.

Astropecten preissii, Miiller § Troschel,
235 ; distrib., 229-30.

Aurora rowi, Dendy, 478, 494.

Australia (West), see Echinoderms.

Avicella, Van Bened., 298.

Avicularia, internal, 318.

Avicularia, Zam., mentd., 298.

Avicularia, Gray, 298, 314, 315.

— flabellata, Gray, 298, 299.

Bactridinm, Reuss, 298.

— hagenowi, Zeuss, 298.

Baculogypsina, Sacco, 638.

— baculata, Silv., 638, 647.

— spherulatus, Cushm., 6388.

Balanoglossus, Delle Ch., 393.

Balantiophorus minutus, Schew., 458, 457.

Barentsia, Hincks, 315.

Basipharynx of Chlorops, 409.

Bathycrinus, Thoms., 95.

Bathydoris, 547.

Beania, Johnst., 298, 304, 306.

— mirabilis, Johnst., 298.

Bees, their mental qualities influence the
evolution of flowers, 287.

Benthesicymus altus, 45.

Berthella, Blainv., 536.

— plumula, 525, 536, 538, 578.

— porosa, Blainy., 536.

Bicellaria, Macg., 299.

Bicellaria, Blainv., 298-300.

— aculeata, Orb., 817, 355.

— aldert, Busk, 299.

— annulata, Mapl., 317.

— ciliata, Blainy., 298, 299.

— grandis, Busk, 306.

— infundibulata, Busk, 306.

Bicellariella, Zev., 299.

— ciliata, Zev., 299.

Bicellarina, Zev., 299.

— alderi, Zev., 299.

Biemna mucrova, Hentsch., 503.

— tubulata, Dendy, 479, 5038.

Bifrons, MacGill., 299.

Bifurcation of Menipea and Scrupocellaria,
321; — of Polyzoa, 320.

Biloculina, O7d., 602.

— depressa, Ord., 602.

— — var. serrata, Bavl., 602.

— elongata, Orb , 603.

— oblonga, Orb., 603.

— opposita, Desh., 603.

— ringens var. striolata, Brady, 602.

— sarsi, Schlumb., 602.

Bimichaelia crassipalpis, Halb.*, 383, 392.

Biogenetic Law restated, 81-101.

Bird-of-Paradise, courtship activities, 278.

Birds, courtship activities in various kinds
of, 253-292.

Bittacomorpha, Westw., 409.

Blackbird, courtship activities, 263.

Biackeock, courtship activities, 270, 277, 287.

Black locust tree, mentd., 217.

Blepharisma laterita, Ehrenb., 453, 457.

Blue Tit, courtship activities, 287.

Bolivina, Or., 621.

— dilatata, Reuss, 621.

— inflata, Heron-All. § Earl., 622.

— limbata, Brady, 622.

— — var. abbreviata, Heron-All. § Earl.*,
622:

— plicata, Ord., 622.

— punetata, Ord., 621.

— robusta, Brady, 621.

— textilaricides, Reuss, 621.

— tortuosa, Brady, 621.

— variabilis, Heron-All. § Earl., 622.

Borradaile, L. A., Mouth-parts of Shore
Crab, 115-142.

Bosmina brevicornis, Hellich, 446.

— coregont, Baird, 444.

— coregoni-longispina, Riihe, 444,

— longirostris, Baird, 425, 446; distrib.,
414, 418, 424.

— obtusirostris, Sars, 421, 422, 423, 425,
443-445 ; distrib., 414, 415, 418.

Bourne, G. C., Raninidz, 25-79.

Bower-bird, courtship activities, 269.

Brachythecium salebrosum, Schimp., 457.

Branching of Bugula, 321, 328, 324.

652 INDEX.

Brephos parthenias, 86.

Brettia, Dyst., 299; mentd., 300.

— pellucida, Dyst., 299.

Breynia australasie, Gray, 251; distrib.,
229, 232.

Briarella microcephala, 570.

— sp., 570.

Bryobia humeralis, /Za/b.*, 385, 392.

— preetiosa, 385.

Bryum, Dill., spp., 450.

Bueula, Oken, 298, 299, 306, 311, 314, 315,
326; its branching, 321, 323, 324;
mentd., 300.

— abyssicola, Kluge, 300.

— angusta, Kluge, 300.

— arevlata, Kluge, 300.

— avicularia, Oken, 299.

— ciliata, Oken, 299.

— dentata, 826, 359.

— flabellata, 299.

— gigantea, Kluge, 300.

— johnston, 326, 359.

— lata, Kluge, 300.

— lewaldit, Kluge, 300.

— leontodon, Busk, 311.

— margaritifera, Busk, 311.

— multispinosa, Kluge, 300.

—- murrayana, 309.

— neritina, Oken, 299,

— retiformis, Kluge, 300.

— scaphoides, Avrkp., 326, 359.

— stnuosa, Busk, 31], 512.

— — var. variabilis, Kluge, 312.

— sp. var. vartospinosa, 300.

— tricornis, Waters, 300.

— versicolor, Busk, 512.

Bugularia, Zev., 299.

— dissimilis, Lev., 299.

Bugulella, Verr., 300.

Bugulina, Gray, 298, 800, 314.

— avicularia, Gray, 298, 300.

— flabellata, 298.

— neritina, 298.

Bugulopsis, Verr., 300, 353.

— peachii, Verr., 800, 321.

Bulimina, Ord., 620,

— elegantissima, Orb., 620, 646.

—limbata var. abbreviata, Heron-All. §
Earl.*, 646.

— polystropha, Reuss, 619,

— williamsoniana, Brady, 621.

Bulla plumula, Mont., 586.

Bunaster lithodes, W. K. Fisher, 2415
distrib., 232.

— ritteri, mentd., 241.

— uniserialis, mentd., 241.

Bunting, courtship activities, 263, 277.

Bustard, courtship activities, 277.

Bythotrephes longimanus, Leyd., 421 ;
distrib., 414, 418.

Caberea, Lamex., 300, 309, 312, 329.

— benemunita, Ley., 354.

— boryi, Aud., 516.

— crassimarginata, Busk, 334, 337.

— dichotoma, Lamz., 300.

— marvonensis, Ley., 337.

— pinnata, Lamw., 300.

— rudis, Busk, 332, 337.

Caberiella, Lev., 300, 329.

— benemunita, Lev., 300.

Calappa, Fabr., 31, 40, 58.

Calligonus scapularis, Berl., 387.

Callitriche, Zznn., in tarns, 423, 424.

Calmanesia, Collinge *, 107, 109.

— methueni, Collinge*, 109-118.

Campascus triqueter, 453.

Camptoplites, Harmer *, 295, 300.

— abyssicola, Harmer *, 300.

— angusta, Harmer *, 300.

— areolata, Harmer *, 300.

— bicornis, Harmer *, 300.

— gigantea, Harmer *, 300.

— lata, Harmer *, 500.

— lewaldii, Harmer *, 300.

— multispinosa, Harmer*, 300.

— retiformis, Harmer *, 300.

— sp. var. variospinosa, Harmer *, 300.

— tricornis, Harmer *, 300.

Camptothecium nitens, Schimp., 450.

Cancer, Linn., 129.

Canda, Lamx., 300, 329.

— arachnoides, Lam.., 300.

Candiella, Gray, 548.

Canthocamptus cuspidatus, Schmez, 422,
424,

— pygmaeus, G. O. Sars, 424.

— zschokkei, Schmezl, 424, 425.

Carabodes affinis, Berl., 380.

— marginatus, Mich., 380.

Caradrina, Hued., mite on, 587.

INDEX. | 653

Carbasea, Gray, 301, 311.

— armata, Busk, 312.

— bombyeina, Busk, 316.

— carbasea, Harmer *, 301.

— dissimilis, Busk, 299.

— episcopalis, Busk, 308.

— hgulata, Busk, 314.

—- papyracea, Gray, 301.

— solander?, Norman, 301.

Carchesium, Ehrenb., mentd., 450.

Carcinology, Raninide, 25-79.

Carcinus meenas, Leach, 35, 40; mouth-
parts, 115-142.

Carterina, Brady, 616.

— spiculotesta, Brady, 616.

Cassella, H. & A. Adams, 554.

Cassidulina, Ord., 622.

— clavata, Brady, 623.

— crassa, Orb., 622.

— subglobosa, Brady, 623.

Catenaria bizornis, Busk, 315.

Catenicella, 320.

Caulibugula, Verr., 301.

— armata, Verr., 301,

Cellarta (pars), 348, 353.

Cellaria, Hil. § Sol., 301, 303, 304, 308,
315,

— cirrata, Ell. § Sol., 801, 313.

— cornuta, 307.

— furciminordes, Ell. & Sol., 301, 302.

— fistulosa, Hincks, 301-308.

— fistulosa, Hincks, 303.

-- flabellum, Il. & Sol., 313.

— johnsoni, Busk, 303.

— loricata, 307.

— pyriformis, Bertol., 305, 306.

— salicornia, Lamx., 301.

— salicornia, Lam., 301.

— salicornia (pars), Pall., 301, 308.

—- salicornia, Padll., 302, 305.

— salicornioides, Lamx., 303, 515.

— sinuosa, Zll., 302.

— sinuosa, Hincks, 303.

— tenella, Lam., 314.

— ternata, Ell. & Sol., 317, 354.

Cellarina, Orb., 304.

Cellurina, Van Bened. (pars), 353.

Cellarina, Van Bened., 308, 304.

— gracilis, Van Bened., 308, 304.

— seabra, Van Bened., 303.

Cellularia (pars), auctt., 353.

Cellularia (pars), Busk, 339.

Cellularia, Pall., 30), 304, 320.

-— avicularia, Pall., 314.

— biloba, Busk, 350.

— cirrata, Busk, 508.

— errata, Ell. & Sol., 340.

— crateriformis, Busk, 350.

— crispa, Pall., 313, 340.

— cuspidata, Busk, 355.

— elonyata, Busk, 350, 351.

— farciminoides, Hl. & Sol., 303.

— flabellum (pars), Ell, & Sol., 340.

— floccosa, Pall., 540.

— loriculata, Pall., 310, 314.

— monotrypa, Busl, 355.

— ornata, Busk, 340.

— peachti, Busk, 300, 355.

— plumosa, Pall., 306.

— quadrata, Busk, 542.

— salicornia, Pall., 303, 315.

— seruposa, Verr., 304.

— ternuta, 304.

— — forma gracilis, Smitt, 354.

Cellularine Polyzoa, 293-361.

Centrechinus savignyi, H. LZ. Clark, 248;
distrib., 232.

— setosus, Jackson, 248; distrib., 232.

Centropyxis, Stein, 457, 470.

—aculeata, Stern, 454, 471.

— — var. spinosa, Cash, 471.

— — var. ecornis, Leidy, 471.

— levigata, Penard, 454, 470.

Centrostephanus tenuispinus, H. L. Clark,
249; distrib., 230, 232.

Cephalodiscus, MeInt., 393.

Ceradocus, A. Costa, 6.

— rubromaculatus, (Stebb.), 2, 6, 18.

Ceraochalina multiformis yar. manaarensis,
Dendy, 478, 500.

— retiarmata, Dendy, 499.

Ceratodus, Agass., 87, note; mentd., 165-
215.

— forsteri t, 179: mentd., 166, 175, 199.

Ceratosoma, Adams § Reeve, 558.

— brevicaudatum, dérah., 528, 558, 569,
572, 577, 579.

— calidonicum, P%sch., 558.

— corallinum, Odhn., 523.

— cornigerum, Adams § Reeve, 558.

— oblongum, Abrah., 558, 560.

— tenue, Abrah., 558.

654 INDEX.

Ceratosoma trilobata, 570.

Ceratozercon bicornis, Halb.*, 375, 392.

Cercaripora, Fisch., 304, 309,

— argillacea, 304,

— hgulata, 304.

— truncata, 304.

Cercobodo, Avass., 467.

Cercomonas crassicauda, Alex., 452, 457,
458, 465.

— sp., 457.

Ceriodaphnia pulchella, Sars, 425; distrib.,
414.

— quadrangula, distrib., 414, 415, 416.

Ceriopora globulus, Reuss, 688.

Chelifer dubius (Camb.), 364.

Chalina palmata, Ridl. § Dendy, 499.

— sp., 478, 500.

Chaperia, Jull., 304.

acanthina, Harmer *, 304.

— australis, Jull., 304.

Chartella, Gray, 304, 311.

— papyracea, 304.

Chaunosia, Busk, 304.

— hirtissima, Busk, 304.

Cheyletus eruditus, Schr., 387.

— venustissimus, C. LZ. Koch, 387.

Chilodon cucullulus, Ehrenb., 457.

Chlamydophrys stercorea, Cienk., 453, 457.

Chlidonia, Lam., 305, 306.

— pyriformis, 305.

Chhidonia, Hiibn., 305.

Chlidonia, Herr.-Schiitt., 305.

Chlidonias, Rafin., 305.

Chlorella Beyer., food of Daphnia, 416.

Chloropisca glabra, 405, 408.

Chlorops tzniopus, Mey., head capsule,
399-410.

Chromodoris, Ald. & Hane., 553.

— magnifica, Quoy & Gaim., 553.

Chrondrilla australiensis, Carter, 478, 496.

Chydorus piger, 425, 437.

— spheericus, 414.

Cibicides refulgens, Montf., 635.
Cidarites annulifera, Lam., 248.
Cilicea stylifera, Whitel., 14.
Ciliceeopsis, Hansen, 13, 14.

— dakini, Tatt.*, 13.

— granulata, 14.

— whiteleggei, 14.

Cilliba, Heyd., 378.

— vegetans, Hull, 377.

Cillibano dinychoides, Huil, 379.

Cinclidium stygium, Stw., 450.

Cinetoscids, Martens, 305.

Cladochalina euplax, Lendent., 499.

Cladomonas sp., 457, 462.

— fruticosa, Stein, 457, 463.

Clark, H. L., Some Echinoderms from
West Australia, 229-251.

Cleanthus, Leach, 536.

Clypeus of Chlorops, 403.

Coition of Lyctus, 590.

Coleoptera, female reprod. organs, 59.

Collinge, W. E., Hluma celatum, 103;
Tsopods fr. Madagascar, 107.

Coloburiscus humeralis (Walk.), wing-
veins, 153.

Colpoda, Schrank, mentd., 450.

—cucullus, Lhrenb., 453, 457.

Colotois pennaria, 86.

Columnaria, Lev., 305.

— borealis, Lev., 305.

Columnaria, Goldf., 305.

Colymbus immer, M. 7. Briinn., courtship
activities, 265.

— stellatus, Pontopp., courtship activities,
253-292.

Comatella nigra, 4. H. Clark, 232; distrib.,
231.

— stelligera, 4d. H. Clark, 282; distrib.,
231.

Comatula jacguinot?, J. Muller, 234.

— pectinata, A. H. Clark, 233; distrib.,
231.

— purpurea, mentd., 233.

— solaris, Lam., 235; distrib., 231.

Comanthus annulata, dA. H. Clark, 233;
distrib., 231.

— luteofusea, mentd., 234.

— parvicirra, A. H. Clark, 254; distrib.,
231.

— polvenemis, 4. H. Clark, distrib., 231.

Conchopoma gadiformis, Kner t+, struc-
ture, 198-201, 213-214.

Coniston plankton, 421.

Conocephalus conicus, Dum., mite on, 384.

Copromonas sp., 457.

— subtilis, Dobell, 465.

Cornucopina, Lev., 306, 327, 359.

— grandis, Lev., 306.

— infundibulata, Lev., 306.

Cornuspira, Schultze, 610

INDEX. 655

Cornuspira involvens, Reuss, 610.

— selseyensis, Heron-All. § Harl., 610.

Corycia coronata var. simplex, Penard, 452,
470.

— flava, Greef, 452, 470.

Corynoporella, Hincks, 306.

— tenuis, Hincks, 306.

Coryphella rufibranchialis, 569, 570,

Corystes, Latr., 31, 57, 58, 61.

Corythion dubium, Taran., 453, 475.

— schaudinni, Schussl., 473.

Coscinasterias calamaria, Perrier, 244;
distrib., 229.

Coscinoderma pyriforme, Lendenf., var. a,
479, 512, 513.

— —, var. B, 479, 513.

Cosmonotus, HW’. Adams, 27, 61.

— Grayi, Adams § White, 26.

Cothurnicella, W. Thoms., 306.

— dedala, W. Thoms , 306.

Courtship in birds, 253-292.

Crab, mouth-parts, 115-142.

Crane, courtship activities, 277.

Craspedozoum, MacGill., 306, 339, 347.

— ligulatum, MacGill., 306, 344.

— membraniporoides, MacGill., 306.

— roborutum, MacGill., 306, 345.

— spicatum, MacGill., 3806, 348, 344,
347.

Craterites, Heron-All. § Earl.*, 611, 612,
646.

— rectus, Heron-All. § Larl. *, 611, 646.

Crella schmidti, Ridl., 505.

Crepis, Judl., 306.

— longipes, Jull., 306.

Crinoidea from West Australia, 252-235,

Crista, auctt., 807.

Crisia, Lame., 306, 507, 308, 521.

— appendiculata, 317.

— boryt, Aud. (?), 316.

— ciliata, Zamz., 306, 307.

— cornuta, 317.

— delilii, Aud., 303.

— eburnea, Luma., 306, 307.

— elegans, Lamx., 342.

Cristellaria, Zam., 625.

— gibba, Orb., 625.

— rotulata, Brady, 625.

Crisularia, Gray, 306.

— fastigiata, Gray, 306.

Crustacea of the tarns, 425; mentd., 285.

Crustacean plankton of English Lakes,
411-447.

Ctenodus, Agassizt, structure, 188-199;
mentd., 163, 171, 195-215.

— cristatus j, 189, 195.

— elegans +, 164.

— ellipticus t, 164.

— imbricatus ft, 164.

— “interruptus Barkas”t, 172, 190-196,
2112.

—murchisoni, Ward +, mentd., 196.

— obliquus +, 164, 190.

— sp.t, mentd., 176.

— tardus, Friisch t, sp. dub., 196.

— tuberculatus +, 166, 195.

Cuckoo, courtship activities, 276.

Cyclidium glaucoma, Ehrenb., 453, 457.

Cyclodorippe, 28, 54.

Cyclops, O. F. Muell., in tarns, 425.

— abyssorum, Sars, 421, 435, 436, 437 ;
distrib., 414, 415, 418.

— agilis, Koch, 425.

— capillatus, Sars, 438, 459.

— crinitus Gract., 437-439.

— diaphanus, Fisch., 488.

— fuscus, Schmerl, 425.

— lacustris, G. O. Sars, 436,
—leuckarti, Claws, 425, 4
414-416. ;

— nanus, G. O. Sars, 425.

-— robustus, G. O. Sars, 439.

— scutifer, G. O. Sars, 436, 487.

— strenuus, Fisch., 486, 437.

—venustus, Norm. § Scott, 425, 437,
459. :

— vernalis, isch., 458, 459.

— — yar., 439.

— vicinus, Ulj., 456.

— viridis, Misch., 425, 439.

Cyclostomes, mentd., 285.

Cylindrecium, auctt., 309.

Cymbzremus monilipes, Halb. *, 381.

Cymbalopera, Hagen, 630.

—millettii, Heron-All. § Earl., 630.

— poeyi, Brady, 630.

Cymodoce, Leach, 14.

— bicarinata, Stebb., 16.

— longistylis, Miers, 16.

— mammifera, Hasw., 14.

— pelsarti, Tatt.*, 15.

— pilosa, M.-Edw., 16.

437.
35; distrib.,

656 ; INDEX.

Cymodoce zanzibarensis, Stebb., 16.

Cymonomops, Ale., 28.

Cymonomus, M.-Edw., 55.

Cyphoderia ampulla var, vitrea, Wailes,
472.

— sp., Schlumb., 453.

Cyrtolelaps mucronatus, Berl., 365.

Dabchick, courtship activities, 253 et seq.

Dameeus, Rajfin., 374.

Dameosoma maculosa, Warb. § Pearce,
380, 392.

— minus, Paoli, var. lamellata, Halb, *,
380, 392.

Damiria australiensis, Dendy, 505.

— schmidta, Tops., 505.

Damsel-flies, mentd., 143.

Daphnia, O. F. Muell., 440.

— cucullata, mentd., 418.

— galeata, 421; distrib., 414.

— hyalina, Leyd., 441-444; distrib., 414-
416, 418.

— — subsp. galeata, Sars, 441-444.

— — subsp. lacustris, Sas, 441, 442.

— —f. lucernensis, Burck., 444.

— lacustris, 444; distrib., 414.

— — var. vicina, Rich., 443.

— longispina, O. F. Mueil., 440-443.

— pellucida, 441.

— pulex, Zatr., 441,

— variabilis, Langh., 441.

Dendoricella schmidti, Hentsch., 505, 479.

Dendrobeania, Lev., 306.

— murrayand, Ley., 306.

Dendrocrinus, Hall, 95.

Dendrodoris, Ehzenb., 560, 561.

— lugubris, Ehrenb., 560, 561.

— mammosa, 523, 562, 578,

— nigra, 523, 561.

Dendrolelaps bicornis, Hull, 366.

— cornutus, Halb. *, 366.

Dendronotus, Ald. & Hance., 544.

Dendrophrya, 8. Wright, 613.

— radiata, S. Wright, 618.

Dendy, A., & L. M. Frederick, Sponges
from Abrolhos, 477.

Dercitopsis ceylonica, Dendy, 489.

—mammillaris, Dendy, 479, 489.

— minor, Dendy, 479, 489.

Desmacella tubulata, Dendy, 503.

Diacara, Budde-Lund, 108.

Diachoris, Busk, 306.

— crotali, Busk, 306.

— hirtissima, Heller, 304.
Diachoseris, Ortm., 306.

— discodernie, Ortm., 306.
— hexaceras, Ortm., 306.
— magellanica, Ortm., 306.

Diadema savianyi, Michelin, 248.
Diaphanosoma brachyurum, Scowrf., 425,
426 ; distrib., 414, 415, 418, 423.

— gracilis, 426.

Diaptomus gracilis, G. 0. Sars, 421-425;
distrib., 414.

— hireus, Brady, 435.

— laciniatus, Zilijeb., mentd., 419.

— laticeps, G. O. Sars, 422, 424, 425, 435;
distrib., 414, 415, 418, 419.

— wierzejskii, Rich., mentd., 419.

Dicranodromia, M.-Edw., 51.

Didymia, Busk, 295, 306, 307.

— simplex, Busk, 306. i

Didymia, Le Pell. § Serv., 306.

Didymozoum, Harmer, nom. noy.*, 295,
306, 307, 328, 325, 359.

— simplex, Harmer *, 306.

Difflugia constricta, Lhrenb., 452, 457, 470.

— fallax, Penard, 452, 470.

— globulus, Duj., 452, 457, 470.

— lucida, Penard, 452, 470.

— pyriformis, Perty, 452, 470.

Ditfusilina, Heron-All. § Barl.*, 614, 646.

— humilis, Heron-All. § Earl, *, 614, 646.

Dileptus, Dy., mentd., 450.

Dimastigameeba, Blochm., 467.

— gruberi, Schard., 457, 469.

Dimetopia, DU. [=Trachymene, Rudge],
299.

Dimetopia, Busk, 299, 307.

— cornuta, Busk, 307.

— spicata, Busk, 307.

Dimorphism in Ptychodera, 393.

Dimorphozoum, Lev., 307.

— nobile, Lev., 307.

Dinychus carinatus, Berl., 377.

Dinycura, Berl., subgen., 378.

Diplodidymia, Reuss +, 307, 315.

— alata, Reuss f, 316.

— complicata, Reuss, 807.

Dipnoi, see Paleeozoic Dipnoi.

Dipterus, Sedy. § Murch. t, mentd., 163-
179, 190, 194, 200, 204, 215.

INDEX. 657

Dipterus platycephalus, Ag. +, structure,
207-212, 215; mentd., 197, 199,
205.

— vyalenciennesi, Sedg. § Murch. +, 205-
207; mentd., 211-212.

Discorbina, Park. § Jones, 630.

— araucana, Brady, 631.

— baceata, Heron- All. § Earl., 631.

— concinna, Brady, 630.

— cora, Heron- All. § Earl. *, 630, 631.

— elobularis, Brady, 631.

— harmeri, Heron-All. § Earl., 635.

— isabelleana, Brady, 630.

— lauriei, Heron-All. § Harl.*, 635, 654,
647.

— mediterranensis, JHeron-All. § Eari.,
631.

— nitida, Heron-All. § Earl., 630.

— patelliformis, Brady, 633.

— polystomelloides, Park. § Jones, 682.

— pulvinata, Brady, 633, 646.

— pyramidalis, Heron-All. § Earl. *, 654,
647.

— varescens. Brady, 632.

—reniformis, Heron-All. § Earl., 632.

— rimosa, Park. § Jones, 632.

— rosacea, Brady, 631.

— rugosa, Brady, 632.

— tabernacularis, Brady, 633, 646.

— — yar. levis, Lieb. § Schub., 634.

— tabernacularis, Sideb., 635.

— turbo, Brady, 631.

— vyalyulata, Brady, 631, 682.

— yilardeboana, Heron-All. § Emil. *, 631.

Distiproboscis of Chlorops, 407.

Dittosaria, Busk +, 307.

— wetherellii, Busk +, 307.

Diver, Red-throated, courtship activities,
253-292; Black-throated, 265 et seq.

Dolabritera, Gray, 534.

— dolubrifera, Rang, 584.

—- pelsartensis, O’Don.*, 523, 534, 530,
577, 578.

Donatia arabica, Tops., 495.

— ylobostellata, Tops., 495.

— ingalli, Dendy, 495.

— japonica var. albanensis, Hentsch., 495.

— multistella, Dendy *, 478, 495.

— robusta, Dendy *, 478, 495.

Doridicola, Leydig, 569.

Doridopsis, Ald. & Hane., 561.

Doridopsis gemmacea, Ald, & Hane., 561.

— mammosa, auct., 562.

— nigra, auct., 561.

Doriopsis, Pease, 560, 561.

— granulosa, Pease, 560.

— mammosa, auct., 562.

— nigra, auct., 561.

Dorippe lunata, M.-Hdw., 47, 48.

Doriprismatica, Orb., 553, 554.

— atromarginata, Cuv., 555.

Doris lugubris, Gravenh., 561.

— mammosa, Abrah., 562.

— mgra, Stimp., 561.

— palhda, Rupp. & Leuck., 553.

Doto coronata, 569.

Dove, courtship activities, 275, 285.

Dragonflies, mentd., 143.

Drammenia, Thor, 387.

— crassipalpis, Thor, 387, 392.

— elongata, Thor, 387, 388.

Dromia, Fabs., 31, 33, 36, 87, 44, 51.

— vulgaris, Bouvier, 31, 35, 40, 42, 50, 77,
78.

Drosophila, Fal/., 88; courtship activities,
270.

Ducks, courtship activities, 290.

Duvaucelia, Risso, 543.

Dynamena, Lev., 307.

— barbata, Lev., 307.

Kehinocardium cordatum, Gray, 251;
distrib., 229-230.

Echinoderms from West Australia (Clark),
229-251; mentd., 285.

Echinodictyum bilamellatum, Ridl., 479,
504,

— elegans, Hallm., 504,

Echinoidea from West Australia, 248-251.

Kchinometra mathaei, Blainv., 250;
distrib., 232.

— setosa, Leske, 248.

Echinonema vasiplicata, Carter, 504.

Hehinospira, Avohn, 92, 93.

Echinus, Linn., 96.

— cordutus, Pennant, 251.

— erythrogrammus, Valenciennes, 250.

— gratilla, Linn., 249.

— mathaet, Blainy., 250.

— pallidus, Lam., 249.

Keret, Snowy, courtship activities, 265 et
seq.

658 INDEX.

Ehrenbergina, Reuss, 628.

— serrata, Reuss, 623.

Elasmopus, A. Costa, 8.

— pectinicrus, 8. Bate, 9, 10.

— serrula, Walker, 9, 10.

— subearinatus, Chilt., 2, 9, 10.

Eluma, Budde-Lund, 103, 104.

— celatum, Collinge, 103-106.

— helleri, Verhoef, 106.

— purpurascens, Budde-Lund, 103, 104.

Emma, Gray, 307, 318, 3825, 327, 329,
356.

— buskii, Harmer *, 357, 359.

—cervicornis, MacGill., 357.

— crystallina, Gray, 307, 357.

— cyathus, Harmer *, 357, 359.

— tricellata, Busk, 357.

Enchelys farcimea, Ehrend., 457.

— sp., 453, 457.

Encionema australiensis, Sollas, 492.

Eneyrtid, mentd., 225.

English Lakes, Crustacean plankton, 411—
447.

Ennerdale Plankton, 420, 421.

Entosiphon sulcatum, Stein, 452.

Entosolenia lineata, Williams, 623.

Ephemera, wing-veins, 152.

Epicranial suture of Chlorops, 899-403.

Epicrius lelaptoides, Berl., 370.

Epipharynx of Chlorops, 407, 408.

Episeius, Hull, subgen., 371, 372, 373.

— italicus, 384.

Epistomia, Flem., 307, 511, 314.

— bursaria, Flem., 307.

Eraunis escularia, Schiff., 86.

Ergasilus, Nordm., 569.

Erina, Swainson, 307.-

Erina, Canu, 307.

— patagonica, Canu, 307.

Erisocrinus, Meek, 95.

Erylus proximus, Dendy, 478, 494.

Eschara fistulosa, L., 308, 308.

— foliacen, Linn., 310.

— papyrea, Pall., 301.

Esperella plumosa, 479, 503.

Esperia parasitica, Carter, 502.

Hucratea, Lam., 307, 310, 311, 314.

— appendiculata, Lamx., 317.

— cordiert, Aud., 305.

— cornuta, Lamx., 307.

—- lafontui, Aud., 298.

Eucratea loricata, Zamz., 307, 312, 317.
— vesiculosa, Hamm., 305.

Eucratia, Flem., 308 = preec.
Eugamasus loricatus, Wank., 364.
Euglena, Zhrenb., in Spitsbergen, 450.
Euglypha ciliata, Hhrenb., 453, 471.

— levis, Perty, 458, 457, 472.

— strigosa, Leidy, 453, 457, 471.

— — f. g abra, 457.

— — yar. glabra, Wailes, 4
— tuberculata, Dy., 453, 4
Euoplozoum, Harmer *, 295, 308, 359.

— cirratum, Harmer *, 308, 326.
Eupagurus, Brandt, 31.

Kuphrurus, Rajin., 564.

HEuryale aspera, Lam., 246; distrib., 230.
Eurystheus, Bate, 10.

— afer, Stebb., 11.

— atlanticus, Stebb., 2, 10, 11, 18.

— gardineri, Walker, 11.

— lamellatus, Baird, 425.

— zeylanicus, Walker, 10, 11.
Eusthenopteron +, mentd., 215.

Euthyris bombycina, Busk, 308.

— episcopalis, 308.

Euthyris, Hincks, gen. dub., 308, 516.

— obtecta, Hinchks, 308.

Euthyroides, Harmer, 308.

— episcopalis, Harmer, 308.

Evolution of Courtship in birds, 253-292.
Hyes of Chlorops, 405.

58, 471.
72.

Falearia, Haworth, 308.

Falearia, Oken, 308.

— anguina, 308.

— cornuta, 308.

— eburnea, 308.

— falcata, Oken, 308.

Fareimia, Wlem., 308.

— fistulosa, Flem., 308.

— oculata, Waters, 308.

— sinuosa, Hass., 302, 3038.

— salicornia, Hass., 302.

— spathulosa, Hass., 303.

Farciminaria, Busk, 309.

— aculeata, Bush, 309.

— atlantica, 305.

Female genitals of lLyctus, 585-590;
reproductive organs in Coleoptera, 591.

Filicella, S. Wood }, 509.

INDEX. 659

Filicella anguinea, S. Wood (?), 309.

Fischerina, Zerg., 610.

— pellucida, Millett, 610.

Fishes, méntd., 285.

Fissurina bicarinata, Terq., 624.

— levigata, Reuss, 624.

Fistulana, O. F. Muell., 309.

— longicornis, O. F. Muell., 309.

— multicornis, O. F. Muell., 309.

— muscoides, O. F. Muell., 309.

— ramosa, O. F. Muell., 309.

— sunplex, O. F. Muell., 309.

Fistularia, Linn., 809.

Flabellaria, Gray, 309.

— setacea, Flem., 309.

— spiralis, Gray, 309.

Flabellaria, Zam., 309.

Flabellaris (pars), Waters, 310, 339,

— flabellum, Waters, 310, 340.

—- roborata (pars), Waters, 310, 845.

— — var. higulata, Waters, 344.

Flabelltina, Ley., 310, 339.

— roborata, Ley., 310, 345,

Flabellina, Voigt, 310.

Flagellates, food of Daphnia, 416.

Floutern Tarn, crustacea, 423.

Flustra, Zinn., 310, 311.

— abyssicola, G. O. Sars, 315.

— acanthina, Quoy & Gaim., 304.

— biseriata, Busk, 312.

— bombycina, Kill. & Sol., 308, 314.

— carbasea, Ell. & Sol., 301.

— foliacea, Linn., 310.

— membraniporoides, Busk, 306, 345.

— militaris, Waters, 317.

— murrayana, Bean, 306.

— nobilis, Hineks, 307.

— papyracea, Wl. & Sol., 804.

— reticulum, Hincks, 315,

— setacea, Flem., 309.

Foraminifera of Lord Howe Island, 599-
646.

Frew, J. G. H., Morphology of head of
Chlorops, 399-410.

Fromia andamanensis,
distrib., 232.

— elegans, H. L. Clark, 240; distrib., 232.

— milleporella, Gray, mentd., 240.

Frondicularia, Defr., 625.

— scottii, Heron- All. § Earl., 625, 646.

Fronto-clypeus of Chlorops, 403.

Koehler, 289;

Galathea, Fabr., 31.

Galvina viridula, 570.

Gamasellus granulatus, Halb,*, 364, 391.

— mucronatus, Halb. *, 365.

— subnudus (?), Berl., 365, 391.

Gamasus bicornis, Kram., 375.

— crassitarsis, Halb. *, 363, 391.

— halleri, G. & R. Can., 369.

— magnus, Avam., 364.

— mucronatus, G. & R. Can., 365.

— oudemansi, Berl., 364.

— remiger, Kram., 574.

Gammaropsis atlantica, Stebb., 10.

— gardinert, Walker, 10.

— zeylanicus, Walker, 10.

Gammarus tenellus, Dana, 8.

Gannet, courtship activities, 278.

Garstang, W., Recapitulation, 81-101.

Gaudryina, Ord., 620.

— rugosa, Orb., 620.

— scabra, Brady, 620.

Gemellaria, Sav., 314.

Gemellaria, van Bened., 310-312.

— avicularis, Piep., 317.

— loriculata, Blainy., 310.

Gemicellaria, Blainy., 310.

— bursaria, Blainy., 311.

— loriculata, Blainy., 310, 311.

Genital system of Lyctus brunneus, 581.

Gilchrist, J. D. F., Dimorphism in Ptycho-
dera capensis, 393-398,

Gill, E. L., see Watson, D. M.S.

Gleditschia triacanthos, Zinn., mentd.,

Mio
Glenodinium, Ehrenb., in Spitsbergen,
450.

Globigerina, Orb., 627,

— bulloides, Orb., 627.

— cretacea var. eggeri, Heron-All. §: Earl.,
627.

— rubra, Ord., 628.

— triloba, Reuss, 627.

Glossze of Chlorops, 408.

Glossodoris, Ehrenb., 553, 554.

— elisabethina, 570.

— pallida, 553.

— westraliensis, O’Don., 554, 577, 579.

— wantholeuca, Ehrenb., 555.

Goats Water crustacea, 424.

Golderest, courtship activities, 277.

Goniobranchus, Pease, 553.

660 INDEX.

Goniobranchus vibrata, Pease, 553.

Goniodoris, Forbes, 553, 564.

Gonodontis bidentata, 86.

Gonostomum atline, Stein, 457.

Grantiopsis cylindrica, Dendy, 477, 518.

— — var. fruticosa, Dendy *, 477.

— infrequens, Dendy & Row, 483.

Graphiocrinus, Hon., 95,

Graptoleberis testudinaria, 425.

Grebe, courtship activities, 2538 ef seq.

Grimmia commutata, Hweb., 451.

Grubia, Czern., 12, 13.

— brevidactyla, Chev., 13.

— setosa, Stebb., 2, 12.

Guillemots, courtship activities, 267, 281.

Gulls, courtship activities, 281.

Gumey, R., Crustacea of English Lakes,
411-447.

Gymnopleura, Bourne *, 55.

Gypsina, Carter, 637, 639.

—- globulus, Brady, 658.

— inherens, Brady, 637, 647.

— rubra, Heron-All. § Earl., 638.

— vesicularis, Brady, 638.

Gyrinus, Linn., 67.

Haddonia, Chapm., 615, 616.

— torresiensis, Chapm., 615, 646.

Halbert, J. N., Acari, 363-392.

Halichondria phakellioides, Dendy *, 478,
498, 519.

Halophila, Gray, 311.

— johnstone, Gray, 311.

— johnstonie, Gray, 311.

Haplocrinus, 97, ftnote.

Haplophragmium, Reuss, 613.

— anceps, Brady, 614.

— canariense, Brady, 613.

— compressum, Goés, 613.

— emaciatum, Brady, 615.

Harmer, Sir S. F., Cellularine Polyzoa,
293-361.

Hartmanella hyalina, Alex., 452, 457, 469.

— sp. &, Cutl., 452.

Tauerina, Ord., 609.

— compressa, Orb., 609.

— ornatissima, Brady, 609.

Ifead capsule of Chlorops, 399-410.

Heliocidaris erythrogramma, Agassiz §
Desor, 250; distrib., 229, 282.

Heliocidaris tuberculata, mentd,, 250.

Helkesimastix firecicola, Woode. § Lep., 452,
457, 461.

Hemichordata, 393.

Hemipeneus spinidorsalis, 45.

Heptagenia, mentd., 161.

Hermannia arrecta, Nicol., 381.

— granulata, Nicol., 381.

Hermanniella granulata, Halb.*, 381.

Heron-Allen, E., & A. Earland, Foramini-
fera of Lord Howe Island, 599-646.

Herons, courtship activities, 265 et seq.

Herpetomonas, Kent, 460.

Heterocella, Canw7, 311.

— fragilis, Canu }, 311.

Heteroflustra, Ley., 311.

Heteromita, 458.

— globosa, Stein, 450, 451, 457, 458.

— lens, Mwell., 452, 457, 458.

Heterostegina, Orb., 641.

— depressa, Orb., 641.

Hexabranchus, Lhrenb., 545.

—imperialis, ent, 523, 545, 546, 577,
578.

— pretextus, Lhrend., 545.

Hiantopora, Mae Gill., 311.

— ferox, MacGiill., 311.

Hickson, S. J., Two Sea-pens from
Abrolhos, 21-28.

Highlow Tarn, crustacea, 424.

Himantozoum, Harmer *, 295, 311, 326.

— leontodon, Harmer *, 311.

— margaritifera, Harmer *, 311.

— mirabile, Harmer,*, 311.

— sinuosa, Harmer *, 311, 312.

Hippa, Fadr., 31, 56.

Hippospongia intestinalis, Dendy*, 479,
511,

Hircinia sp. ?, 502.

Holoeladina pustulifera, Carter, 638.

Holopedium gibberum, Zadd., 412, 414—
416, 418, 421, 422, 440.

Holopsamma crassa, Carter, 511.

Holopus, Orb., 97.

Holostapis vagabundus, Berl., 366.

Homarus, Edw., 39, 48.

Homolodromia, M.-Edw., 51.

Hoplitella, Zev., 312.

— armata, Lev., 312.

Humming-birds, courtship activities, 277,
278.

INDEX.

Huxley, J. S., Courtship activities in the
the Red-throated Diver (Colymbus
stellatus, Pontopp.), 255-292.

Huxleya, Dyster, 312.

— fragilis, Dyster, 312.

Hyalosphenia minuta, Cash, 457, 471.

Hybernia, Zaty., 88.

— aurantiaria, 88.

— defoliaria, 88.

— marginaria, 88,

Hybocrinus, Bill., 94.

Hydracarina, 387, 389.

Hydryphantes, 388.

— bayeri, Pisar., 389, 392.

— bayeri [var.] nonundulata, Jets, 389,
392.

— crassipalpis, Koen., 389.

— — f. lacustris, Halb.*, 389, 392,

— dispar, Halb.*, 389, 392,

— placationis, Thon, 389, 392.

— planus, Thon, 389.

— prolongatus, Thon, 388, 389, 392.

— ruber, De Geer, 388, 389, 392.

Hyocrinus, Sedgzw., 97.

Hypnum, ZLinn., 5 spp., 451.

Hypoaspis aculeifer, Can., 367.

— subglabra, Oudem., 375.

Hypograntia infrequens, Carter, 485, 486.

Hypopharynx of Chlorops, 408.

Idalia aspersa, 569.

Tlia nucleus, 54.

Insects, Primitive, mentd., 285.
Internal avicularia, 318.

Iphis halleri, Berl., 369.

Tridia, Heron-AU. § Earl., 612.
— diaphana, Heron-All. § Earl. 612.
Ismalia monstrosa, 571.

Tsodictya permollis, Bowerb., 497.
Tsoétes, Zinmn., in tarn, 423.
Tsopoda from Abrolhos, 1.

i

Tsopods, terrestrial, 103, 107.

Jubella, Jull., 312.

— enucleata, Jull., 312.

Kalykenteron elegans, Lendent., 504.
— silev, Lendenf., 504.
Kenella, Zev., 311, 312.

LINN. JOURN.—ZOOLOGY, VOL. XXXY.

661

Kenella biseriata, Zev., 314.

Kentrodoris maculosa, 523.

Kestrel, courtship activities, 285, 289.

Khaya, 4. Juss., sp., mentd., 217, 219.

Kinetoskias, Kor. § Dan., 305, 312, 359,

— arabianensis, Roberts, 308.

—- arborescens, Dan., 312.

— cyathus, 326.

— snuthi, Dan., 312.

— smittii, Dan. (emend.), 312.

Kingfisher, its brilliant colouring
matic, 288.

apose-

Labelle of Chlorops, 408.

Labidostoma cornuta, 379.

— luteum, Aram., 379.

Labium of Chlorops, 406.

Labrum of Chlorops, 404.

Leelaps aculeifer, Can., 367.

—agilis, C, L. Koch, 367.

— fimbriatus, Halb.*, 368, 391.

— halleri, G. Can., 369.

— latisternus, Halb.*, 367, 391.

— simplex, Halb.*, 368, 591.

— stabulasis, C. LZ. Koch, 367.

Lafoea dafoyt, Blainy., 317.

— cornuta, Lamz., 317.

Lagena, Walk. § Boys, 628.

—annectans, Burr. § Holl., 624.

— bicarinata, Ihillett, 624.

— clathrata, Brady, 624.

— levigata, Brady, 624.

levis, Welliams., 624.

— lineata, Brady, 623.

— marginata, Brady, 624,

— squamosa, Cushm., 624,

Lakes, English, crustacean plankton, 411-
419.

Lamellaria, Oken, 92, 93.

Lamprometra gyges, 4d. H. Clark, 254;
distrib., 231. ;

Laplysia, Linn., 527, 528.

— depilans, Linn., 528.

Lasioseius, 363.

— glaber, Berl, var. curtipes, Halb. *, 370,
391.

— — var. minor, Trdédg., 370.

— gracilis, Halb.*, 369, 391.

—italicus, Berl., 371, 3872,
392.

374,

Ora
V0;

AT

for)
for)
bo

Lasioseius lelaptoides, Halb.*, 370, 371.

— major, Zalb.*, 371-873, 392.

— michaeli, Halb.*, 371-378, 392.

— minusculus, Berl., 874.

— minutus, Berl., 374.

— muricatus, C. Z. Koch, 370, 871.

— ometes, Oudem., 370, 391, 392.

— sphagni, Halb.*, 371, 391.

— subglaber, Halb.*, 373, 374, 392.

— tenuipes, Halb.*, 371-373.

Lasius flavus, Mayr, 379.

— mixtus, Mayr, 367.

— niger, Fubr., 379.

Law, biogenetic, 81.

Leiosoma palmicinctum, Mich., 379. -

Lelapia, 477.

—antiqua, Dendy *, 487, 519;
478, 479,

— australis, Carter, 488, 489.

— nipponica, 479, 488, 489.

Lenticulites complanata, Detr., 641.

— rotulata, Lam., 625.

Lepralia ferox, MacGill., 311.

Leptodora kindtii, Zidéjeb., 421; distrib.,
414, 415, 417, 419.

Leptomonas jaculans, Ber/., 460.

Leuealtis clathria, Haeck., 477, 483.

Leucandia microrhaphis, Dendy, 482.

Leucetta chagosensis, Dendy, 477, 482.

— microraphis, Haeck., 477, 482.

— primgenia var. microraphis,
482.

Leuconia dura, Poléj., 482.

Leucosia, Fabr., 26.

Leucosolenia grisea, Dendy *, 480, 477,
518.

— protogenes, Haeck., 477, 480.

Leucothoé, Leach, 6.

— spinicarpa, Chilt., 6.

Levers Water, crustacea, 423.

Liacarus palmicinctum, Mich., 379.

Limnicythere incisa in Africa, 420.

— mirabilis in Switzerland, 420.

Limnocalanus grimaldii,
435.

— macrurus, G. O. Sars, 421, 426-436,
447 ; distrib., 414, 415, 418, 420.

Liriope, Gistel, 542.

Lithodes, Zaty., 28, 56.

Tittorella, Berg., abundant in
423,

mentd.,

Haeck.,

Guerne, 427-

tarns,

INDEX.

Lituola cenomana, Carp., 614.

Tntuolina irregularis var. compressa, Goés,
613.

Ljania bipapillata, S. Thor., 388.

Lobelia, Zinn., in tarns, 423.

Locust tree, mentd., 219.

Lomanotus genei, 570.

Lord Wowe Island Foraminifera, 599-
646.

Loricaria, Zinn., mentd., 310, 312.

Loricaria, Lamx., 312.

-— egyptiaca, Aud., 310, 317.

— americana, Lamx., 312.

— europea, Lamx., 312.

Loricula, Curtis, 312.

Loricula, Templet., 312.

— loricata, Templet., 312.

Low Water crustacea, 423.

Luidia maculata australie, Déderlein, 235 ;
distrib., 229-230.

Lyctus brunneus, Steph., method of ovi-
position and the egg of, 217-227;
genital syst., 581-597.

— canaliculatus, Fabr., 217, 218, 581.

— carbonarius, Waltl., 218.

— cavicollis, Zec., mentd., 224,

— linearis, Goéze, 224, 590; mentd., 217—
222.

— oblongus, Oliv., 217.

— parallelopipedus, Welsh., mentd.; 224.

—planicollis, Ze Conte, mevtd., 218-
225.

— pubescens, Panz., mentd., 218.

— sp., mentd., 218.

— striatus, Melsh., 217.

— unipunctatus, Herbst, 217.

Lyprobius cristatus, Budde-Lund, 107.

Lyre-bird, courtship activities, 277.

Lyveidus, 26, 31-79.

— channeri, Wood-Mason, 26, 69.

-— tridentatus, de Huan, 25, 29, 30, 69,

77—79.

Macrocheles, Berl., subgen., 366.

— yagabundus, Halb.*, 566.

Madagascar, ‘wo Isopods from, 107.

Madrella ferruginosa, Ald. § Hance.,
525.

Meera, Leach, 8.

— mastersii, Stebb., 9.

INDEX. 663

Meera tenella, Walker, 2, 6.

— spinosa, Hasw., 8.

Mahogany, mentd., 217, 219.

Male genitals of Lyctus, 583-585,

Mandibles absent from Chlorops, 406.

Maplestonia, MacGill., 812.

— cirrata, MacGill., 312.

Matuta, Hartm., 58, 67.

Maxillary palps of Chlorops, 409.

Mayflies, wing venation in, 143-162.

Mediproboscis of Chlorops, 407.

Megalichthys, Agass.t, mentd., 211-212.

Megalopastes arenifibrosa, Dendy *, 509,
479.

Megamera mastersti, Hasw., 9.

— subcarinata, Hasw., 9.

— thompsoni, Miers, 9.

Megapus gibberipalpis, Piers., 388.

Melicerta, Schrank, 317.

Melicerita, W.-Edw., 313.

— charlesworthit, Orb., 313.

—- charlesworthii, 8. Wood, 313.

— dubia, Busk, 313.

Melhicertina, Ehrenb., 313, 317.

Melosira, Spreng., food of Daphnia, 416.

Membranicellaria, Zev., 313.

— dubia, Lev., 313.

Membranipora magnilabris, Busk, 304.

— roborata, Hincks, 306, 310, 345, 347.

— spinosi, Quoy & Gaim., 304.

Mentpea, Lamx. (pars), 348, 353, 356.

Menipea, Lame., 306, 318, 315, 338, 3:
328-330; Craspedozoum group, 339.

— aculeata, Busk, 355.

— apicata, Harmer *, 343-346.

— benemunita, Busk, 300, 331, 384.

— buskii, W. Thoms., 357.

— buskii, Waters, 313.

— cervicornis, MacGill., 357.

— cirrata, Lame., 313.

— errata, Busk, 339, 340.

— cirrhata, Lamx., 313.

— clausa, Busk, 351.

— compacta, Hincks, 354.

— crispa, Marcus, 313, 340, 359.

— erystallina, MacGill., 357.

— cyathus, W. Thoms., 357.

— duplex, Ley., 352.

— fasciculata, MacGill., 335.

— flabellum, Lame, 313, 340.

Menipea flabellum, Busk, 510, 340.
— flagellifera, Busk, 345.

— fuegensis, Busk, 356.

— funiculata, 331.

— gracilis, Busk, 354.

— gracilis, Norm., 304.

— jeffreysi, Norm., 351, 352.

— ligulata, Zev., 319, 344-346, 360.
— lonyispinosa, Yan. & Ok., 556.

— murginata, Hincks, 332.

— maroniensis, Bush, 318, 341, 842, 347,
360, 361.

— maroniensis (pars), Busk, 3835, 386
341, .

— multiseriata, Bush, 347, 348.

— occidentalis, Trask, 354.

— ornata, 340.

— patagonica, Busk, 521, 341.

— pateriformis, Busk, 339.

— pribilofi, Robertson, 354.

— quadrata, 542.

— roborata, Zev., 318, 319, 343-346, 348,
360.

— smitti, Norm., 352.

—- spicata, 319, 328, 359, 360.

— sympodia, Yan. & Ok., 355.

— ternata, Hincks, 354.

— tricellata, MacGill., 357.

— triseriata, Busk, 319, 341, 342, 347, 359,
361. ;

— yectifera, Harmer *, 255, 319, 339, 346,
348, 360.

Mesephemera cellulosa (Hagen)}, wing-
yeins, 151-152.

Metazoa, 83.

Miliolina, Williamson, 604.

— auberiana, Brady, 607.

— — yar. semireticulata, Heron-All. &
Harl*, 607, 646.

— bertheliniana, Brady, 605.

— hicostata, Gioés, 607.

— bosciana, Millett, 606.

— circularis, Brady, 604.

— contorta, Goés, 608.

— cultrata, 605.

— envieriana, Brady, 607.

— dilatata, Heron-All. § Earl., 604,

— ferussacii, Brady, 608.

— fichteliana, Brady, 405.

— fusea, Heron-All. § Earl., 608.

— kerimbatica, Heron-All. § Earl.*, 608.

664

Miliolina labiosa, Brady, 604.
leevigata, Heron—All, § Harl., 606.
—- linnzeana, Brady, 609.
— oblonga, Heron-All. 5 Earl. 606,
— parkeri, Brady, 608.
— reticulata, Brady, 608.
— rotunda, .Willet, G06.
— schauinsland, Rhumbl., 601.
—seminuda, Heron-Au. § Larl., 605.
— seminulum, Brady, 606.
— stelligera, Heron-All. § Earl., 609.
— striata, Heron-All. § Earl., 609.
— suborbicularis, Millet, G05.
— subrotunda, Grady, 605.
—triangularis, Jones, Parker S Brady,
607.
— tricarinata, Brady, 6065.
— trigonula, Brady, 605.
— undosa, Brady, 607.
— undulata, Heron-All. § Larl.*, 607.
—yalvularis, Brady, 604.
— vulearis, Heron-All. § Earl., 606.
— webbiana, Heron-All. § Bayt, G05.
Mitiolites planulata, Lam., 603.
— trigonula, Lam., 606.
Milivloculina undulata, Ueron-All, & Karl,
608.
Millepora miniacea, Pall., 659
Millericrinus, Orb., 95.
Molege cristatus, 424.
Mononota, Piep., 313, 317.
Monsella, Canwt, 315.
— eocena, Canut, 315.
Moorhen, courtship activities, 285.
Moraria brevipes, G. O. oe 437,
Morgan, Miss A., her theory of wing-
veins in Dragonflies criticised, 143,
161.
Morpholovy of head of Chlorops, 399.
Mortensenia, Déderlein, mentd., 250
— oblonga, Déderlein, 250. -
Mouth-parts of Chlorops, 899-410; of
Crab, 115-142.
Mus sylvaticus, Zen. | Sphalm. sylvestris),
367.
Mycetobia, Merg.. 403.
Mycetophila, Wezg., 403.
Myonyssus decuminus, Tiradb., 367.
Myviophyllum, Zenz., in tarn, 425.
Mysis oculata, 4838.
— relicta, 420, 433.

INDEX.

Naegleria gruberi, Alex., 469.

Nagara cristata, Budde-Lund, 107,
Nannonyx kidderi, Chilt., 3.

Naresia, WV. Thems., ow

— cyathus, W. Thoms., 3

Nassula, Lhrenb., 450.

Nautilus beccarit, Linn., 637.

— crispus, Linn., 640.

— depressulus, Walk. & Jacob, 639.

— lvbatulus, Walk. & Jacob, 635.

— macellus, Ficht. & Moll, 640.

— melo, Ficht. & Moll, 612.

— pertusus, Forsk., 610.

— repandus, Ficht. & Moll, 686,

— striatus-punctatus, Ficht. & Moll, 640.
— umbilicatulus, Mont., 640.

Nebela collaris, Letdy, 453, 454, 471.

— lageniformis, Penard, 457, 471.
Necromantes, Gistel, 542.

Nectria macrobrachia, H. L. Clark*, 23

distrib., 280, 232. t
— ocellata, Perrier, mentd., 257.

— sella, Gray, mentd., 237.

Nellia, Bush, 314.

— oculata, Busk, 308, 314.

— simplex, Busk, 314.

— tenella, 314.

Nembrotha, Bergh, 567, 568, 569.

— nigerrima, Bergh, 567.

— purpureolineata, O’Don., 525, 568, 578,
579.

Neoceratodus forstertt, 179.

Neogenetic and neomorphic, use defined, 99.

Neophyllobius elegans, Berl., 384.

— saxatilis, Halb.*, 384, 392.

Nephrops, Leach, 32, 45, 46.

— norvegicus, 39, 41, 45, 78

Nightingale, courtship activities, 263.

Nolella, Gosse, 309.

Nomenclature of Polyzoa, 294, 297-517

Nonionina, Ovb., 639.

— canariensis, Orb., 613.

— depressula, Brady, 659.

— pauperata, Balk. § Wright, 640.

— umbilicatula, Brady, 640.

Notamia, Busk, 507.

Notamia, Rafin., mentd., 514.

Notamia, Flem., 514.

— bursaria, Flem., 314.

— loricata, Flem., 314.

Notaspis lacustris, Michael, 384,

INDEX.

Notaspis monilipes, Mich., 381.

Notodoris, Bergh, 564, 566.

— citrina, Bergh, 564, 567. ,

— gardineri, hot, 523, 565, 567,578, 579.

— minor, Eliot, 567.

Notoplites, Harmer *, 295, :
329, 530, 339, 343, 347, ae

— minlionetoms, Harmer *, 352.

— avicularie, Harmer *, 351.

— biloba, Harmer *, 350.

— crateriformis, Harmer *

— drygalskii, Harmer *

— elongatus, Harmer *, 351, 353.

— jeffreysii, Harmer *, 351.

— marsupiatus, Harmer *, 351, 35

—normani, Harmer *, 353.

— perditus (?), Harmer *, ¢

— rostratus, Harmer *,
359.

— simplex (?), Harmer *, 35:

— smittii, Zarmer*, 352.

— tenuis, Harmer *, 352.

— yanhoffeni, Harmer *, 353.

— watersi, Harmer *, 355.

Notopoides, 58, 66, 67, 68.

— latus, Henders., 26, 60.

Notopus, Haan, 31, 52, 41-79.

— Beyrichii t, 52.

— dorsipes, Labr., 25, 31, 48, 60-79.

— ovalis, Henders., 26.

Notosceles, Bowne '
70; genus deser., 73.

— Onan Bowne*, 25, 31, GO, 74,
No:

Nubecularia, Def, 601.

— bradyi, Aillett, 601.

— inflata, Brady, 601.

— lucifuga, Defr., 601, 646.

— schauinslandi, Heron-Adll. § Larl.*, 601,
646; mentd., 599, 600, 602.

Nuclearia simplex, Cienk., 457, 473.

Nymphea, Zinn., in Highlow Tarn, 424.

2h

Oak, “ Wainscot,” mentd., 219.

Occiput of Chlorops, 405.

Ocelli of Chlorops, 405.

Odonata, mentd., 144.

O'Donoghue, C. H., Opisthobranchiata
from Abrolhos, 521

Oicomonas termo, Kent, 452, 457.

665

Oligometra carpenteri, Bell, mentd., 235.

— serripinna, A. H. Clark, 235; distrib.,
231.

Onchidiopsis, Bergh, 92.

Onchoporella, Busk, 308, 314, 316.

— bombycina, Busk, 314.

— buskii, Harmer *, 295, 514.

— diaphana, Busk, 314.

— ligulata, Busk, 314.

Oniscigaster distans, Zatun, wing-veins,

155, 158, 161.

Ontogenies, phyletic succession, 83.

Operculina, Ord., G41.

— complanata, Orb., G4.

Ophiactis savignyi, Ljungman, 246 ;
distrib., 229, 232.

Ophiocoma brevipes, Peters, distrib., 282 ;
mentd., 247.

— — var. variesata, H. L. Clark, 247.

— dentata, mentd., 247

— déderleini, mentd., 247.

— insularia, mentd., 247.

— variegata, H. A. Smith, 247.

Ophiolepis imbricata, Muller & Troschel,
248,

— savignyt, Miller & Troschel, 246.

Ophionereis porrecta, Lyman, 247 ; distrib.,
232)

Ophioplocus imbricatus, Lyman, 248 ;
distrib., 232. :

Ophiothrix longipeda (Lam.), distrib., 232.

— michaelseni, Aoedler, 246 ; distrib., 250,
232.

— spongicola, Stimp., 246; distrib., 229
280, 282.

— stelligera, Zyman, 247; distrib., 280,
232.

Ophiuroidea from West Australia, 246-248.

Ophrydium, Zhrend. ?, 450.

Opisthobranchiata from Abrolhos, 521.

Opisthograptis crateegata, 86.

Orbitolina spherulata, Park. & Jones,
638.

— resicularis, Park. & Jones, 638.

Orbitolites, Lam., 610.

— complanata, Lam., 611, 612.

— — var. plicata, Zam., 611.

— duplex, Carp., 610.

Oyiposition and the egg of Lyctus brunneus,
Steph., 217-227.

Ovipositor of Lyctus, 586-595.

666 INDEX.

Oribata lueasii, Nie., 584.
Oribatula oblonga, 380.

— plantivaga, Berl., 380.

— tibialis, Berl., 580.
Ornithoorina, Orb., 314.
Ornithopora, Ord., 514.
avicularia, Ord., 314.
Ornithoporina, Ord., 514.

— ayicularia, Ord., 314.

— flabellata, Ord., 315.

— flabellata (?), 315.
Orthothecium chryseum, Schimp., 451.
Owl, courtship activities, 289.
Oxytricha pellionella, Kent, 457.
— sp, 453.

Pachychalina sp. (?), 500.

Paleinachus, Woodw., 52.

Paleeogenetic and paleomorphic, use
defined, 99.

Paleozoic Dipnoi, structure of (Watson &
Gill), 163-216.

Palps of Chlorops, 409.

Papilio dardanus, Fabr., mimicry in, 290,

Paraglossee of Chlorops, 408.

Paralelapia nipponica, Hozawa, 488.

Parapostgenal thickening in Chlorops,

406,
Parasterina crassa, Esher, 243; distrib.,
230.

Parawaldechia, Stebb., 3.

—lkidderi, Tatt.*, 2, 3, 18.

— Thomsoni, Stebb., 3.

Parharpinia, Stedb., 4, 5.

— villosa, Stebb., 2, 4, 18.

Parocciput of Chlorops, 405.

Patellina, Williams., 629.

— corrugata, Williams., 629.

Patiria (2) crassa, Gray, 245.

Payonina, Ord., 619.

— flabelliformis, Ord., 619.

Peacock, courtship activities, 277, 280, 287,
290.

Peewit, courtship activities, 277.

Pelican, courtship activities, 269, 282.

Peltigera, Willd., mite on, 379.

Peneroplis, Montf., 610.

— carinatus, Ord., 610.

—- pertusus, Brady, 610.

Penguins, courtship activities, 282, 285.

Pennatulacea from Abrolhos, 21.

Pentagonaster, C. F. Schulize, mentd., 243.

— diibeni, Gray, mentd., 238.

— stibarus, H. Z. Clark, 208; distrib.,
230, 232.

Pentlandia t, mentd., 212-213.

Peregrine, courtship activities, 280.

Periscyphops preeconius, Budde-Lund, 108.

Petalomonas angusta, 457, 465.

Petalostegus, Zev., 315.

— hicornis, Lev., 315.

Petrasia dura, Vosm., 478, 498.

Petrels, courtship activities, 281.

Petricia obesa, H. L, Clark *, 241; distrib.,
230, 232.

— vernicina, mentd., 242,

Phalanterium consociatum, Fres., 463, 465.

— digitatum, Stein, 463, 465.

— solitarium, Sandon *, 468, 464, 475;
mentd., 457.

Phaneropleuron t, mentd., 204, 212-215.

Phaulocylliba virgata, Halb.*, 377-379.

Pheasants, courtship activities, 277, 280,
290.

Phidiana inea, 570.

— lynceus, 570.

Philyra brevis, 40, 54.

— levis, 72, 79.

Philyxia undecimspinosa, 54.

Phloeodictyon abrolhosenis, Dendy*, 478,
500, 519.

— seychellense, Dendy, 501.

— sp., 501.

Phoronis, Wright?, 393, 398.

Phryganella, Penard, 470.

— nidulus, Penard, 470.

Phyletie succession, 83.

Phyllirhoé lichtensteini, Bschsch,, 522.

Phyllomitus sp., 452, 465.

Phyllotegeus palmicinctum, Halb.*, 379.

Phouus bate’, Hasw., 5, 6.

— villosus, Hasw., 4, 5.

Physcia, DC., mite on, 380.

Pigeon, courtship activities, 276, 285.

Placobranchus, Hass., 539, 540.

— expansa, O’ Don. *, 523, 589, 578.

Placopsilina, Ord., 614.

— cenomana, Orb., 614.

Planicellaria, Orb. +, 315.

— eocena, Meun. & Perg. t, 513.

— fenestrata, Orb. t, 315.

INDEX.

Planicellaria oculata, Ord. +, 315.

Planispirina, Seg., 609.

—auriculata, Lyg., 609.

Plankinastrella mammillaris, Lendent., 489.

Plankton, Crustacean, of Hnglish Lakes,
411-447.

Planorbulina, Orb., 635.

— larvata, Brady, 635.

-— mediterranensis, Orb., 635.

— rubra, Orb., 638.

— vulgaris var. larvata, Park. & Jones,
635.

Plecaniwn rugosum, Reuss, 618.

Plectoptera, wing venation in (Tillyard),
143-162,

Pleurobranchus (pars), Bergh, 536.

— ornatus, Cheesm., 538.

Policella australis, Gray, 21, 22.

Polyaspinus cylindricus, Berl., 876, 392.

Polymastia, Bowerb., 509.

—manmumillaris, Bowerb., 479, 509.

Polymorphina, Ord., 625.

— communis, Ord., 625.

— problema, Ord., 625.

— regina, Brady, Park. § Jones, 626.

Polyphemus pediculus, Straus, 425; distrib.,
414, 418, 424.

Polypterus larvee, 85.

Polystomella, Zam., 640.

— crispa, Brady, 640,

— macella, Brady, 640,

— millettiana, Heron-All. § Harl,, 640.

— striato-punctata, Brady, 640.

— verriculata, Millett, 640.

Polytrema, Zzsso, 639.

—miniaceum, Brady, 639.

— — var. alba, Carter, 639.

Polyzoa, Cellularine, 293.

Pontharpinia, Stebd., 4, 5.

— rostrata (Dana), 5, 6.

— uncinata, Chev., 6.

— villosa, Tatt., 6.

Porcellana, Alein, 28, 31, 56.

Porcellanopagurus, Fvdh., 28, 56.

Poricellaria, Ord. +, 515, 522.

— alata, Orb. 7, 315.

Porcellio cristatus, Dollf., 107.

Porocrinus, Bill., 94.

Portunion, Girard, 86, 90.

Postgenze of Chlorops, 405.

667

Potamobius, Leach, 30.

Powder-post beetles, 217-227.

Prionocidaris baculosa var. annulifera,
Mortensen, 248; distrib., 252.

— bispinosa var. chinensis, Déderlein,
248; distrib., 232.

Prodinychus, Berl., subgen., 379.

Prorodon, Ehrenb., mentd., 450.

Protereisma sp. +, wing-veins, 144-161.

Protocarcinus, 52.

Protolelaps, Berl., subgen., 363.

Protozoa, 83; from Spitsbergen, 449.

Psammopemma crassum, Lendenf., 479,511.

Pseudoboletia indiana, A. Agassiz, 249;
distrib., 232.

Pseudoesperia carteri, Dendy*, 479, 500,
503, 519.

— enigmatica, 500.

— trichophora, Dendy *, 479, 509.

Pterodorts, Khrenb., 553.

— picta, Khrenb., 568.

Pteroides griseum, 25.

— hymenocaulon, Broch, 21, 25.

— malayense, 23.

— sp., 22.

Ptilinum of Chlorops, 404.

Ptychodera capensis, Gilehr., 593-398.

— flava, 397.

— minuta, 397.

— natalensis, Gilchi., 393.

— proliferans, Gilchr., 893-398.

Pulvinulina, Park. § Jones, 636.

— canariensis, Brady, 636.

— concentrica, Park. § Jones, 636.

— crassa, Brady, 636.

— elegans, Brady, 637.

— lateralis, Brady, 636.

— repanda, Brady, 636.

— truncatulinoides, Heron-All. § Earl.,
636.

Pygopodes, courtship activities, 266.

Quadrula irregularis, Arch., 453, 457, 471.
Quercus Robur, LZinn., mentd., 219.
Quinqueloculina auberiana, Orh., 607.

— bicostata, Orb., 607.

— bosciana, Orb., 606.

— contorta, Orb., 608.

— curvteriana, Orb., 607.

— dilatata, Orb., 604.

— ferussacit, Orb., 608.

668

Quinqueloculina fusca, Brady, 608.
— kerimbatica, Cushm., 608.

— ornatissima, Karrer, 609.

— schauinsland?, Cushm., 601.

— seminuda, Orb., 605.

— stelligera, Orb., 609.

— striata, Orb., 609.

— triangularis, Orb., 607.

— undosa, Karrer, 607.

— undulata, Orb., 607.

— vulgaris, Orb., 606.

— “with oblique ridges,’ Park., 608.

Rachochoris, Morch, 560.
— laciniata, auct., 560.
Ramulina, R. Jones, 627
— grimaldii, Schlwmob., 697, 646.
— proteiformis, Flint, 627.
— sp.?, Heron-All. § Earl., 627.
Ranilia, MW. Edw., 48, 61, 63.
—muricata, M. Edw., 26, 61.
Ranina, Zam., 25-72
— dentata, Latr., 25, 29, 36, 39, 48, 77.
— marestiana, 52.
Raninella Trigeri, 52.
— elongata, 52.
Raninide, 25-79.
Raninoides, Ldw.,
738, 75.
— personatus, White, 25, 77-79.
— serratifrons, Henders., 74.
taphidiophrys brunii, Penard, 473.
Razorbills, courtship activities, 281.
Recapitulation restated, 81-101.
Redshank, courtship activities, 287.
Redstart, courtship activities, 287.
Remora, Wiil., 90.
Reniera aquieductus, O. Schmidt, 478, 496.
— cribricutis, Dendy, 478, 497.
— dura, Nardo, 498.
— permollis, Tops., 497.
— spp., 478, 498.
teproduction, asexual, in Ptychodera, 393.
Reproductive organs in Coleoptera, 591.
Retepora cornea, Busk, 315.
Retiflustra, Lev., 311, 5165.
— cornea, Harmer *, 315,
— cribriformis, Lev., 315.
— reticulum, 315.
— schonaui, Lev., 515,
Rhabdophaga, Westw., 403.
Rhabdozoum, Hincks, 315.

66, 68, 69, 70, 71,

INDEX.

Rhabdozoum wilsoni, Hincks, 315.

Rhacodes inscriptus, Koch, 105.

Rhaphignathus clavatus, Berl., 585.

— longipilis, Halb.*, 386.

— patrius, Berl., var. brevipalpis,
386, 392.

— — var. truncatus, Halb.*, 585.

— sp., 386.

Robinia Pseudacacia, Linn., mentd., 219.

Rosalina araucana, Orb., 631.

— globularis, Orb., 651.

— lateralis, Terqg., 636.

— mediterranensis, Orb., 631.

— poeyt, Orb., 680.

— rugosa, Orb., 632.

— valvulata, Orb., 632.

— vilardebouna, Orb., 631.

Rostanga pulchra, 566,

Rotalia, Lam., 637.

— beccarii, Orb., 637.

— beccarii (pars), Balk. & Wright, 637.

— elegans, Orb., 637.

—- perlucida, Heron- All. § Eari., 637.

—— precincta, Karrer, 656.

-— spiculotesta, Carter, 617.

Rotalina canariensis, Orb., 636.

—— crassa, Orb., 636.

— ochracea, Williams., 616.

— rosacea, Orb., 631.

— truncatulinoides, Orb., 636.

— wngeriana, Orb., 635.

Rutt, courtship activities, 2

Triig.,

77

Saccohydra, Bill., 315.

— problematica, Bill., 315.
Sagenella frondescens, Brady, 613.
Sagenina, Chapm., 615.

—— frondescens, Chapm., 618.
Sagenodus, Owen, structure,
mentd., 163, 190-215.

— copeanus {, mentd., 181.
— imbricatus t, 178.

— inequalis, Owen +, mentd., 164.
— obliquus +, 178.

Sagrina, Park, § Jones, 626.

— columellaris, Brady, 626.

— raphanus, Brady, 627.

Sainoui on, Sandon *, 459, 475; mentd., 450,
— mikroteron, Sandon *, 459 ; mentd., 450,
460, 475.

— oxu, Sandon *,

468, 475.

163-188 ;

460; mentd., 451, 467,

Salicornaria, Schinz, 315.

Salicornaria, Schweig., 315.

— dichotoma, Busl, 314.

— dichotoma, Schweig., 308, 314, 315.

— farciminoides, Johnst., 303.

Salicornia, Blainv., 515.

Salmacis alevandri, Bell, 249.

— virgulata, 4g. § Des., distrib., 232.

— virgulata alexandri, Doderlein, 249.

Salmo alpinus, 420.

Salpingia, Copp., 316.

— hassallii, Copp., 316.

Sandon, A., Protozoa from Spitsbergen, 449.

Sarciflustra, Jull. § Calv., 311, 516.

— abyssicola, Judl. § Calv., 316.

Scaumenacia*t, mentd., 212-218.

Schnudtia dura, Bals., 498.

Scirus inermis, 77dg., 383, 392.

Sctulus minutus, Halb., 374.

— spathuliger, Veon., 3874.

Seiurus vulgaris Linn., 369.

Scruparia, Oken, 316.

— chelata, Oken, 316, 317.

— clavata, Hincks, 316.

— diaphana, Busk, 314.

— reptans, Verr., 316.

Scrupocellaria (pars), 348.

Scrupocellaria, Van Bened., 304, 316, 329,
357, 358 ; bifurcation, 321, 322.

—- abyssicola, Kluge, 385,

— antarctica, Waters, 352.

— avicularie, Yan. & Ok., 351.

— benemunita, Jull., 334.

— hbertholletii, Aud., 358.

— bifurcata, Kluge, 356.

— caberioides, Kluge, 335.

— drygalskii, Kluge, 352.

— ferox, 326, 359.

— flagellifera, Kluge, 343.

— fuegensis, Waters, 356.

— funiculata, Waters, 335.

— gaussi, Kluge, 388.

— marsupiata, Jull., 351.

— perdita, Kluge, 353.

— simplex, Kluge, 353.

— smittti, Waters, 352,

— solida, Kluge, 338.

— tenuis, Kluge, 352.

— vanhdffeni, Kluge, 353.

— waters, Kluge, 353.

Serupocellaridee, 528, 338.

LINN. JOURN.—ZOOLOGY, VOL. XXXV.

INDEX. 66%

Sceuparia, Gray, 316,

Scupocellaria, Gray, 316.

Scylla pelagica Linn., 522.

Sea-pens from Abrolhos, 21.

Seius becornis, Can. & Fan., 375.

— cornutus, Kram., 366.

— togatus, C. L. Koch, 374.

Seiulus plumosus, Oudem., 369.

Sejus hicornis, Can., 375.

Selbia, Gray, 316.

— zelandica, Gray, 316.

Semiflustra, Orb., 308, 316.

— bombycina, Ord., 308, 316.

Serpula marginata, Walk. § Boys, 624.

— seminulum, Linn., 606.

Sertularia anguina, Linn., 297, 298, 304, 308.

— avicularia, Linn., 299, 800, 314.

— bursaria, Linn., 311, 314.

— chelata, Linn., 307, 316, 317.

— ethata, Linn.. 298, 306, 307.

— cornuta, Linn., 307, 308, 317.

— eburnea, Linn., 306, 307.

— fastigiata, Linn., 306.

— loricata, Linn., 307, 310, 312, 314.

—neritina, Linn., 297, 299.

— repens, Will. & Sol., 316.

— scruposa, Linn., 316.

— spiralis, Olivi, 309.

Sida, Strauss (crustacean), 415.

— crystallina, O. F. Muell., 421, 422,
439; distrib., 414,

— limnetica, Burck., 440.

Sigmosceptrella fibrosa, Dendy, 479, 507.

Siphluride, mentd., 144.

Skua, mentd., 268, 260.

Sparrow-Hawk, courtship activities, 289.

Sparrow, House-, courtship activities, 267.

Spatangus australasie, Leach, 261.

Spathidium, Zooss, 450.

Spherostoma, MacGull., 544, 564.

— dakini, O Don., 523, 548, 545, 577, 578.

— hombergii, 542.

— jamesonii, MacGill., 542.

Sphagnum pools crustacea, 424.

Spiralaria, Busk, 311, 316.

— florea, Busk, 416.

Spiralis, Lev., 316.

Spirastrella vagabunda, Red/., 479, 507.

Spinillina, Ehrend., 628. ‘

—campanula, Heron-Ail. § Earl. *, 628,
629, 646.

48

670

Spirillina decorata, Brady, 628.

— — yar. unilatera, Chapm., 628.

— limbata, Brady, 628.

— lucida, Srdeb., 629.

— revertens, Rhwmbl., 629.

—vivipara, Hhrenb., 628.

Spiroculina, Orbd., 603.

—acutimargo, Brady, 603, 604.

affixa, Terg., 604.

—antillarum, Oré., 603.

— canaliculata, Ord., 603.

— planiculata, Brady, 603.

— tenuirostra, Karrer, 604, 646.

Spiromonas angusta, Kent, 452, 463.

—angustator, Duj., 457.

Spirophora bacterium, Lendenf., 507.

— digitata, Lendenf., 507.

Spiroplecta, Hhrenb., 620.

— biformis, Brady, 620.

Spitsbergen Protozoa, 449.

Splanchnotrophus, Hane. § Norm., 570.

— brevipes, 569, 570.

— gracilis, Hane. § Norm., 569, 571, 573,
579.

— sacculatus, O’ Don., 571, 573.

— sp., 570.

Spongelia dakini, Dendy *, 479, 511.

— incerta, Hyatt, 513.

— spinosa, Hyutt, 513.

Sponges from Abrolhos, 477.

Spongia bilamellata, Lam., 504.

— palmata, Lam., 499.

Steganoporella magnilabris, 304,

Stellaster incei, Gray, 238; distrib., 232.

Stelletta australiensis, Carter, 492.

— brevis, Hentsch., 478, 490.

— debilis, Thiele, 478, 491.

— sigmatrizna, Lendenf., 478, 491.

Stephanocrinus, Conr., 94.

Stigmzeus anthrodes var. reticulatus,
Halb. *, 387, 392.

Stirparia, Goldst., 295, 317.

— annulati, Goldst., 317.

Stirparia, Zewck., mentd., 317.

Stirpariella, Harmer, nom. nov. *, 295,301,
317, 325-327, 359.

—annulata, Harmer *, 317.

— carabaica, Lev., 326.

— zanzibariensis, Waters, 325.

Stolonella, Zincks, 317.

— clausa, Hincks, 317.

INDEX.

Strophidium, term defined, 33.

Stylonychia sp., 453.

Suberites aaptus, Lendenf., 508,

— spissus, Tops., 508.

Swans, courtship activities, 278, 281,
289,

Swartzia montana, Lindb,, 450.

Sycon gelatinosum, Blainv., 477, 488.

Synnota, Piep., 317.

Synnotum, inchs, 310, 313, 317.

— egyptiacum, Harmer *, 317.

— aviculare, Hincks, 317.

Synura uvella, Lhrend., 450.

Tabanus, Linn., 403, 409.

Tarns, crustacea, 422.

Tarsonemus brevipes, 382.

— culmicolus, Rewt., 381.

— floricolus, 382.

laticeps, Halb.*, 381, 392.

— spirifex, March., 382.

Tattersall, W., Abrolhos Amphipoda and
Tsopoda, 1-19.

Tauroglossus, Speng., 393.

Tentorium of Chiorops, 399-403.

Terncellaria, Orb., 317, 353.

— aculeata, Orb., 317,

Terrestrial [sopods, 103, 107.

Tethea robusta, Bowerb., 495.

Tethya cliftont, Ridl., 495.

— globostellata, Lendent., 495.

— ingalli, Wirkp., 495.

— lyncurium, Row, 495.

— multastella, Lendenf., 495.

Tethyophena silivcifica, O. Schmidt, 508.

Tethys, Linn., 527, 528.

— denisoni, 523, 531, 577, 578,

— gigantea, 522, 523, 528, 529, 531, 5388,
577, 578.

— punciata, 529, 531, 532.

Textularia, Defi., 617.

— abbreviata, Ord., 618.

—agelutinans, Ord., 618.

— — var. abbreviata, Park. & Jones, 615,
618, 619.

— — var. biformis, Park. & Jones, 620.

— candeiana, Orb., 618.

— conica, Orb., 619.

— — var. jugosa, Jones § Millett, 619.

— crispata, Brady, 617.

INDEX. 671

Textularia folium, Park. § Jones, 617.

— gramen, Ordb., 615, 618.

— inconspicua var. jugosa, Millett, 617.

— rhomboidalis, Millett, 617.

— rugosa, Brady, 618.

— sagittula, Defr., 618.

— — var. fistulosa, Brady, 618.

— variabils, Williams., 622.

Thalassodendron typica, Whitel., 504.

Theca of Chlorops, 408.

Theria, Hiib., 88.

— rupicapraria, 88.

Thia polita, 57.

Thiolliericrinus, Htallon, 95.

Tillyard, R. J., Wing-venation of the order
Plectoptera or Mayflies, 145-162.

Tinoporus bacuatus, Carp., 638.

Tipula, Linn., 409.

Tormee of Chlorops, 403.

Toxema tubulota, Hallm., 503.

Toxopneustes indianus, Michelin, 249.

Trachya durissima, Carter, 509.

— globosa, Carter, 509.

— — yar. rugosa, Carter, 509.

— horrida, Carter, 509.

— pernucleata, Carter, 508, 509.

Trachycladus levispirulifer, Carter, 479,
506.

Trachynotus egrota var. pyriformis, Berl.,
376.

— egrota, C. L. Koch, 376.

— lambda, C. L. Koch, 376.

— minima, 77dig., 377.

— pi, Berl., var. pauperior, Berl., 377.

— pyriformis, Kram., 376.

— tubilera, C. L. Koch, 376.

Trachytes pyriformis, Halb.*, 376.

Trachyuropoda cxspitum, 379.

— celtica, Halb., 378.

— cordieri, Berl., 378.

— troguloides, Can. § Fan., var. celtica,
Halb.*, 378, 392.

Trevelyana, Kel., 547.

. Tricellaria, Flem., 800, 804, 317, 324, 327,

329, 348, 357, 358.

aculeata, Harmer *, 355, 356, 359.

— aculeata, Orb., 317, 326.

-— longispinosa, Harmer *, 357.

— monotrypa, Harmer *, 355,

— occidentalis, Harmer *, 354,

— — var. dilatata, Ovtm., 327, 359.

Tricellaria peachii, Harmer *, 325-327,
329, 355, 359.

— pribilofi, Harmer *, 354.

— sympodia, Harmer *, 355.

— ternata, Harmer, 317, 324, 354, 359.

— — yar. gracilis Harmer”, B54.

Triloculina circularis, Bornem., 604.

— labiosa, Orb., 604.

— levigata, Orb., 606.

— linneiana, Orb., 609.

— reticulatu, Orb., 608.

— rotundata, Orb., 606.

— tricarinata, Orb., 605.

— trigonula, Orb., 605.

— valvularis, Reuss, 604.

— webbiana, Orb., 605.

Trinema complanatum, Penard, 453, 454,
473.

— enchelys, Lhrenb., 453, 457, 472.

— — var. galeata, Penurd, 453, 473.

— lineare, Penard, 453, 457, 472.

Triopa clavigera, Johnst., 564.

Tripneustes gratilla, Lovén, 249; distrib.,
232.

Tritaxia, Reuss, 619.

— lepida, Brady, 619.

Tritonia, Cuv., 542, 564.

— clavigera, Rafin., 564.

— hombergit, Cuy., 542, 564.

Tritonium, O. F. Muell., 564.

Trochammina, Park. § Jones, 616.

— inflata var., Ballw. & Wright, 616.

— ochracea, Millett, 616.

— rotaliformis, Heron-All. § Earl., 616.

— squamata, Jones § Park., 616.

— vesicularis, G'oés, 616.

Truncatulina, Ord., 635.

— lobulata, Brady, 635.

— precincta, Brady, 636.

— refulgens, Brady, 635.

— ungeriana, Brady, 635.

— variabilis, Orb., 635.

Tuberellu, Kell., 509,

— aaptos, Lendenf., 508.

— Duchassainyr, Tops., 508.

— tethyoides, Kell., 508, 509.

Tubucellaria, Orbd., 315.

Tubularia fistulosa, Linn., 301, 302.

— muscoides, Lian., 309.

— ramosa, Linn., 309.

— salicornis, Ksper, 303.

672 INDEX.

Turnstone, courtship activities, 287.
Tylos latreillei, Aud. & Say., 103.

Ulidium, 8. Wood, 318, 317.

— charlesworthit, S. Wood, 313, 317.

Umbrella-bird, courtship activities, 277.

Unicellaria, Blainy., 317.

— appendiculata, Blainy., 317.

— chelata, Blainv., 317.

— cornuta, Blainv., 317.

— lafoy?, Blainv., 317.

Uniophora dyscrita, H. L. Clark*, 244;
distrib., 230.

Uroglena, Hhrenb., 450.

Uroleptus mobilis, Engelm., 457.

Uronema sp., 457, 468.

— schewiakofli, Buddenb., 468.

Uronemus +, mentd., 163, 215.

—splendens (Ziag.) +, structure, 201-
204, 213-215.

Uropoda obscura, C. L. Koch, 377.

Uroseius acuminatus, C. L. Koch, 377.

Uvigerina, Orb., 626.

— angulosa, Wilhiams., 626.

— porrecta, Brady, 626.

— pygmea, Orb., 626.

— raphanus, Park. & Jones, 627.

— selseyensis, Heron-All. § Earl., 626,
646.

Veretillum australe, Awk. § Broch, 22.
— malayense, Hickson, 21,
Vernuculum leve, Mont., 624.
— oblongum, Mont., 606.

— squamosum, Mont., 624.
— subrotundum, Mont., 605.
Verneuilina, Ord., 619,

—— polystropha, Brady, 616.
— propinqua, Brady, 619.
— spinulosa, Brady, 619.
Vertex of Chlorops, 405.

Vespertilio dasycneme, 369.

Vincularia fragilis, Defr. +, 311.

Virgularia elegans, Gray, 21.

Volvox, Zinn., in Highlow Tarn, 424.

Vorticella, O. F. Mueill., mentd., 450.

— microstoma, 457.

Vorticella, Esper, 305.

— polypina, Esper, 305.

— polypina, L., 305.

Vosmaeropsis mackinnoni, Dendy*, 477,,
483, 518.

Waldeckia, Chevr., 3.

Walnut, “ Italian,’ mentd., 219.

Warblers, courtship activities, 275, 277,.
281.

Wastwater plankton, 421.

Watersia, Lev., 317.

— wmilitaris, Lev., 317.

Watson, D.M.S., aud E. L. Gill, Structure
of certain Paleeozoic Dipnoi, 163—
216.

Xylotrogus brunneus, Steph., 217.
Yvesia spinulata, Hentsch., 505.

Zanclifer, 58, 66, 67.

— caribensis, Frremen., 26, 60,
Zercon bicornis, Berl., 375.

— ornatus, Berl., 375.

— perforatulus, Berl., 375.
—radiatus, Berl., 375.

— tragardhii, Halb.*, 375, 592.
— triangularis, C. L. Koch, 373.
Zerconopsis, Hull, 374.

— minutus, Hull, 374.
Zercoseius, Berl., subgen., 374.
Zygoptera, mentd., 143,
Zygotes, phyletic succession, 83.

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